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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.1180.107333" ID-Pensoft-Pub="1313-2970-1180-257" ID-Pensoft-UUID="D3C115F72BB351AB89F7AF46AF0A7E43" ID-ZooBank="912793FF3FCF46248CCBB42A9FD2BD73" ModsDocID="1313-2970-1180-257" checkinTime="1695824266241" checkinUser="pensoft" docAuthor="Reyes-Puig, Juan Pablo, Urgiles-Merchan, Miguel, Franco-Mena, Daniela, Guayasamin, Juan M., Batallas, Diego &amp; Reyes-Puig, Carolina" docDate="2023" docId="10218311988457C8AE068B321C63C959" docLanguage="en" docName="ZooKeys 1180: 257-293" docOrigin="ZooKeys 1180" docPubDate="2023-09-26" docSource="http://dx.doi.org/10.3897/zookeys.1180.107333" docTitle="Pristimantis kayi Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig 2023, sp. nov." docType="treatment" docUuid="E4854476-B0BD-44DB-8E5F-3D4E2A590962" docUuidSource="ZooBank" docVersion="2" id="D3C115F72BB351AB89F7AF46AF0A7E43" lastPageNumber="257" masterDocId="D3C115F72BB351AB89F7AF46AF0A7E43" masterDocTitle="Two new species of terrestrial frogs of the Pristimantis gladiator complex (Anura, Strabomantidae) from the Ecuadorian Andes, with insights on their biogeography and skull morphology" masterLastPageNumber="293" masterPageNumber="257" pageNumber="257" updateTime="1695824568533" updateUser="ExternalLinkService">
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<mods:title id="F6370D3AA93D617D8FDBF36F9ADCCA6A">Two new species of terrestrial frogs of the Pristimantis gladiator complex (Anura, Strabomantidae) from the Ecuadorian Andes, with insights on their biogeography and skull morphology</mods:title>
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<mods:namePart id="18AB7C1F8F376780F7CC3E32383E2809">Reyes-Puig, Juan Pablo</mods:namePart>
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<mods:affiliation id="D5A2D40AB32F08BE50ABA7BCCBEADFED">Unidad de Investigacion, Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador &amp; Fundacion EcoMinga; Fundacion Oscar Efren Reyes, Departamento de Ambiente, Banos, Ecuador</mods:affiliation>
<mods:nameIdentifier id="C662BBEF608B191E74EEED25D9E94C3C" type="email">juanpabloecominga@gmail.com</mods:nameIdentifier>
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<mods:namePart id="D022F92CA5335F99CEE2D0DC236B3379">Urgiles-Merchan, Miguel</mods:namePart>
<mods:nameIdentifier id="82DF84CE56A556C4F07A74E2F746410B" type="ORCID">https://orcid.org/0000-0002-4606-2237</mods:nameIdentifier>
<mods:affiliation id="11B6DF0FD13D0152AF3132B1AAC45255">Unidad de Investigacion, Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador</mods:affiliation>
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<mods:namePart id="A439A92EBCFBB14DA92C3455D20F21EF">Franco-Mena, Daniela</mods:namePart>
<mods:nameIdentifier id="95D8AE5028E57C39228758DBE5D28A93" type="ORCID">https://orcid.org/0000-0003-3655-2678</mods:nameIdentifier>
<mods:affiliation id="BDE2E9C0502247D8638476E951B2B8AF">Laboratorio de Biologia Evolutiva, Colegio de Ciencias Biologicas y Ambientales COCIBA, Universidad San Francisco de Quito USFQ, Quito, Cumbaya, Ecuador</mods:affiliation>
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<mods:namePart id="53D0F62CF2EE31FFF8FA632F7D1DAF96">Guayasamin, Juan M.</mods:namePart>
<mods:nameIdentifier id="63AC36BE42145799A79B852692363052" type="ORCID">https://orcid.org/0000-0003-0098-978X</mods:nameIdentifier>
<mods:affiliation id="8AB079B1DE60A9F4F664BBBA850F85D5">Laboratorio de Biologia Evolutiva, Colegio de Ciencias Biologicas y Ambientales COCIBA, Universidad San Francisco de Quito USFQ, Quito, Cumbaya, Ecuador</mods:affiliation>
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<mods:namePart id="60A6C7A4E2EC5BE6F3774CDCF3C7E910">Batallas, Diego</mods:namePart>
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<mods:affiliation id="8900D97449EBAD026349D44D46B26687">Laboratorio de Biologia Evolutiva, Colegio de Ciencias Biologicas y Ambientales COCIBA, Universidad San Francisco de Quito USFQ, Quito, Cumbaya, Ecuador &amp; Universidad Complutense de Madrid, Madrid, Spain</mods:affiliation>
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<mods:namePart id="8CA04570B95499A57EE4615ECC3E99F0">Reyes-Puig, Carolina</mods:namePart>
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<mods:affiliation id="803422D034D1E03E3A0324FF15561F05">Unidad de Investigacion, Instituto Nacional de Biodiversidad (INABIO), Quito, Ecuador &amp; Laboratorio de Biologia Evolutiva, Colegio de Ciencias Biologicas y Ambientales COCIBA, Universidad San Francisco de Quito USFQ, Quito, Cumbaya, Ecuador</mods:affiliation>
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<subSection id="9B5F997049C5BAFE7A292F9F62099FC3" pageId="0" pageNumber="257" type="results">
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<paragraph id="477579DE1F0D2D8863F0BE3CCC69CEF2" pageId="0" pageNumber="257">
<taxonomicName id="3BE7AB4AD03368DB889CE3926476851D" LSID="https://zoobank.org/E4854476-B0BD-44DB-8E5F-3D4E2A590962" authority="Juan M. Guayasamin, Miguel Urgilés-Merchán, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi" status="sp. nov.">
Pristimantis kayi Juan M. Guayasamin, Miguel
<normalizedToken id="0B8BC7A5B7160ED854E3208C1DA9B355" originalValue="Urgilés-Merchán">Urgiles-Merchan</normalizedToken>
, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig
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<taxonomicNameLabel id="89B504CBBB5BECAEA4D91D73C12D3B5F" pageId="0" pageNumber="257">sp. nov.</taxonomicNameLabel>
</paragraph>
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<subSubSection id="762968E9B1007CFC1DD917EE86B43B41" pageId="0" pageNumber="257" type="description">
<paragraph id="018063B95387665F73AAF513FAB8460C" pageId="0" pageNumber="257">
<figureCitation id="428BF098AC3D04D801E797BBD17A43B1" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Evolutionary relationships amongst species in the Pristimantis gladiator complex, under the Maximum Likelihood inference criterion (modified from Franco-Mena et al. (2023)). Red font indicates the new sequence." figureDoi="10.3897/zookeys.1180.107333.figure1" httpUri="https://binary.pensoft.net/fig/912029" pageId="0" pageNumber="257">Figs 1</figureCitation>
<figureCitation id="4494BF2F80DC5994708665430DC3B7EF" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Dorsolateral and ventral views in life of Pristimantis gladiator species complex and related species A Pristimantis donnelsoni sp. nov. female, DHMECN 14701, holotype, LRC: 15.42 mm B Pristimantis donnelsoni sp. nov. male, DHMECN 18854, LRC: 13.66 mm from Cerro Candelaria C Pristimantis kayi sp. nov. female, ZSFQ 0775, holotype, LRC: 19.7 mm D Pristimantis kayi sp. nov. male, DHMECN 14450 LRC: 13.58 mm, from Reserva Machay E Pristimantis gladiator sensu stricto female, DHMECN 12463, LRC: 20.38 mm F Pristimantis gladiator sensu stricto, male DHMECN 12474, LRC: 16.87 mm; from Rio Quijos drainage G Pristimantis festae female, DHMECN 16280, LRC: 20.19 mm H Pristimantis festae, male (left), DHMECN 16837, LRC: 16.50 mm; female (right), DHMECN 16836, LRC: 19.80 mm; Photographers: JPRP (A, B), MYM (D, E, F, H); Patricia Bejarano (G); Jose Vieira (C)." figureDoi="10.3897/zookeys.1180.107333.figure5" httpUri="https://binary.pensoft.net/fig/912033" pageId="0" pageNumber="257">, 5</figureCitation>
<figureCitation id="5266DC54C5F90EC4AC1D974B09C4E641" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Skull morphology diagram in dorsal (left column) and ventral (right column) views in four species of the Pristimantis myersi species group: A, B P. donnelsoni sp. nov., DHMECN 4808 C, D P. kayi sp. nov., DHMECN 14447 E, F P. gladiator DHMECN 12465 G, H P. festae DHMECN 9294. Detail of the skull bones abbreviations: pmax = premaxilla; max = maxilla; spheth = sphenethmoides; fpar = frontoparietal; pter = pterigoides; pro = prootic; q = quadratojugal; sp = squamosal; exo = exoccipital; prph = parasphenoides. Frontoparietal fontanelle not coloured." figureDoi="10.3897/zookeys.1180.107333.figure8" httpUri="https://binary.pensoft.net/fig/912036" pageId="0" pageNumber="257">, 8</figureCitation>
<figureCitation id="9779B83B431D2F545101CC2577849795" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Habitat vegetation at type localities and etymology of the two new frogs described herein. Photographs by JPRP and Zane Libke A panoramic view of the upper Pastaza Valley and its adjacent mountains, main geographic barriers promoting speciation at a regional scale, at back Tungurahua Volcano and mountains south of the Pastaza River habitat of Pristimantis donnelsoni sp. nov., image from the slopes of Cerro Mayordomo type locality of Pristimantis kayi sp. nov., in the northern mountains to the Pastaza River B Tungurahua Volcano eruption, from Chamana Reserve C Montane forest in Candelaria Reserve D Don Nelson Palacios (+) during an expedition; we tribute his work with the naming of Pristimantis donnelsoni sp. nov. E Sumaco Volcano seen from Wawa Sumaku, habitat of P. kayi sp. nov. F Montane forest and subparamo in Leito, Patate G Andreas Kay (+); we honour his devotion to nature photography and research in the upper Pastaza Valley and through Ecuador by naming Pristimantis kayi sp. nov." figureDoi="10.3897/zookeys.1180.107333.figure12" httpUri="https://binary.pensoft.net/fig/912040" pageId="0" pageNumber="257">, 12</figureCitation>
<figureCitation id="C451A21116CDCCA999279EB48DF9C888" captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Holotype of Pristimantis kayi sp. nov. in life, adult female, ZSFQ 0775. LRC: 19.7 mm. Photographs by Jose Vieira." figureDoi="10.3897/zookeys.1180.107333.figure13" httpUri="https://binary.pensoft.net/fig/912041" pageId="0" pageNumber="257">, 13</figureCitation>
<figureCitation id="8ADCA09146B2BBE88BA27DCBFBC80BA3" captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Colour variation of Pristimantis kayi sp. nov. A MZUTI 2209, adult female, LRC: 20.15 mm, from Cordillera de Guacamayos B MZUTI 2210, adult female, LRC: 19.90 mm, from Cordillera de Guacamayos C MZUTI 2211, adult male, LRC: 14.30 mm, from Cordillera de Guacamayos D MZUTI 2213, adult male, LRC: 15.10 mm, from Cordillera de Guacamayos E MZUTI 2214, adult female, LRC: 18.73 mm, from Cordillera de Guacamayos F ZSFQ 774, adult female, LRC: 18.96 mm from Sumaco Volcano G ZSFQ 782, adult female, LRC: 30.03 mm, from Sumaco Volcano. Photographs by Lucas Bustamante (A, B, C, D, E) and Jose Vieira (F, G)." figureDoi="10.3897/zookeys.1180.107333.figure14" httpUri="https://binary.pensoft.net/fig/912042" pageId="0" pageNumber="257">, 14</figureCitation>
<figureCitation id="E5BB541B6FF902BD073377F7884C91B1" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Preserved holotype of Pristimantis kayi sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257">, 15</figureCitation>
<figureCitation id="7CE168C9DC95863B6F7E20BCF1BFBA7E" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">, 16</figureCitation>
<figureCitation id="9632A658587787AC64B96FC3D1AA5D68" captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Dorsal and ventral views of preserved type series of Pristimantis kayi sp. nov. A holotype ZSFQ 0775, female; Paratypes B ZSFQ 0779, female C DHMECN 16212, female D MZUTI 5162, female E MZUTI 4657, male F ZSFQ 0773, male G DHMECN 15233, male H DHMECN 18440, male; Photographs by JPRPR and David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure17" httpUri="https://binary.pensoft.net/fig/912045" pageId="0" pageNumber="257">, 17</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="6529DAE8BFF45C7455EB5DA192FEAE80" pageId="0" pageNumber="257" type="type material">
<paragraph id="389EB4E066DC9757EDFD0D18CF82C241" pageId="0" pageNumber="257">Type material.</paragraph>
<paragraph id="FA321F7826078AC4544A8308BC5E3701" pageId="0" pageNumber="257">
<emphasis id="AF07DCC8135CA4F7F95D4A6EA80378AD" bold="true" pageId="0" pageNumber="257">
<emphasis id="2DBAE745A686A8113114E523EDC16879" italics="true" pageId="0" pageNumber="257">Holotype</emphasis>
.
</emphasis>
ZSFQ 0775, adult female (Figs
<figureCitation id="09D69C1D52F229852B5D95432DAEEF87" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Dorsolateral and ventral views in life of Pristimantis gladiator species complex and related species A Pristimantis donnelsoni sp. nov. female, DHMECN 14701, holotype, LRC: 15.42 mm B Pristimantis donnelsoni sp. nov. male, DHMECN 18854, LRC: 13.66 mm from Cerro Candelaria C Pristimantis kayi sp. nov. female, ZSFQ 0775, holotype, LRC: 19.7 mm D Pristimantis kayi sp. nov. male, DHMECN 14450 LRC: 13.58 mm, from Reserva Machay E Pristimantis gladiator sensu stricto female, DHMECN 12463, LRC: 20.38 mm F Pristimantis gladiator sensu stricto, male DHMECN 12474, LRC: 16.87 mm; from Rio Quijos drainage G Pristimantis festae female, DHMECN 16280, LRC: 20.19 mm H Pristimantis festae, male (left), DHMECN 16837, LRC: 16.50 mm; female (right), DHMECN 16836, LRC: 19.80 mm; Photographers: JPRP (A, B), MYM (D, E, F, H); Patricia Bejarano (G); Jose Vieira (C)." figureDoi="10.3897/zookeys.1180.107333.figure5" httpUri="https://binary.pensoft.net/fig/912033" pageId="0" pageNumber="257">5C</figureCitation>
,
<figureCitation id="C926CED76DBF1567F52BDE6B09C428BA" captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Holotype of Pristimantis kayi sp. nov. in life, adult female, ZSFQ 0775. LRC: 19.7 mm. Photographs by Jose Vieira." figureDoi="10.3897/zookeys.1180.107333.figure13" httpUri="https://binary.pensoft.net/fig/912041" pageId="0" pageNumber="257">13</figureCitation>
,
<figureCitation id="D3A2AE1C8CAE2B830B7A192623585B35" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Preserved holotype of Pristimantis kayi sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257">15</figureCitation>
-
<figureCitation id="32ECD0A269588D4CB26A964E9648F436" captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Dorsal and ventral views of preserved type series of Pristimantis kayi sp. nov. A holotype ZSFQ 0775, female; Paratypes B ZSFQ 0779, female C DHMECN 16212, female D MZUTI 5162, female E MZUTI 4657, male F ZSFQ 0773, male G DHMECN 15233, male H DHMECN 18440, male; Photographs by JPRPR and David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure17" httpUri="https://binary.pensoft.net/fig/912045" pageId="0" pageNumber="257">17</figureCitation>
) collected by
<normalizedToken id="145A29541BFE31CCB7264B23B4897B7B" originalValue="José">Jose</normalizedToken>
Vieira, David Brito-Zapata, Jefferson Mora, and Carolina Meneses from
<normalizedToken id="0BE301807D0548FCB8F7F51189A5D70F" originalValue="Volcán">Volcan</normalizedToken>
Sumaco, Sumaco National Park, Napo, Ecuador (
<geoCoordinate id="063F1D9D2594BD1C74150C1D557BF0A2" degrees="0.5696" direction="south" orientation="latitude" precision="5" value="-0.5696">0.5696°S</geoCoordinate>
,
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; 2322 m alt.) on 11 September 2018.
</paragraph>
<caption id="D7BE1F7AC3F6ED458D316AF2663D1B18" doi="10.3897/zookeys.1180.107333.figure13" httpUri="https://binary.pensoft.net/fig/912041" pageId="0" pageNumber="257" start="Figure 13" startId="F13">
<paragraph id="A606DAE160B25FDA5883F2B87EE8C95E" pageId="0" pageNumber="257">
<emphasis id="6466DE2F29E938BBB11B59A59A4DE712" bold="true" pageId="0" pageNumber="257">Figure 13.</emphasis>
Holotype of
<taxonomicName id="AD68E9CC6C858BECDC26279D736C4B2B" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="B76853CC6BA5EA1966C62DDD7817B3FE" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. in life, adult female, ZSFQ 0775. LRC: 19.7 mm. Photographs by Jose Vieira.
</paragraph>
</caption>
<paragraph id="6BAC88B3F4D05FDF90ACF5C11B957C76" pageId="0" pageNumber="257">
<emphasis id="7AD4E1D04A8F9F5E4D245D11EACA7868" bold="true" italics="true" pageId="0" pageNumber="257">Paratypes</emphasis>
(13, 4 ♂ / 9 ♀, Figs
<figureCitation id="478AA859770A27EA99BA020EC1F62FDA" captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Colour variation of Pristimantis kayi sp. nov. A MZUTI 2209, adult female, LRC: 20.15 mm, from Cordillera de Guacamayos B MZUTI 2210, adult female, LRC: 19.90 mm, from Cordillera de Guacamayos C MZUTI 2211, adult male, LRC: 14.30 mm, from Cordillera de Guacamayos D MZUTI 2213, adult male, LRC: 15.10 mm, from Cordillera de Guacamayos E MZUTI 2214, adult female, LRC: 18.73 mm, from Cordillera de Guacamayos F ZSFQ 774, adult female, LRC: 18.96 mm from Sumaco Volcano G ZSFQ 782, adult female, LRC: 30.03 mm, from Sumaco Volcano. Photographs by Lucas Bustamante (A, B, C, D, E) and Jose Vieira (F, G)." figureDoi="10.3897/zookeys.1180.107333.figure14" httpUri="https://binary.pensoft.net/fig/912042" pageId="0" pageNumber="257">14</figureCitation>
,
<figureCitation id="73F907119ECFBC2E890626EEDB4FEDA7" captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Dorsal and ventral views of preserved type series of Pristimantis kayi sp. nov. A holotype ZSFQ 0775, female; Paratypes B ZSFQ 0779, female C DHMECN 16212, female D MZUTI 5162, female E MZUTI 4657, male F ZSFQ 0773, male G DHMECN 15233, male H DHMECN 18440, male; Photographs by JPRPR and David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure17" httpUri="https://binary.pensoft.net/fig/912045" pageId="0" pageNumber="257">17</figureCitation>
). ZSFQ 0779, adult female. ZSFQ 773, adult male. ZSFQ 782, adult female with the same data as the holotype, ZSFQ 783, adult female collected by
<normalizedToken id="FDBE0FED1453B4719C92C4E9061612D0" originalValue="José">Jose</normalizedToken>
Vieira, David Brito-Zapata, Jefferson Mora, and Carolina Meneses from Sumaco National Park, Napo, Ecuador (
<geoCoordinate id="B73EC58825867427708B4A207E159C3F" degrees="0.5696" direction="south" orientation="latitude" precision="5" value="-0.5696">0.5696°S</geoCoordinate>
,
<geoCoordinate id="141BBEE5C1E912B3CB730017514235A1" degrees="77.5753" direction="west" orientation="longitude" precision="5" value="-77.5753">77.5753°W</geoCoordinate>
; 2,476 m alt.) on 1 September 2018; ZSFQ 778, adult male, ZSFQ 780 and 781 with same data as ZSFQ 783. DHMECN 14446, adult female collected by Mario Humberto
<normalizedToken id="A671D469E77DEB81256A6FE38AF0D1D9" originalValue="Yánez">Yanez</normalizedToken>
<normalizedToken id="E54758104F26517471D0C4801304DC63" originalValue="Muñoz">Munoz</normalizedToken>
, JPRP, and DFM from Cerro Mayordomo, Reserva Machay,
<normalizedToken id="ACA7D4D0B5064C98799877983966E2D8" originalValue="Río">Rio</normalizedToken>
Negro,
<normalizedToken id="13E40851D1A41CEAD1CD0789001829FE" originalValue="Baños">Banos</normalizedToken>
de Agua Santa, Tungurahua, Ecuador (
<geoCoordinate id="5DD71622DE44C4312E9205CA9C89574E" degrees="1.3686" direction="south" orientation="latitude" precision="5" value="-1.3686">1.3686°S</geoCoordinate>
,
<geoCoordinate id="7370D7C740FFD69B4D2F4505DB07F550" degrees="78.2693" direction="west" orientation="longitude" precision="5" value="-78.2693">78.2693°W</geoCoordinate>
8; 2972 m alt.) on 2 March 2018. DHMECN 15226 and 15237, adult females collected by Kelsey Huisman and Eduardo
<normalizedToken id="D9BFD67AD941B89D85B61459FC87322A" originalValue="Peña">Pena</normalizedToken>
from Vizcaya, Reserva Naturetrek, Ulba,
<normalizedToken id="9B065F25B5BCA629CDD2BF0DE1B4F0BC" originalValue="Baños">Banos</normalizedToken>
de Agua Santa, Tungurahua, Ecuador (
<geoCoordinate id="968EFDCD05A838C29EF836054C010CBA" degrees="1.396" direction="south" orientation="latitude" precision="55" value="-1.396">1.396°S</geoCoordinate>
,
<geoCoordinate id="66E87655258B60C9A73B55320F3BF37C" degrees="78.3942" direction="west" orientation="longitude" precision="5" value="-78.3942">78.3942°W</geoCoordinate>
; 3150 m alt.) on 12 December 2019. DHMECN 15231 and 15233, adult females collected by Kelsey Huisman and Eduardo
<normalizedToken id="3CFCD1C9A16764B42D98FD89BA28205B" originalValue="Peña">Pena</normalizedToken>
from Cerro Mayordomo, Reserva
<normalizedToken id="8F8B1926351774EA090353B7862181AB" originalValue="Ecológica">Ecologica</normalizedToken>
Machay,
<normalizedToken id="F76AF03C41F17DB413460DD25C8BA376" originalValue="Río">Rio</normalizedToken>
Verde,
<normalizedToken id="04A111119DB059411039EB019FB7EE84" originalValue="Baños">Banos</normalizedToken>
de Agua Santa, Tungurahua, Ecuador (
<geoCoordinate id="0517CA05CC5B1D4C6AE31BF63794C33E" degrees="1.368" direction="south" orientation="latitude" precision="55" value="-1.368">1.368°S</geoCoordinate>
,
<geoCoordinate id="EF94C3D797E75120E116F11DA102C2AA" degrees="78.2692" direction="west" orientation="longitude" precision="5" value="-78.2692">78.2692°W</geoCoordinate>
; 2969 m alt.) on 24 November 2019. DHMECN 16212, adult female collected by JPRP fromVizcaya, Reserva Naturetrek, Ulba,
<normalizedToken id="362FF5205E68106D424B22B250E03E9F" originalValue="Baños">Banos</normalizedToken>
de Agua Santa, Tungurahua, Ecuador (
<geoCoordinate id="10D12B5C0021FAC51272E8731967E5F6" degrees="1.3962" direction="south" orientation="latitude" precision="5" value="-1.3962">1.3962°S</geoCoordinate>
,
<geoCoordinate id="A1A0F9C1CAF931D44B56B47835C11C87" degrees="78.3941" direction="west" orientation="longitude" precision="5" value="-78.3941">78.3941°W</geoCoordinate>
; 3062 m alt.) on May 2021. DHMECN 18440, adult male collected by JPRP, Eduardo
<normalizedToken id="CFB30E69E112EB6B8713D2570FF14771" originalValue="Peña">Pena</normalizedToken>
, and Edgar
<normalizedToken id="925477785C50C83D94CC2C73DF23614A" originalValue="Martínez">Martinez</normalizedToken>
, from Los
<normalizedToken id="5E58D7E2AF3DD86E034F3527545A1EEE" originalValue="Mortiños">Mortinos</normalizedToken>
, Leito, Patate, Tungurahua, Ecuador (-1.315235, -78.452793; 3448 m alt.) on 6 July 2022. MZUTI 2209, adult female collected by Juan M. Guayasamin and Lucas Bustamante from Cordillera de los Guacamayos, Napo, Ecuador (
<geoCoordinate id="F74C712A9DC9F61B3DA8E068FFB32710" degrees="0.3769" direction="south" orientation="latitude" precision="5" value="-0.3769">0.3769°S</geoCoordinate>
,
<geoCoordinate id="11382516745CC52A2F1143EA81761E80" degrees="77.5048" direction="west" orientation="longitude" precision="5" value="-77.5048">77.5048°W</geoCoordinate>
; 2190-2247 m alt.) in March 2013. MZUTI 2010, 2011, 2013 and 2014 with the same data as MZUTI 2209.
</paragraph>
<caption id="E775F803D6CDA95BDE200781E45EF1C8" doi="10.3897/zookeys.1180.107333.figure14" httpUri="https://binary.pensoft.net/fig/912042" pageId="0" pageNumber="257" start="Figure 14" startId="F14">
<paragraph id="E2EF4D55052D339B199662D34983AD8B" pageId="0" pageNumber="257">
<emphasis id="AC00BC7361C54F97880E913F41C1750C" bold="true" pageId="0" pageNumber="257">Figure 14.</emphasis>
Colour variation of
<taxonomicName id="E40BAD0219B16D1E71BF0351CB802EC7" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="944DB8EB3111C4E119DCEC5C31B447AB" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov.
<emphasis id="E9E7B8B1E3525D3E9F507A6CC0105FFA" bold="true" pageId="0" pageNumber="257">A</emphasis>
MZUTI 2209, adult female, LRC: 20.15 mm, from Cordillera de Guacamayos
<emphasis id="4BB7B27ABCEDD9043A33AB669E33BC6D" bold="true" pageId="0" pageNumber="257">B</emphasis>
MZUTI 2210, adult female, LRC:19.90 mm, from Cordillera de Guacamayos
<emphasis id="7A6A002D6BD7305770BBADC7E5D2339C" bold="true" pageId="0" pageNumber="257">C</emphasis>
MZUTI 2211, adult male, LRC: 14.30 mm, from Cordillera de Guacamayos
<emphasis id="8254308707CD9C410B3810EABFF5B839" bold="true" pageId="0" pageNumber="257">D</emphasis>
MZUTI 2213, adult male, LRC: 15.10 mm, from Cordillera de Guacamayos
<emphasis id="6C090167353220C183772AD4E511A1B5" bold="true" pageId="0" pageNumber="257">E</emphasis>
MZUTI 2214, adult female, LRC: 18.73 mm, from Cordillera de Guacamayos
<emphasis id="B835AE732075540C3C629EE4418AEA03" bold="true" pageId="0" pageNumber="257">F</emphasis>
ZSFQ 774, adult female, LRC:18.96 mm from Sumaco Volcano
<emphasis id="166E937783757640EE4F8E99F7D3CA05" bold="true" pageId="0" pageNumber="257">G</emphasis>
ZSFQ 782, adult female, LRC: 30.03 mm, from Sumaco Volcano. Photographs by Lucas Bustamante (
<emphasis id="947DB7FD5CAD9329AA7F9892E2002DC1" bold="true" pageId="0" pageNumber="257">A, B, C, D, E</emphasis>
) and Jose Vieira (
<emphasis id="FFC240E8762940EFD8C4652B4F0AA6EF" bold="true" pageId="0" pageNumber="257">F, G</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="F6A3D428D87B4FD35A3643745A021993" pageId="0" pageNumber="257" type="generic placement">
<paragraph id="344DB1C3CC46683B8C6A4451E3F85B93" pageId="0" pageNumber="257">Generic placement.</paragraph>
<paragraph id="F520A4C31B0D25C940AB5509EB5A6592" pageId="0" pageNumber="257">
As defined by
<bibRefCitation id="EE88F5D6D653B738DB09159AB6018080" DOI="https://doi.org/10.5962/bhl.title.7951" author="Lynch, JD" journalOrPublisher="Special Publications" pageId="0" pageNumber="257" pagination="1 - 236" refId="B19" refString="Lynch, JD, Duellman, WE, 1997. Frogs of the genus Eleutherodactylus (Leptodactylidae) in Western Ecuador: Systematics, ecology, and biogeography. The University of Kansas Natural History Museum. Special Publications 23: 1 - 236, DOI: https://doi.org/10.5962/bhl.title.7951" title="Frogs of the genus Eleutherodactylus (Leptodactylidae) in Western Ecuador: Systematics, ecology, and biogeography. The University of Kansas Natural History Museum." url="https://doi.org/10.5962/bhl.title.7951" volume="23" year="1997">Lynch and Duellman (1997)</bibRefCitation>
,
<bibRefCitation id="393138A30A4C6C2AB55E8A81054A10A0" DOI="https://doi.org/10.11646/zootaxa.1737.1.1" author="Hedges, SB" journalOrPublisher="Zootaxa" pageId="0" pageNumber="257" pagination="1 - 182" refId="B8" 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="B8C0242767A3E344928454EB55D63BF5" DOI="https://doi.org/10.7717/peerj.14715" author="Franco-Mena, D" journalOrPublisher="Zoosystematics and Evolution" pageId="0" pageNumber="257" refId="B6" refString="Franco-Mena, D, Guayasamin, JM, Andrade-Brito, D, Yanez-Munoz, MH, Rojas-Runjaic, FJM, 2023. Unveiling the evolutionary relationships and the high cryptic diversity in Andean rainfrogs (Craugastoridae: Pristimantis myersi group). PeerJ 11: e14715. https://doi.org/10.7717/peerj.14715" title="Unveiling the evolutionary relationships and the high cryptic diversity in Andean rainfrogs (Craugastoridae: Pristimantis myersi group). PeerJ 11: e 14715." url="https://doi.org/10.7717/peerj.14715" year="2023">Franco-Mena et al. (2023)</bibRefCitation>
, the
<taxonomicName id="713204A0D7B8CA08F783487FDF0E32BC" baseAuthorityName="Goin &amp; Cochran" baseAuthorityYear="1963" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis myersi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="myersi">
<emphasis id="0094A5961E0CBC21665DCF63FED6AF5B" italics="true" pageId="0" pageNumber="257">Pristimantis myersi</emphasis>
</taxonomicName>
group (subgenus
<taxonomicName id="1370FE51248D8DCA36DC55B5B24DAF1D" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis (Trachyphrynus)" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="subGenus" subGenus="Trachyphrynus">Pristimantis Trachyphrynus</taxonomicName>
) contains frogs with the following combination of traits: (1) small body size (SVL in females &lt;34.6 mm; in males &lt;20.5 mm); (2) short snout; (3) robust body; (4) Toe V longer than Toe III, Finger I shorter than II; (5) digital discs narrow or slightly expanded (expanded in
<taxonomicName id="B333C7B57637AFD611E32D7EC1C02EA3" lsidName="P. floridus" pageId="0" pageNumber="257" rank="species" species="floridus">
<emphasis id="EAC6BC1B0FBEDA28ED46CA949A98DAD1" italics="true" pageId="0" pageNumber="257">P. floridus</emphasis>
</taxonomicName>
); and (6) cranial crests absent. In addition, all species in the group are found on low vegetation or at ground level, or even underground. The morphology of the new species agrees with all the aforementioned diagnostic traits.
</paragraph>
</subSubSection>
<subSubSection id="4AA17CFFC8CBD4879F51E9EEA9DF553B" pageId="0" pageNumber="257" type="diagnosis">
<paragraph id="F7FFDC34317917A5AE744313FB7733FF" pageId="0" pageNumber="257">Diagnosis.</paragraph>
<paragraph id="B12DD55FCAA0282F1983400660DB1475" pageId="0" pageNumber="257">
(1) Skin on dorsum shagreen with small scattered, rounded warts; upper flanks with numerous low warts; dorsolateral and w-shaped occipital folds present (Figs
<figureCitation id="58655E75F591E0840F5626AF896BFA67" captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Holotype of Pristimantis kayi sp. nov. in life, adult female, ZSFQ 0775. LRC: 19.7 mm. Photographs by Jose Vieira." figureDoi="10.3897/zookeys.1180.107333.figure13" httpUri="https://binary.pensoft.net/fig/912041" pageId="0" pageNumber="257">13</figureCitation>
-
<figureCitation id="F18660B9E5837D304A425E16F4668DC8" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Preserved holotype of Pristimantis kayi sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257">15</figureCitation>
); (2) tympanic membrane and tympanic annulus well differentiated (Fig.
<figureCitation id="B742F1BEC28812C5FEAAD605E525DD03" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Habitat vegetation at type localities and etymology of the two new frogs described herein. Photographs by JPRP and Zane Libke A panoramic view of the upper Pastaza Valley and its adjacent mountains, main geographic barriers promoting speciation at a regional scale, at back Tungurahua Volcano and mountains south of the Pastaza River habitat of Pristimantis donnelsoni sp. nov., image from the slopes of Cerro Mayordomo type locality of Pristimantis kayi sp. nov., in the northern mountains to the Pastaza River B Tungurahua Volcano eruption, from Chamana Reserve C Montane forest in Candelaria Reserve D Don Nelson Palacios (+) during an expedition; we tribute his work with the naming of Pristimantis donnelsoni sp. nov. E Sumaco Volcano seen from Wawa Sumaku, habitat of P. kayi sp. nov. F Montane forest and subparamo in Leito, Patate G Andreas Kay (+); we honour his devotion to nature photography and research in the upper Pastaza Valley and through Ecuador by naming Pristimantis kayi sp. nov." figureDoi="10.3897/zookeys.1180.107333.figure12" httpUri="https://binary.pensoft.net/fig/912040" pageId="0" pageNumber="257">12</figureCitation>
); tympanic annulus not sexually dimorphic; tympanum in males 7% SVL, tympanum in females 7% SVL; (3) snout rounded in dorsal, protruding in lateral view (Fig.
<figureCitation id="A61C64643F6F3C1592431D3F391B8002" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Preserved holotype of Pristimantis kayi sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257">15</figureCitation>
); (4) upper eyelid usually bearing one conical or subconical tubercle and many low tubercles; upper eyelid about in males 63% IOD, in females 59% IOD; cranial crests absent; (5) dentigerous processes of the vomer evident, each process bearing 2-3 teeth; (6) sphenethmoides on its dorsal view with short blunt anterior border, is short and blunt in ventral view, posterior border present oblique articulation with frontoparietals; posterior border of frontoparietal with an irregular border not projected in dorsal view; zygomatic ramus of squamosal elongated anteriorly, in dorsal view; procesus cultriform of the parasphenoides reaching posterior level of the vomers, rounded anterior border (Fig.
<figureCitation id="C1248BD4F4EB52DE6D3AD6F3141635B8" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Skull morphology diagram in dorsal (left column) and ventral (right column) views in four species of the Pristimantis myersi species group: A, B P. donnelsoni sp. nov., DHMECN 4808 C, D P. kayi sp. nov., DHMECN 14447 E, F P. gladiator DHMECN 12465 G, H P. festae DHMECN 9294. Detail of the skull bones abbreviations: pmax = premaxilla; max = maxilla; spheth = sphenethmoides; fpar = frontoparietal; pter = pterigoides; pro = prootic; q = quadratojugal; sp = squamosal; exo = exoccipital; prph = parasphenoides. Frontoparietal fontanelle not coloured." figureDoi="10.3897/zookeys.1180.107333.figure8" httpUri="https://binary.pensoft.net/fig/912036" pageId="0" pageNumber="257">8</figureCitation>
); (7) males with vocal slits, nuptial pads absent; (8) finger I shorter than finger II; disc on Finger I not expanded; discs on Finger II-IV slightly expanded (Figs
<figureCitation id="20B001C01A68EE478674EB56C148D316" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Pristimantis donnelsoni sp. nov. dorsal and ventral views of the preserved types series A DHMECN 4779, female B DHMECN 4782, female C female, ZSFQ 1097 D holotype, 14701, female E DHMECN 4781, male F DHMECN 16613, male G DHMECN 16610, male H ZSFQ 1098, male I DHMECN 4778, male. Photographs by JPRP and David Brito Zapata." figureDoi="10.3897/zookeys.1180.107333.figure6" httpUri="https://binary.pensoft.net/fig/912034" pageId="0" pageNumber="257">6</figureCitation>
,
<figureCitation id="EF7160850364B3C6648BA2919A097FBA" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Preserved holotype of Pristimantis kayi sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257">15</figureCitation>
); (9) fingers with thin lateral fringes; (10) two or three low ulnar tubercles present; (11) heel with low to conical tubercle; outer edge of tarsus with low conical tubercles; (12) inner metatarsal tubercle oval, about 1.5-2
<normalizedToken id="3F17A2159240C0FDCB6373997C623B03" originalValue="×">x</normalizedToken>
the length of round outer metatarsal tubercle; (13) toes with thin lateral fringes; webbing absent; Toe V slightly longer than Toe III; discs slightly expanded (Figs
<figureCitation id="D6A628E01E47C04B2D4FD4683D71C9BA" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Preserved holotype of Pristimantis kayi sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257">15</figureCitation>
,
<figureCitation id="FE577A510FF610C605EC7A4DE62D0F5B" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">16</figureCitation>
); (14) in life, dorsum brown with darker and lighter markings; venter light grey to black, with or without orange to red spots of different sizes; groin with orange to red spots that are more conspicuous in females than males (see Colour variation; Figs
<figureCitation id="B2CCDC74EDAC5A6175F03CB8362E1A9D" captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Colour variation of Pristimantis kayi sp. nov. A MZUTI 2209, adult female, LRC: 20.15 mm, from Cordillera de Guacamayos B MZUTI 2210, adult female, LRC: 19.90 mm, from Cordillera de Guacamayos C MZUTI 2211, adult male, LRC: 14.30 mm, from Cordillera de Guacamayos D MZUTI 2213, adult male, LRC: 15.10 mm, from Cordillera de Guacamayos E MZUTI 2214, adult female, LRC: 18.73 mm, from Cordillera de Guacamayos F ZSFQ 774, adult female, LRC: 18.96 mm from Sumaco Volcano G ZSFQ 782, adult female, LRC: 30.03 mm, from Sumaco Volcano. Photographs by Lucas Bustamante (A, B, C, D, E) and Jose Vieira (F, G)." figureDoi="10.3897/zookeys.1180.107333.figure14" httpUri="https://binary.pensoft.net/fig/912042" pageId="0" pageNumber="257">14</figureCitation>
,
<figureCitation id="F4B516CD84DCB417B7D9080462118FE3" captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Dorsal and ventral views of preserved type series of Pristimantis kayi sp. nov. A holotype ZSFQ 0775, female; Paratypes B ZSFQ 0779, female C DHMECN 16212, female D MZUTI 5162, female E MZUTI 4657, male F ZSFQ 0773, male G DHMECN 15233, male H DHMECN 18440, male; Photographs by JPRPR and David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure17" httpUri="https://binary.pensoft.net/fig/912045" pageId="0" pageNumber="257">17</figureCitation>
); (15) SVL in adult males, 11.8-15.6 mm (mean = 13.7, SD = 2.7, n = 30), SVL in adult females 12.3-20.2 mm (mean = 16.3, SD = 5.5, n = 13); (16) call composed of 1 to 7 notes, notes have a mean duration of 36.0
<normalizedToken id="2747DE6D4E28D88FAF67C2F1566A00BA" originalValue="±">+/-</normalizedToken>
18.3 ms, the mean interval between notes is 43.8
<normalizedToken id="AFC36EE7A416E93C90BC58CA786750C9" originalValue="±">+/-</normalizedToken>
36.2 ms, emitted at a mean rate of 15.7
<normalizedToken id="7F5016029EEE6C990D6E2A421F3BC8DB" originalValue="±">+/-</normalizedToken>
6.1 notes/second. The notes are composed of 2 to 13 pulses. Pulses have a mean duration of 3.9
<normalizedToken id="9E05927D9D59B1D5E1FA1421A7C460FC" originalValue="±">+/-</normalizedToken>
1.1 ms, with a mean interval between pulses of 4.2
<normalizedToken id="DB54D120019ABEFE733C8DA4302F07BA" originalValue="±">+/-</normalizedToken>
2.8 ms, emitted at a mean rate of 202.2
<normalizedToken id="99D08CBB9AB307D694DD38FAEB4362E7" originalValue="±">+/-</normalizedToken>
119.7 pulses/second; (17) dominant frequency of 3.19
<normalizedToken id="C847B3FA089D45C218CF5781816CE051" originalValue="±">+/-</normalizedToken>
0.05 kHz, with 2-6 partial harmonics in the spectrogram; (18) call duration of 165.1
<normalizedToken id="8BB893C81399D5B8F4AA0336B5BEAECE" originalValue="±">+/-</normalizedToken>
130.1 ms.
</paragraph>
<caption id="47131BB5EE15275B9A8C6E0B6373F487" doi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257" start="Figure 15" startId="F15">
<paragraph id="B5AD7BE6170F4C68E1F0502B4072FE2A" pageId="0" pageNumber="257">
<emphasis id="8D6708A3B2F98A1F9B036C2F0D327DEC" bold="true" pageId="0" pageNumber="257">Figure 15.</emphasis>
Preserved holotype of
<taxonomicName id="0E099BA04063B4B96D5B090FF5749275" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="DE9A39B480894AE09CE7AC2A81FEC96C" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata.
</paragraph>
</caption>
<caption id="9582A854D0DA375DE2C18927BBF10909" doi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257" start="Figure 16" startId="F16">
<paragraph id="B6CBF2F8F3D878DDAC681BB4C47F1FE9" pageId="0" pageNumber="257">
<emphasis id="3BCD032A5E8EAA86D4207636AF7B14C9" bold="true" pageId="0" pageNumber="257">Figure 16.</emphasis>
Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of
<taxonomicName id="E8B8EBC864074933EC42E7A43C5DF311" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="7870720A0B2E89CECED833B9A27724F8" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. Photographs David Brito-Zapata.
</paragraph>
</caption>
<caption id="7251F489CB278751195FBFB8BF920837" doi="10.3897/zookeys.1180.107333.figure17" httpUri="https://binary.pensoft.net/fig/912045" pageId="0" pageNumber="257" start="Figure 17" startId="F17">
<paragraph id="556823E3A088E99CFCA109C27E0DBD53" pageId="0" pageNumber="257">
<emphasis id="383AFD97F7F036D17DA9BFE40B3F88B3" bold="true" pageId="0" pageNumber="257">Figure 17.</emphasis>
Dorsal and ventral views of preserved type series of
<taxonomicName id="DDD8DE8AE3547AF6A235551D1132982A" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="64FC7AFECF4D0F257E9D3EC417C7B8A1" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov.
<emphasis id="DE5450CCF8EC15166604F689D676531E" bold="true" pageId="0" pageNumber="257">A</emphasis>
holotype ZSFQ 0775, female; Paratypes
<emphasis id="5669690CA5E7EFB668BA8BC70F6D68E6" bold="true" pageId="0" pageNumber="257">B</emphasis>
ZSFQ 0779, female
<emphasis id="CBE065445BBAB2173418765E01D845AD" bold="true" pageId="0" pageNumber="257">C</emphasis>
DHMECN 16212, female
<emphasis id="D5872AB737405CEDB5B2D1E43472E963" bold="true" pageId="0" pageNumber="257">D</emphasis>
MZUTI 5162, female
<emphasis id="43A663C53C9F213AAB892C00666A6DC0" bold="true" pageId="0" pageNumber="257">E</emphasis>
MZUTI 4657, male
<emphasis id="6C509023AB977A586607E4A311027F67" bold="true" pageId="0" pageNumber="257">F</emphasis>
ZSFQ 0773, male
<emphasis id="B8668E365489C1A5350411CBAF247FF7" bold="true" pageId="0" pageNumber="257">G</emphasis>
DHMECN 15233, male
<emphasis id="F9E3BD46F34229BAA188F948DE0F028A" bold="true" pageId="0" pageNumber="257">H</emphasis>
DHMECN 18440, male; Photographs by JPRPR and David Brito-Zapata.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="B87F479872C1303B3E6D5627EC7A494B" pageId="0" pageNumber="257" type="comparisons">
<paragraph id="8DC53ABD63809E9B4A30EAA4F7D4C837" pageId="0" pageNumber="257">Comparison with other species</paragraph>
<paragraph id="DE1DD3A1DC08237C947A52E149877628" pageId="0" pageNumber="257">
<emphasis id="B871CB1C71C6676E1A41A7BD4053E2B7" bold="true" pageId="0" pageNumber="257">
(Figs
<figureCitation id="6A6B7F17F667413D938A4472B42E9CA5" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Dorsolateral and ventral views in life of Pristimantis gladiator species complex and related species A Pristimantis donnelsoni sp. nov. female, DHMECN 14701, holotype, LRC: 15.42 mm B Pristimantis donnelsoni sp. nov. male, DHMECN 18854, LRC: 13.66 mm from Cerro Candelaria C Pristimantis kayi sp. nov. female, ZSFQ 0775, holotype, LRC: 19.7 mm D Pristimantis kayi sp. nov. male, DHMECN 14450 LRC: 13.58 mm, from Reserva Machay E Pristimantis gladiator sensu stricto female, DHMECN 12463, LRC: 20.38 mm F Pristimantis gladiator sensu stricto, male DHMECN 12474, LRC: 16.87 mm; from Rio Quijos drainage G Pristimantis festae female, DHMECN 16280, LRC: 20.19 mm H Pristimantis festae, male (left), DHMECN 16837, LRC: 16.50 mm; female (right), DHMECN 16836, LRC: 19.80 mm; Photographers: JPRP (A, B), MYM (D, E, F, H); Patricia Bejarano (G); Jose Vieira (C)." figureDoi="10.3897/zookeys.1180.107333.figure5" httpUri="https://binary.pensoft.net/fig/912033" pageId="0" pageNumber="257">5</figureCitation>
,
<figureCitation id="636F8B862C415716BE1FECF76D4671CA" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Skull morphology diagram in dorsal (left column) and ventral (right column) views in four species of the Pristimantis myersi species group: A, B P. donnelsoni sp. nov., DHMECN 4808 C, D P. kayi sp. nov., DHMECN 14447 E, F P. gladiator DHMECN 12465 G, H P. festae DHMECN 9294. Detail of the skull bones abbreviations: pmax = premaxilla; max = maxilla; spheth = sphenethmoides; fpar = frontoparietal; pter = pterigoides; pro = prootic; q = quadratojugal; sp = squamosal; exo = exoccipital; prph = parasphenoides. Frontoparietal fontanelle not coloured." figureDoi="10.3897/zookeys.1180.107333.figure8" httpUri="https://binary.pensoft.net/fig/912036" pageId="0" pageNumber="257">8</figureCitation>
,
<figureCitation id="54D1C5D3B28D0B2F88CCA63BF90C76C3" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Boxplot of body size (SVL) and tympanic diameter (TD) in four closely-related species of the Pristimantis myersi species group. * Asterisks represent species with significant differences." figureDoi="10.3897/zookeys.1180.107333.figure9" httpUri="https://binary.pensoft.net/fig/912037" pageId="0" pageNumber="257">9</figureCitation>
).
</emphasis>
<taxonomicName id="D7264B8363E83F3BDCA058B8145F79CE" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="19EE946900680BB662F7CCABE26591D7" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. is most similar to
<taxonomicName id="13CBE8D5C05D9F6050A9FE1B4CDB37DB" lsidName="P. gladiator" pageId="0" pageNumber="257" rank="species" species="gladiator">
<emphasis id="4D92F7CC1ACED2E379509A57D69F7EF5" italics="true" pageId="0" pageNumber="257">P. gladiator</emphasis>
</taxonomicName>
(Lynch, 1976),
<taxonomicName id="950DB7098A203460052BDCF3DD03E1F1" lsidName="P. festae" pageId="0" pageNumber="257" rank="species" species="festae">
<emphasis id="A509245A34E6A9FF9E920E88A6F231F2" italics="true" pageId="0" pageNumber="257">P. festae</emphasis>
</taxonomicName>
(Peracca, 1904), and
<taxonomicName id="F860D0683251B6D0D88C4BB9D372AB3C" lsidName="P. donnelsoni" pageId="0" pageNumber="257" rank="species" species="donnelsoni">
<emphasis id="715A18ACCE2F2DEAD1B418301A446245" italics="true" pageId="0" pageNumber="257">P. donnelsoni</emphasis>
</taxonomicName>
sp. nov.
<taxonomicName id="4E358CCF7F598287ED5F7CEC969BE7EC" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="40E66EE64DF4BF74BA71DA1C52D6F2B0" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. differs from
<taxonomicName id="652171FE586908F236BBEA03A66D0B2D" lsidName="P. gladiator" pageId="0" pageNumber="257" rank="species" species="gladiator">
<emphasis id="FE4DF5C22892AC7485598A97C5A251F8" italics="true" pageId="0" pageNumber="257">P. gladiator</emphasis>
</taxonomicName>
and
<taxonomicName id="961A10DF79FD786D958A7B50D6F72A23" lsidName="P. festae" pageId="0" pageNumber="257" rank="species" species="festae">
<emphasis id="706B969ACDB9EDE630E370188AC2C0A8" italics="true" pageId="0" pageNumber="257">P. festae</emphasis>
</taxonomicName>
by being smaller (Kruskal-Wallis Chi2 = 25.39, p = &lt;0.001, Table
<tableCitation id="9B420872DFACFB02CB827AEE86CCFBC2" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Measurements of four closely-related species of Pristimantis from the eastern Ecuadorian Andes." httpUri="http://table.plazi.org/id/070103EF30A585DD1BF82638AFEF4D1D" pageId="0" pageNumber="257" tableUuid="070103EF30A585DD1BF82638AFEF4D1D">2</tableCitation>
, Fig.
<figureCitation id="91A04FCF936018A306DEECEF840383E2" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Boxplot of body size (SVL) and tympanic diameter (TD) in four closely-related species of the Pristimantis myersi species group. * Asterisks represent species with significant differences." figureDoi="10.3897/zookeys.1180.107333.figure9" httpUri="https://binary.pensoft.net/fig/912037" pageId="0" pageNumber="257">9</figureCitation>
). Additionally,
<taxonomicName id="EEFB3D97ECED3FF2CC8DC7B55C7A8AE3" lsidName="P. kayi" pageId="0" pageNumber="257" rank="species" species="kayi">
<emphasis id="BB5B9CCAE2DEB7B7898FF16BBFF6432D" italics="true" pageId="0" pageNumber="257">P. kayi</emphasis>
</taxonomicName>
sp. nov. differs from
<taxonomicName id="0ADD6A419E0C08120F1BC01CC2DC21DF" lsidName="P. festae" pageId="0" pageNumber="257" rank="species" species="festae">
<emphasis id="72A2149889E5D53CCF57A3592BF77604" italics="true" pageId="0" pageNumber="257">P. festae</emphasis>
</taxonomicName>
by having orange to red spots on the groin (white to reddish large spots in
<taxonomicName id="56BBD3575212D98C64DDC5CDA6559BCA" lsidName="P. festae" pageId="0" pageNumber="257" rank="species" species="festae">
<emphasis id="8C3A84F84786911DD3AF34F9A232D7A5" italics="true" pageId="0" pageNumber="257">P. festae</emphasis>
</taxonomicName>
; Fig.
<figureCitation id="E1194D5ECB282903902EB4CC40D1DA67" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Dorsolateral and ventral views in life of Pristimantis gladiator species complex and related species A Pristimantis donnelsoni sp. nov. female, DHMECN 14701, holotype, LRC: 15.42 mm B Pristimantis donnelsoni sp. nov. male, DHMECN 18854, LRC: 13.66 mm from Cerro Candelaria C Pristimantis kayi sp. nov. female, ZSFQ 0775, holotype, LRC: 19.7 mm D Pristimantis kayi sp. nov. male, DHMECN 14450 LRC: 13.58 mm, from Reserva Machay E Pristimantis gladiator sensu stricto female, DHMECN 12463, LRC: 20.38 mm F Pristimantis gladiator sensu stricto, male DHMECN 12474, LRC: 16.87 mm; from Rio Quijos drainage G Pristimantis festae female, DHMECN 16280, LRC: 20.19 mm H Pristimantis festae, male (left), DHMECN 16837, LRC: 16.50 mm; female (right), DHMECN 16836, LRC: 19.80 mm; Photographers: JPRP (A, B), MYM (D, E, F, H); Patricia Bejarano (G); Jose Vieira (C)." figureDoi="10.3897/zookeys.1180.107333.figure5" httpUri="https://binary.pensoft.net/fig/912033" pageId="0" pageNumber="257">5</figureCitation>
) and slightly expanded discs on outer fingers (discs not expanded in
<taxonomicName id="33B9D28C09A3C88984AE3AD9AEC829E4" lsidName="P. festae" pageId="0" pageNumber="257" rank="species" species="festae">
<emphasis id="06E05004E9651007629DEC802A5B9085" italics="true" pageId="0" pageNumber="257">P. festae</emphasis>
</taxonomicName>
).
<taxonomicName id="D3340B98B2031DBA695E97703677DFC9" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="F6E76817776647EBA420DCB473DECBA5" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. is similar to the allopatric
<taxonomicName id="3ACF4EDCBF72E89B15328CD3F27B14E0" lsidName="P. donnelsoni" pageId="0" pageNumber="257" rank="species" species="donnelsoni">
<emphasis id="EC6AA5C9916BCE3947EE3812A6699FEF" italics="true" pageId="0" pageNumber="257">P. donnelsoni</emphasis>
</taxonomicName>
sp. nov.; however, the diameter of the tympanum in males is larger in
<taxonomicName id="8928CC1A357F0EE6E46602BC199FBDDB" lsidName="P. kayi" pageId="0" pageNumber="257" rank="species" species="kayi">
<emphasis id="1AB7948946C70569BC307AE01A437FD4" italics="true" pageId="0" pageNumber="257">P. kayi</emphasis>
</taxonomicName>
sp. nov. compared to
<taxonomicName id="C592ED0E485D816417BF6C0A7E3AD5B8" lsidName="P. donnelsoni" pageId="0" pageNumber="257" rank="species" species="donnelsoni">
<emphasis id="6781ECC541C83F3F5C78DA644B915AE6" italics="true" pageId="0" pageNumber="257">P. donnelsoni</emphasis>
</taxonomicName>
sp. nov. and the other closely-related species (i.e.
<taxonomicName id="AA10BD78A5274C060B9E6569D8981578" lsidName="P. festae" pageId="0" pageNumber="257" rank="species" species="festae">
<emphasis id="F0926CB591BA8BA96099A053027A4056" italics="true" pageId="0" pageNumber="257">P. festae</emphasis>
</taxonomicName>
and
<taxonomicName id="4E3F20250321BAFD6012F100EA5410DB" lsidName="P. gladiator" pageId="0" pageNumber="257" rank="species" species="gladiator">
<emphasis id="527ADE239DEAFC989C7683373447C66D" italics="true" pageId="0" pageNumber="257">P. gladiator</emphasis>
</taxonomicName>
) (Kruskal-Wallis Chi2 = 10.28, p = &lt;0.05*, Table
<tableCitation id="3047C938C1A9DED990D57BDB1D85B705" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Measurements of four closely-related species of Pristimantis from the eastern Ecuadorian Andes." httpUri="http://table.plazi.org/id/070103EF30A585DD1BF82638AFEF4D1D" pageId="0" pageNumber="257" tableUuid="070103EF30A585DD1BF82638AFEF4D1D">2</tableCitation>
, Fig.
<figureCitation id="334D50FA7DF814D491E8CF19FAC13819" captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Boxplot of body size (SVL) and tympanic diameter (TD) in four closely-related species of the Pristimantis myersi species group. * Asterisks represent species with significant differences." figureDoi="10.3897/zookeys.1180.107333.figure9" httpUri="https://binary.pensoft.net/fig/912037" pageId="0" pageNumber="257">9</figureCitation>
). The most conspicuous differences in the skull morphology between the two new species are listed below (Fig.
<figureCitation id="747CE39273B3BEE39873CDF91DB277C2" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Skull morphology diagram in dorsal (left column) and ventral (right column) views in four species of the Pristimantis myersi species group: A, B P. donnelsoni sp. nov., DHMECN 4808 C, D P. kayi sp. nov., DHMECN 14447 E, F P. gladiator DHMECN 12465 G, H P. festae DHMECN 9294. Detail of the skull bones abbreviations: pmax = premaxilla; max = maxilla; spheth = sphenethmoides; fpar = frontoparietal; pter = pterigoides; pro = prootic; q = quadratojugal; sp = squamosal; exo = exoccipital; prph = parasphenoides. Frontoparietal fontanelle not coloured." figureDoi="10.3897/zookeys.1180.107333.figure8" httpUri="https://binary.pensoft.net/fig/912036" pageId="0" pageNumber="257">8</figureCitation>
): in
<taxonomicName id="7C335445114967EF4D3397B415298105" lsidName="P. kayi" pageId="0" pageNumber="257" rank="species" species="kayi">
<emphasis id="7F859A777370884E766EF7D8EF933E2A" italics="true" pageId="0" pageNumber="257">P. kayi</emphasis>
</taxonomicName>
sp. nov., the parasphenoid is much more developed than in
<taxonomicName id="0155D089C82558EEA291484E98AFEED9" lsidName="P. donnelsoni" pageId="0" pageNumber="257" rank="species" species="donnelsoni">
<emphasis id="2959461111C0A79A3B35B6BFA7A655EA" italics="true" pageId="0" pageNumber="257">P. donnelsoni</emphasis>
</taxonomicName>
sp. nov., with its lateral alary processes almost reaching the squamosal; also, the palatine bones are shorter in
<taxonomicName id="AE577404DF9F1656274AAFB11140A1EE" lsidName="P. kayi" pageId="0" pageNumber="257" rank="species" species="kayi">
<emphasis id="03F9F90AF6FD72BC0ED32DC56488E7D2" italics="true" pageId="0" pageNumber="257">P. kayi</emphasis>
</taxonomicName>
sp. nov. than in
<taxonomicName id="2125C0364F25B4E98BE669D0D1CED329" lsidName="P. donnelsoni" pageId="0" pageNumber="257" rank="species" species="donnelsoni">
<emphasis id="AB8BE5D836014C3CDA93AA9904FA9E89" italics="true" pageId="0" pageNumber="257">P. donnelsoni</emphasis>
</taxonomicName>
sp. nov. Finally, the zygomatic ramus of the squamosal is much longer in
<taxonomicName id="F39F15F175307B89C0E852EFCE3ED864" lsidName="P. kayi" pageId="0" pageNumber="257" rank="species" species="kayi">
<emphasis id="73E3821B9B9F1252D3B096EB728A22D2" italics="true" pageId="0" pageNumber="257">P. kayi</emphasis>
</taxonomicName>
sp. nov. than in
<taxonomicName id="494EF62129485A106DA3CB108B81EEE7" lsidName="P. donnelsoni" pageId="0" pageNumber="257" rank="species" species="donnelsoni">
<emphasis id="F9F8E01CCA5B3C7F71E5A097777438C8" italics="true" pageId="0" pageNumber="257">P. donnelsoni</emphasis>
</taxonomicName>
sp. nov. (Fig.
<figureCitation id="565595625F33CE2B17053EC92D8B0216" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Skull morphology diagram in dorsal (left column) and ventral (right column) views in four species of the Pristimantis myersi species group: A, B P. donnelsoni sp. nov., DHMECN 4808 C, D P. kayi sp. nov., DHMECN 14447 E, F P. gladiator DHMECN 12465 G, H P. festae DHMECN 9294. Detail of the skull bones abbreviations: pmax = premaxilla; max = maxilla; spheth = sphenethmoides; fpar = frontoparietal; pter = pterigoides; pro = prootic; q = quadratojugal; sp = squamosal; exo = exoccipital; prph = parasphenoides. Frontoparietal fontanelle not coloured." figureDoi="10.3897/zookeys.1180.107333.figure8" httpUri="https://binary.pensoft.net/fig/912036" pageId="0" pageNumber="257">8</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="1E106F95F704D52D3809F65AC8C0CAE0" pageId="0" pageNumber="257" type="description">
<paragraph id="34218A35D61A65F148BBA7F113776D8F" pageId="0" pageNumber="257">Description of the holotype</paragraph>
<paragraph id="A82E98705AB1099AB93B1BDB5AAEAA8C" pageId="0" pageNumber="257">
<emphasis id="708CAE349F609E38E788142F8612E534" bold="true" pageId="0" pageNumber="257">
(Figs
<figureCitation id="D5D139C8020FE7385D827A8761746BE2" captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Holotype of Pristimantis kayi sp. nov. in life, adult female, ZSFQ 0775. LRC: 19.7 mm. Photographs by Jose Vieira." figureDoi="10.3897/zookeys.1180.107333.figure13" httpUri="https://binary.pensoft.net/fig/912041" pageId="0" pageNumber="257">13</figureCitation>
,
<figureCitation id="439D34AF5F923ABA559848237C33C5B6" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Preserved holotype of Pristimantis kayi sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257">15</figureCitation>
,
<figureCitation id="479386CEE31AA6EAAC308A0AF001B8EB" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">16</figureCitation>
).
</emphasis>
Adult female (ZSFQ 0775) robust body; head slightly longer than wide, not as wide as body, head width 35.27% of SVL (19.7); head length 38% of SVL; snout rounded in dorsal view, protruding in lateral view; eye-nostril distance 9% of SVL; with small papilla at tip (Fig.
<figureCitation id="F3A62E272F1573A3541FAE1E6AA4999F" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Evolutionary relationships amongst species in the Pristimantis gladiator complex, under the Maximum Likelihood inference criterion (modified from Franco-Mena et al. (2023)). Red font indicates the new sequence." figureDoi="10.3897/zookeys.1180.107333.figure1" httpUri="https://binary.pensoft.net/fig/912029" pageId="0" pageNumber="257">1</figureCitation>
); in lateral view, distinct rostral ridge; loreal region slightly concave; nostrils protruding, laterally directed; interorbital area flat, wider than upper eyelid (upper eyelid width 62% of OID); cranial ridges absent; upper eyelid with one conical and several non-conical tubercles; tympanic membrane well defined, pigmented like surrounding skin; tympanic ring distinct, round; supratympanic fold present, obscuring anterodorsal and posterodorsal edges of the ring; tympanic diameter 46% of eye length; several low to conical tubercles situated in the area just posterior to tympanum. Choanae are small, with an oval to square shape, not concealed by the palatal shelf of maxillary; the dentigerous process of the vomer oblique, widely separated, posteromedial to choanae, each bearing two small teeth; tongue slightly longer than wide, granular, with a conspicuous notch along the posterior border.
</paragraph>
<paragraph id="8D38FCAF132956111CB9394AE88EB65A" pageId="0" pageNumber="257">
The skin of the head is greyish; the dorsum greyish, with scattered small tubercles, some of which are aligned over a W-shaped occipital marking, forming folds; upper flanks with numerous low warts; venter slightly areolate; discoidal fold absent; the cloacal sheath is absent; fingers with thin lateral bangs; length of fingers I &lt;II &lt;IV &lt;III; palmar tubercle round, thenar tubercle oval (Fig.
<figureCitation id="6EB9A50BF95C051718592233A88464BB" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">16</figureCitation>
); subarticular tubercles round, not prominent; supernumerary palmar tubercles not evident; disc sheath of Finger I not expanded; those of Fingers II-IV slightly expanded; all disc sheaths with nearly elliptical ventral pads defined by grooves (Fig.
<figureCitation id="45B85E43B29E1E9D3A6A4F160291461D" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">16</figureCitation>
).
</paragraph>
<paragraph id="63989DA751437719D44420BDF712D913" pageId="0" pageNumber="257">
Hind limbs relatively robust; tibia length 47% SVL; foot length slightly smaller than tibia length (foot length 44% SVL); tarsal tubercles present; small, conical tubercle on heel; toes with narrow lateral fringes (Fig.
<figureCitation id="AB5D457D4E27FDF9707087BB7C995209" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">16</figureCitation>
); subarticular tubercles round, not prominent; inner metatarsal tubercle oval, two times the size of outer tubercle; few low supernumerary plantar tubercles present (Fig.
<figureCitation id="C3382E444B835AB82C26DC51CDDD027E" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">16</figureCitation>
); disc covers of Toes slightly expanded; toes with defined pads; disc pads nearly elliptical; toe lengths I &lt;II &lt;III &lt;V &lt;IV (Fig.
<figureCitation id="94DC1F8166B047C6171694A03B104B8D" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">16</figureCitation>
); the tip of Toe V reaches the middle level of penultimate subarticular tubercle of Toe IV; the tip of Toe III almost reaches the proximal border of penultimate subarticular tubercle of Toe IV.
</paragraph>
</subSubSection>
<subSubSection id="D3B32480305B799651D0002E4C4C35C5" pageId="0" pageNumber="257" type="description">
<paragraph id="3475A87510F5916C9B5462C154DA4D84" pageId="0" pageNumber="257">Measurements of the holotype</paragraph>
<paragraph id="3DD3ECD9155034C7BEAB241CF19D5058" pageId="0" pageNumber="257">
<emphasis id="1E193C08DDB9E380D9944D304EDE8D14" bold="true" pageId="0" pageNumber="257">(in mm).</emphasis>
Adult female, ZSFQ 0775. SVL = 19.7; Tibia Length = 9.3; Foot Length = 8.6; Hand Length = 5.0; Head Length = 7.5; Head Width = 7.0; Eye Diameter = 2.4; Tympanum Diameter = 1.1; forearm length = 4.4; snout length = 5.6; Tarsus length = 5.3; Thigh Length = 8.3; Upper arm Length = 3.1; Interorbital Distance = 2.6; Upper Eyelid Width = 1.6; Internarial Distance = 2.1; Eye-Nostril distance = 1.8; Finger III Width = 0.8; Toe IV Width = 0.3.
</paragraph>
</subSubSection>
<subSubSection id="BE13B16AB0128FFAC58836BC7C38959D" pageId="0" pageNumber="257" type="colour of holotype in life">
<paragraph id="5D8C8FAB7B9D4DFA04CA207C877106A6" pageId="0" pageNumber="257">Colour of holotype in life</paragraph>
<paragraph id="622BE4DCF390A601752E6352916FB6CF" pageId="0" pageNumber="257">
<emphasis id="A597AB77C3A13C45B2C5141892A693F0" bold="true" pageId="0" pageNumber="257">
(Fig.
<figureCitation id="7C1AAE91481F83A484B338000E9ED272" captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Holotype of Pristimantis kayi sp. nov. in life, adult female, ZSFQ 0775. LRC: 19.7 mm. Photographs by Jose Vieira." figureDoi="10.3897/zookeys.1180.107333.figure13" httpUri="https://binary.pensoft.net/fig/912041" pageId="0" pageNumber="257">13</figureCitation>
).
</emphasis>
Head, sides of the head, dorsum, flanks and limbs brown, with darker interorbital bar, lips banded with light and dark brown; broad dark brown supratympanic band extending from region anterior to tympanum to before arm insertion. Flanks lighter than dorsum with oblique, dark bands, several scattered whitish markings near the groin and a white spot in the axilla; limbs banded in shades of dark brown. The throat and belly are dark brown and covered with scattered small yellowish-white spots; the lower ventral part has orange spots; the ventral surfaces of the forelimbs are lighter brown. Iris golden with dark brown reticulations, with a reddish-copper horizontal bar.
</paragraph>
</subSubSection>
<subSubSection id="E13036D521432620E6C71EAE887FD810" pageId="0" pageNumber="257" type="colouration of holotype in ethanol">
<paragraph id="B3BAECF96FD5A5D0429A117524B07858" pageId="0" pageNumber="257">Colouration of holotype in ethanol</paragraph>
<paragraph id="AFB86DCD874A7A4F33624A658605AEC1" pageId="0" pageNumber="257">
<emphasis id="DFD7C32267FD9810BFAB110F9F06FF41" bold="true" pageId="0" pageNumber="257">
(Figs
<figureCitation id="95275412622AE0264C9CEE25E2440BA0" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Preserved holotype of Pristimantis kayi sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257">15</figureCitation>
,
<figureCitation id="8FC4E67DD8B77065AE9A2274F6CAB908" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">16</figureCitation>
).
</emphasis>
The holotype has the following colour pattern: head pale brown with darker interorbital and labial bars; black supratympanic stripe; dorsum brown with several darker markings. Flanks grey to black with minute white spots and lighter diagonal stripes. Groin black with large white spots. The dorsal surfaces of limbs are pale brown with dark brown bars. Cloacal region pale brown, delimited by supracloacal black stripe. The anterior and posterior surfaces of the thighs are brown with several medium to large cream spots. Throat and chest cream-brown with darker marks on the centre (Fig.
<figureCitation id="E751FABB39172CE9A34EF0C3D8287684" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Preserved holotype of Pristimantis kayi sp. nov. ZSFQ 0775, on its dorsal, ventral and lateral views. Photographs by David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure15" httpUri="https://binary.pensoft.net/fig/912043" pageId="0" pageNumber="257">15</figureCitation>
). Venter dark grey, with minute to large white spots. Palms and soles are mostly cream, with some pigmentation of Finger IV and Toe V (Fig.
<figureCitation id="6E7F21D1E3B4256BB16DD59F1399D269" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Detailed ventral view of hand and feet of the holotype ZSFQ 0775 of Pristimantis kayi sp. nov. Photographs David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure16" httpUri="https://binary.pensoft.net/fig/912044" pageId="0" pageNumber="257">16</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C11FEDB4EEB5A9F6E2CF7B28A64391ED" pageId="0" pageNumber="257" type="osteology of the skull">
<paragraph id="E84DC265631AFBD40DA4125ADA9DEB55" pageId="0" pageNumber="257">Osteology of the skull.</paragraph>
<paragraph id="84D61DE7D26D88D169E2BE6B65715886" pageId="0" pageNumber="257">
The skull of the adult female paratype DHMECN 14447 is illustrated on its dorsal and ventral surfaces in Fig.
<figureCitation id="BBE0E464E9908834002DEA38ED04B4E3" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Skull morphology diagram in dorsal (left column) and ventral (right column) views in four species of the Pristimantis myersi species group: A, B P. donnelsoni sp. nov., DHMECN 4808 C, D P. kayi sp. nov., DHMECN 14447 E, F P. gladiator DHMECN 12465 G, H P. festae DHMECN 9294. Detail of the skull bones abbreviations: pmax = premaxilla; max = maxilla; spheth = sphenethmoides; fpar = frontoparietal; pter = pterigoides; pro = prootic; q = quadratojugal; sp = squamosal; exo = exoccipital; prph = parasphenoides. Frontoparietal fontanelle not coloured." figureDoi="10.3897/zookeys.1180.107333.figure8" httpUri="https://binary.pensoft.net/fig/912036" pageId="0" pageNumber="257">8</figureCitation>
. We describe the main skull bones with diagnostic characters in Table
<tableCitation id="B660DD0CF369A1F617E4B2DD1764FEBD" captionStart="Tablе 3" captionStartId="T3" captionText="Table 3. Skull diagnostic characters between four related species of Pristimantis gladiator complex." httpUri="http://table.plazi.org/id/D9FCB4E7CAA28E280FB2217A37F05D4B" pageId="0" pageNumber="257" tableUuid="D9FCB4E7CAA28E280FB2217A37F05D4B">3</tableCitation>
. Skull is slightly longer than wide. Dorsally paired nasals overlap the sphenethmoid; the sphenethmoid articulates posteriorly with the frontoparietals, posterior border of frontoparietal is irregular and not projected. A large frontoparietal fontanelle, connected to two parietal fontanelles, is delimited by the sphenethmoid and the frontoparietals.
</paragraph>
<paragraph id="6005D367146802292A0F0DF536DE21CC" pageId="0" pageNumber="257">
In ventral view, anterior margin of the sphenethmoid is rounded. The palatines are relatively short and overlap the sphenethmoid on its anterior portion. Vomers are narrowly separated medially; each vomer has three distinctive rami; the dentigerous process of the vomer does not reach the level of the palatine and almost contacts the tip of the cultriform process of the parasphenoid. The large parasphenoid presents the shape of an inverted cross; the alary processes are directed transversely, partially covering the otic area and almost reaching the squamosal; the posterior process of the parasphenoid almost reaches the foramen magnum. In the dorsal view, the long zygomatic ramus of the squamosal presents an elongated and acuminated anterior border that extends towards the level of the maxilla (Fig.
<figureCitation id="3692C03C6633DB8D0745F50CAE16A568" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Skull morphology diagram in dorsal (left column) and ventral (right column) views in four species of the Pristimantis myersi species group: A, B P. donnelsoni sp. nov., DHMECN 4808 C, D P. kayi sp. nov., DHMECN 14447 E, F P. gladiator DHMECN 12465 G, H P. festae DHMECN 9294. Detail of the skull bones abbreviations: pmax = premaxilla; max = maxilla; spheth = sphenethmoides; fpar = frontoparietal; pter = pterigoides; pro = prootic; q = quadratojugal; sp = squamosal; exo = exoccipital; prph = parasphenoides. Frontoparietal fontanelle not coloured." figureDoi="10.3897/zookeys.1180.107333.figure8" httpUri="https://binary.pensoft.net/fig/912036" pageId="0" pageNumber="257">8</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="66CDF44100EEE5CDCEE8AB572F5CF680" pageId="0" pageNumber="257" type="variation">
<paragraph id="28817E2F6DF1D4E30E3925B60D174A85" pageId="0" pageNumber="257">Variation</paragraph>
<paragraph id="4740ECF9F5DC2086DE22B87DEB77C5CD" pageId="0" pageNumber="257">
<emphasis id="09D31E31BA55B3C61EBC71B445C6A50F" bold="true" pageId="0" pageNumber="257">
(Figs
<figureCitation id="3693FFF491F5461FFFF5F19F63FBA71C" captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Colour variation of Pristimantis kayi sp. nov. A MZUTI 2209, adult female, LRC: 20.15 mm, from Cordillera de Guacamayos B MZUTI 2210, adult female, LRC: 19.90 mm, from Cordillera de Guacamayos C MZUTI 2211, adult male, LRC: 14.30 mm, from Cordillera de Guacamayos D MZUTI 2213, adult male, LRC: 15.10 mm, from Cordillera de Guacamayos E MZUTI 2214, adult female, LRC: 18.73 mm, from Cordillera de Guacamayos F ZSFQ 774, adult female, LRC: 18.96 mm from Sumaco Volcano G ZSFQ 782, adult female, LRC: 30.03 mm, from Sumaco Volcano. Photographs by Lucas Bustamante (A, B, C, D, E) and Jose Vieira (F, G)." figureDoi="10.3897/zookeys.1180.107333.figure14" httpUri="https://binary.pensoft.net/fig/912042" pageId="0" pageNumber="257">14</figureCitation>
,
<figureCitation id="C7160CC82B1E30AADED026D78F71D8DF" captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Dorsal and ventral views of preserved type series of Pristimantis kayi sp. nov. A holotype ZSFQ 0775, female; Paratypes B ZSFQ 0779, female C DHMECN 16212, female D MZUTI 5162, female E MZUTI 4657, male F ZSFQ 0773, male G DHMECN 15233, male H DHMECN 18440, male; Photographs by JPRPR and David Brito-Zapata." figureDoi="10.3897/zookeys.1180.107333.figure17" httpUri="https://binary.pensoft.net/fig/912045" pageId="0" pageNumber="257">17</figureCitation>
).
</emphasis>
Dorsal surfaces with various shades of brown, with or without darker bands or bars. Darker facial markings such as labial, canthal and interorbital bars are usually present. Dorsal mark is an inverted
<normalizedToken id="2236145C2CDD0243D1FC120567470D6A" originalValue="“V”">&quot;V&quot;</normalizedToken>
in the coccygeal region; other patterns include arrangements of thin lines arranged longitudinally. Dorsal surface of hands and feet with irregular brown spots. The fore- and hind limbs are banded with dark brown, separated by light brown interspaces. Flanks with various shades of brown, usually lighter than dorsum. Hind surfaces of thighs with black or dark brown bars and cream interspaces. Groin with orange and sometimes yellowish-white spots on an orange background. Belly brown; sometimes mottled. The belly may have numerous orange to yellowish-white spots. In some specimens, there is a yellowish-white cross on the belly. Iris is golden to bronze with fine black reticulations and a reddish-copper horizontal middle stripe.
</paragraph>
</subSubSection>
<subSubSection id="F9E3CF5C10BE8B3E1F280E468FEE3343" pageId="0" pageNumber="257" type="description">
<paragraph id="D51FEB13CA4A3EA511FD7ABABC4A2FA7" pageId="0" pageNumber="257">Call description</paragraph>
<paragraph id="D96EFDB7765F822C2898F2FA346794E2" pageId="0" pageNumber="257">
<emphasis id="98652DDAEDE8EC42AD24E8852C544403" bold="true" pageId="0" pageNumber="257">
(Fig.
<figureCitation id="1E5960ABB3A8D6EBF983E4618AC1B961" captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Call of Pristimantis kayi sp. nov. (adult male, MZUTI 852). The species has different types of calls, represented as follows: A single note call B pulsed note calls C complex calls (non-pulsed and pulsed notes)." figureDoi="10.3897/zookeys.1180.107333.figure18" httpUri="https://binary.pensoft.net/fig/912046" pageId="0" pageNumber="257">18</figureCitation>
).
</emphasis>
The description is based on the call of an adult male (MZUTI 852), recorded during the night (21:40 h) by Italo Tapia on 7 June 2012, at the Cordillera de los Guacamayos, Province of Napo, Ecuador. The male was calling from leaf litter at ground level. The air temperature was 12.2 °C. The recording consisted of 21 calls, 37 notes and 856 pulses. The call of
<taxonomicName id="C3F6BB03133687ACB59400CE1E34EF7B" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="A368E150B28D5A27AD7FCAAA7F254B97" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. is composed of the emission of several elements with different characteristics, ranging from single notes to pulsed notes. The call (Fig.
<figureCitation id="A0393FBF852ADC7F65CB582F3438507C" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Evolutionary relationships amongst species in the Pristimantis gladiator complex, under the Maximum Likelihood inference criterion (modified from Franco-Mena et al. (2023)). Red font indicates the new sequence." figureDoi="10.3897/zookeys.1180.107333.figure1" httpUri="https://binary.pensoft.net/fig/912029" pageId="0" pageNumber="257">1</figureCitation>
) has a mean dominant frequency of 3.19
<normalizedToken id="D2DC4E28151288F1AEDEB2E5F1559BA4" originalValue="±">+/-</normalizedToken>
0.05 kHz, with 2-6 partial harmonics in the spectrogram. It has a mean duration of 165.1
<normalizedToken id="53485B43F3777BD2CC5F27E1DF90FAFA" originalValue="±">+/-</normalizedToken>
130.1 ms. The mean interval between calls is 4565.6
<normalizedToken id="E74E9111FEA158D3889B7B78E50E1F43" originalValue="±">+/-</normalizedToken>
5742.3 ms, emitted at a mean rate of 32.6
<normalizedToken id="A3D03D2C3617D21EEA8DB31207BE2084" originalValue="±">+/-</normalizedToken>
23.2 calls/minute. Calls are composed of 1 to 7 notes. Notes have a mean duration of 36.0
<normalizedToken id="A9FF97EF92584BC995E6CB54FBD33E46" originalValue="±">+/-</normalizedToken>
18.3 ms. The mean interval between notes is 43.8
<normalizedToken id="E459DE0F7EFE84799590BE98BD04099D" originalValue="±">+/-</normalizedToken>
36.2 ms, emitted at a mean rate of 15.7
<normalizedToken id="CB356A22690C37D9EB99F0E68DD9586A" originalValue="±">+/-</normalizedToken>
6.1 notes/second. The notes are composed of 2 to 13 pulses. Pulses have a mean duration of 3.9
<normalizedToken id="FABA3617A9A5CAE71DA1A019A748B979" originalValue="±">+/-</normalizedToken>
1.1 ms, with a mean interval between pulses of 4.2
<normalizedToken id="F07B63E81BAA3B8C052762230D699E90" originalValue="±">+/-</normalizedToken>
2.8 ms, emitted at a mean rate of 202.2
<normalizedToken id="1B2E4C0D39099E09032C4E852FFD5694" originalValue="±">+/-</normalizedToken>
119.7 pulses/second (Table
<tableCitation id="B194F9147B9871AAC0BBA31CBFE01051" captionStart="Table 4" captionStartId="T4" captionText="Table 4. Spectral and temporal measurements of the call of Pristimantis kayi sp. nov. Abbreviations are: DF = Dominant frequency; NH = Number of visible harmonics; 2 f 0 - 7 f 0 = Harmonic frequencies; CD = Call duration; IC Intervals between calls; CR = Call rate; NC = Notes / call; ND = Note duration; IN = Intervals between notes; NR = Note rate; PN = Pulses / Note; PN = Pulse duration; IP = Interval between pulses; PR = Pulse rate. The abbreviations used in the call types correspond to: CNS = Single note calls; CNP = Pulsed note calls; CCO = Complex calls. The abbreviations used in units of measurement correspond to: kHz = kilohertz; ms = milliseconds; s = seconds; / min = per minute; / s = per second." httpUri="http://table.plazi.org/id/CA5F83F326BA52DE8737B2C5B440DBA7" pageId="0" pageNumber="257" tableUuid="CA5F83F326BA52DE8737B2C5B440DBA7">4</tableCitation>
). The call of
<taxonomicName id="6BB6D6D5C5158E999E90EA9CA3B3D41C" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="04E53FFE680B6E1B333C67B29801528D" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. does not present stereotyped notes. Due to their structural elements (i.e. notes-pulses), notes can be classified into three different types: Single note calls (CNS; Fig.
<figureCitation id="1C57D7FC11331C2986B6BB0D3BB528CD" captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Call of Pristimantis kayi sp. nov. (adult male, MZUTI 852). The species has different types of calls, represented as follows: A single note call B pulsed note calls C complex calls (non-pulsed and pulsed notes)." figureDoi="10.3897/zookeys.1180.107333.figure18" httpUri="https://binary.pensoft.net/fig/912046" pageId="0" pageNumber="257">18A</figureCitation>
), Pulsed note calls (CNP; Fig.
<figureCitation id="DA8F7C6A98B33D26001113A24A2A5406" captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Call of Pristimantis kayi sp. nov. (adult male, MZUTI 852). The species has different types of calls, represented as follows: A single note call B pulsed note calls C complex calls (non-pulsed and pulsed notes)." figureDoi="10.3897/zookeys.1180.107333.figure18" httpUri="https://binary.pensoft.net/fig/912046" pageId="0" pageNumber="257">18B</figureCitation>
) and Complex calls (CNP; Fig.
<figureCitation id="86DD316657556681AE420DA895307E8A" captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Call of Pristimantis kayi sp. nov. (adult male, MZUTI 852). The species has different types of calls, represented as follows: A single note call B pulsed note calls C complex calls (non-pulsed and pulsed notes)." figureDoi="10.3897/zookeys.1180.107333.figure18" httpUri="https://binary.pensoft.net/fig/912046" pageId="0" pageNumber="257">18C</figureCitation>
), which contain single (non-pulsed) and pulsed notes in its structure. Temporal and spectral measurements of the different call types are shown in Table
<tableCitation id="4F73458A7166A86DF294045445CD8F63" captionStart="Table 4" captionStartId="T4" captionText="Table 4. Spectral and temporal measurements of the call of Pristimantis kayi sp. nov. Abbreviations are: DF = Dominant frequency; NH = Number of visible harmonics; 2 f 0 - 7 f 0 = Harmonic frequencies; CD = Call duration; IC Intervals between calls; CR = Call rate; NC = Notes / call; ND = Note duration; IN = Intervals between notes; NR = Note rate; PN = Pulses / Note; PN = Pulse duration; IP = Interval between pulses; PR = Pulse rate. The abbreviations used in the call types correspond to: CNS = Single note calls; CNP = Pulsed note calls; CCO = Complex calls. The abbreviations used in units of measurement correspond to: kHz = kilohertz; ms = milliseconds; s = seconds; / min = per minute; / s = per second." httpUri="http://table.plazi.org/id/CA5F83F326BA52DE8737B2C5B440DBA7" pageId="0" pageNumber="257" tableUuid="CA5F83F326BA52DE8737B2C5B440DBA7">4</tableCitation>
.
</paragraph>
<caption id="EC075F6F480D76F6B7A9F919757587AA" doi="10.3897/zookeys.1180.107333.figure18" httpUri="https://binary.pensoft.net/fig/912046" pageId="0" pageNumber="257" start="Figure 18" startId="F18">
<paragraph id="5B679EDE1097FC5BEE001E835D0C3B84" pageId="0" pageNumber="257">
<emphasis id="C69435FB7D415BB085C9ACBE745C6DCC" bold="true" pageId="0" pageNumber="257">Figure 18.</emphasis>
Call of
<taxonomicName id="2D3D8248DC19C7A37D3E61276EDB1130" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="2A9A501A4244913EDE60EEDF1DA6812D" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. (adult male, MZUTI 852). The species has different types of calls, represented as follows:
<emphasis id="C686E7D9EA169ADA54B0867C5D56F3C3" bold="true" pageId="0" pageNumber="257">A</emphasis>
single note call
<emphasis id="48DEC4F426CD881BEEFD9CA8958E54CC" bold="true" pageId="0" pageNumber="257">B</emphasis>
pulsed note calls
<emphasis id="68CE324EE805D27ACD07BC556F61F09E" bold="true" pageId="0" pageNumber="257">C</emphasis>
complex calls (non-pulsed and pulsed notes).
</paragraph>
</caption>
<caption id="CA5F83F326BA52DE8737B2C5B440DBA7" ID-Table-UUID="CA5F83F326BA52DE8737B2C5B440DBA7" httpUri="http://table.plazi.org/id/CA5F83F326BA52DE8737B2C5B440DBA7" pageId="0" pageNumber="257" start="Table 4" startId="T4">
<paragraph id="3BAE13F3B2BF2D27B4094FDCE4178FF1" pageId="0" pageNumber="257">
<emphasis id="52DAC37F26FE33DE6F013CBA0B490446" bold="true" pageId="0" pageNumber="257">Table 4.</emphasis>
Spectral and temporal measurements of the call of
<taxonomicName id="40E3F90128A93F9B573D2265EE352192" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="020CE1212893FB147B199199B3867E80" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. Abbreviations are: DF = Dominant frequency; NH = Number of visible harmonics; 2f0-7f0 = Harmonic frequencies; CD = Call duration; IC Intervals between calls; CR = Call rate; NC = Notes/call; ND = Note duration; IN = Intervals between notes; NR = Note rate; PN = Pulses/Note; PN = Pulse duration; IP = Interval between pulses; PR = Pulse rate. The abbreviations used in the call types correspond to: CNS = Single note calls; CNP = Pulsed note calls; CCO = Complex calls. The abbreviations used in units of measurement correspond to: kHz = kilohertz; ms = milliseconds; s = seconds; /min = per minute; /s = per second.
</paragraph>
</caption>
<paragraph id="46A7C06AB64289EF36FB44A14D475DB0" pageId="0" pageNumber="257">
<table id="88E70016BF129D2CBA22272B9D1D7781" pageId="0" pageNumber="257">
<tr id="F92A3FC5F42515C676DE8AF220CF3743" pageId="0" pageNumber="257">
<th id="1142B288091DD8CA00FC6B2D6332248D" colspan="1" pageId="0" pageNumber="257" rowspan="2">Parameters</th>
<th id="F32D41C64EA7BE3E296AEAACD064B63A" colspan="1" pageId="0" pageNumber="257" rowspan="1">Call</th>
<th id="440044559644D91039CE30D7D7092608" colspan="3" pageId="0" pageNumber="257" rowspan="1">Call Types</th>
</tr>
<tr id="BFAE593A71D4ABCE1BB2C35044B0BDBD" pageId="0" pageNumber="257">
<th id="537C05164216FFAF902F2B6073A416FE" colspan="1" pageId="0" pageNumber="257" rowspan="1">(general)</th>
<th id="FA85EBAA93326694BEC0F5BF94B16F39" colspan="1" pageId="0" pageNumber="257" rowspan="1">Type CNS</th>
<th id="2684A8AFA9CFA56B167BBA845D9B888F" colspan="1" pageId="0" pageNumber="257" rowspan="1">Type CNP</th>
<th id="D355AA953177D596D563EE831709F5F9" colspan="1" pageId="0" pageNumber="257" rowspan="1">Type CCO</th>
</tr>
<tr id="FFDF78F293428513A9887A70BF2C7E2F" pageId="0" pageNumber="257">
<td id="2B4F7FA89A1B7EA2C813777DB85D75F9" colspan="1" pageId="0" pageNumber="257" rowspan="1">CD (ms)</td>
<td id="A1EFCF10B0132C2A8693FC5045708FED" colspan="1" pageId="0" pageNumber="257" rowspan="1">
10-463 (165.13
<normalizedToken id="83201783B2B76C1A571F2B82380C3F4B" originalValue="±">+/-</normalizedToken>
130.06)
</td>
<td id="D8EA40B02E87B6FFFB16988DE797A4DA" colspan="1" pageId="0" pageNumber="257" rowspan="1">
10-213 (75.30
<normalizedToken id="B6711A6C149ACADFC10DDAA125E34B6C" originalValue="±">+/-</normalizedToken>
83.53)
</td>
<td id="D4D01748F12122E767E4F84A8BEB654A" colspan="1" pageId="0" pageNumber="257" rowspan="1">
40-463 (258.29
<normalizedToken id="5030FB8D2CECB5F5C76F4B682D1F1701" originalValue="±">+/-</normalizedToken>
105.37)
</td>
<td id="9AAD803F99EEA1F03FEF2C295911B9E4" colspan="1" pageId="0" pageNumber="257" rowspan="1">
91-189 (114.29
<normalizedToken id="65DD343B9CC9AF4FF1246E920EA9494B" originalValue="±">+/-</normalizedToken>
34.30)
</td>
</tr>
<tr id="F80053C0BB0C487B61433DF5D789D5A3" pageId="0" pageNumber="257">
<td id="81F2100368C11A0CBB110CEE219CA217" colspan="1" pageId="0" pageNumber="257" rowspan="1">IC (ms)</td>
<td id="8886539C8CE020BBF523E83DC1F83B32" colspan="1" pageId="0" pageNumber="257" rowspan="1">
528-32821 (4565.61
<normalizedToken id="62F89B822DFE2CA10471FF048AC2F0A7" originalValue="±">+/-</normalizedToken>
5742.25)
</td>
<td id="DEB409D1B2D7E26CD6CC011EEA70A558" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
<td id="C4A11341E9E410131BA099C4B66CC5A4" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
<td id="F55F746AC925BCB01C8FC9153F00E9AF" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
</tr>
<tr id="D5F8FAD39794C2D33434C6A860C0D93C" pageId="0" pageNumber="257">
<td id="07A0D8EC182B40CA0D5FF5B435B28198" colspan="1" pageId="0" pageNumber="257" rowspan="1">CR (/min)</td>
<td id="2B52EFE080B23CCB1B3A50D17006C794" colspan="1" pageId="0" pageNumber="257" rowspan="1">
1.82-112.78 (32.61
<normalizedToken id="5E8F7E8948447C91E41679536777DA2E" originalValue="±">+/-</normalizedToken>
23.23)
</td>
<td id="79415DF7AFFA93B352956BC0F476F269" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
<td id="EAD06F41CE864E41DC2BFBA04D6B6EE9" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
<td id="1A033D9322E459DBDFECB930D24A5FCC" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
</tr>
<tr id="A5A42436093C6B36B9AC7C1DA8908D01" pageId="0" pageNumber="257">
<td id="DD7BAAB5ABCE8285DFBEA1711699C94C" colspan="1" pageId="0" pageNumber="257" rowspan="1">NC</td>
<td id="B65DB1AB506B220184512A99FA3AD2A1" colspan="1" pageId="0" pageNumber="257" rowspan="1">
1-7 (2.62
<normalizedToken id="C30A3F81268346E6F3BA45A3F238A40A" originalValue="±">+/-</normalizedToken>
1.68)
</td>
<td id="89FA7B8CCD26C8E6829B85A7978606F8" colspan="1" pageId="0" pageNumber="257" rowspan="1">
1-2 (1.32
<normalizedToken id="348F93AB3AC91C690565EB2355556EB7" originalValue="±">+/-</normalizedToken>
0.48)
</td>
<td id="F2A3A2BBFCBBAE3F79319ABE5E6E049B" colspan="1" pageId="0" pageNumber="257" rowspan="1">
1-7 (3.76
<normalizedToken id="FECA403C8601C44DB1AB1F7B94A09EB0" originalValue="±">+/-</normalizedToken>
1.65)
</td>
<td id="27712D74D97C081E0B118DD0B8137219" colspan="1" pageId="0" pageNumber="257" rowspan="1">2</td>
</tr>
<tr id="9987B75B3BB0651321369F1666EDB76C" pageId="0" pageNumber="257">
<td id="375E17A616C0BF06AEC7B7EE873266F8" colspan="1" pageId="0" pageNumber="257" rowspan="1">ND (ms)</td>
<td id="A74D024A769CF7C0EA347D0AC7137270" colspan="1" pageId="0" pageNumber="257" rowspan="1">
10-111 (36.01
<normalizedToken id="F6F9A52786AB162CA4C4A7E7867D1538" originalValue="±">+/-</normalizedToken>
18.32)
</td>
<td id="FEBCE1966D2860D125D2321800B90CC3" colspan="1" pageId="0" pageNumber="257" rowspan="1">
10-21 (13.82
<normalizedToken id="AB777C1EC81A38FEDAC74CE1A916A44A" originalValue="±">+/-</normalizedToken>
2.76)
</td>
<td id="F7BE8D2295B3C99272C5564DFA2147AB" colspan="1" pageId="0" pageNumber="257" rowspan="1">
12-111 (39.85
<normalizedToken id="EE31F15009EF10A7FDAA9480458B46D1" originalValue="±">+/-</normalizedToken>
17.07)
</td>
<td id="6AF59206D2B04F7F0152A482B1381078" colspan="1" pageId="0" pageNumber="257" rowspan="1">
15-35 (21.40
<normalizedToken id="43C0A2A86DE47CE717D092FEED3BD3BF" originalValue="±">+/-</normalizedToken>
5.28)
</td>
</tr>
<tr id="E00077AD7041D61EC1E30CC244E12D25" pageId="0" pageNumber="257">
<td id="5ADBCBD2E4489D2A225450803DB53FED" colspan="1" pageId="0" pageNumber="257" rowspan="1">IN (ms)</td>
<td id="28EDC765FB09E4D2E35225D3B3B6A4D7" colspan="1" pageId="0" pageNumber="257" rowspan="1">
16-184 (43.81
<normalizedToken id="851221A31815CDD59637E40FFF9977A1" originalValue="±">+/-</normalizedToken>
36.18)
</td>
<td id="54A40D6C5DA1F11EE9F42971904731ED" colspan="1" pageId="0" pageNumber="257" rowspan="1">
134-184 (155.70
<normalizedToken id="0DD74AFC345F7B3A25A4CAEB911B3DC5" originalValue="±">+/-</normalizedToken>
17.45)
</td>
<td id="92F15C2905C406E0A1C0723CF60D8031" colspan="1" pageId="0" pageNumber="257" rowspan="1">
16-69 (31.94
<normalizedToken id="262726C56181BD614E4C8A3BBE36570F" originalValue="±">+/-</normalizedToken>
8.73)
</td>
<td id="F1E80808609F2DCBFBCB4CA8AAA06861" colspan="1" pageId="0" pageNumber="257" rowspan="1">
49-150 (70.86
<normalizedToken id="F20C0AA219121F41D1E040522F7B9A13" originalValue="±">+/-</normalizedToken>
36.13)
</td>
</tr>
<tr id="4C6CBF2898C49C7757B27F48382FCC33" pageId="0" pageNumber="257">
<td id="8AA56384BD71B48BAB927D445F6E9995" colspan="1" pageId="0" pageNumber="257" rowspan="1">NR (/s)</td>
<td id="0399C10343BB5E4E1034168A5C7F3315" colspan="1" pageId="0" pageNumber="257" rowspan="1">
5.13-35.71 (15.68
<normalizedToken id="15EA6749C988830BB7F7F75B5645ED1E" originalValue="±">+/-</normalizedToken>
6.05)
</td>
<td id="E8468401AC46643D6F1D88FE73AB0D8B" colspan="1" pageId="0" pageNumber="257" rowspan="1">
5.13-6.94 (6.14
<normalizedToken id="6AED017CFDCB5B87A545246EB38BE824" originalValue="±">+/-</normalizedToken>
0.69)
</td>
<td id="7CF41C17EEAB288F704750FD3514608C" colspan="1" pageId="0" pageNumber="257" rowspan="1">
10.42-35.71 (16.95
<normalizedToken id="92FE56191C240DD1921583A04CC301E0" originalValue="±">+/-</normalizedToken>
5.22)
</td>
<td id="09B2262BA9B1AFADED8ABAEAE8C53C68" colspan="1" pageId="0" pageNumber="257" rowspan="1">
5.75-14.93 (11.55
<normalizedToken id="F7E08FF3FC81884F34E81A954CF9088C" originalValue="±">+/-</normalizedToken>
3.08)
</td>
</tr>
<tr id="D72B900EFDF48406221E02A79EB83599" pageId="0" pageNumber="257">
<td id="4155E73F66A768D684B8C12D70BA1AF1" colspan="1" pageId="0" pageNumber="257" rowspan="1">PN</td>
<td id="E58E2B90DA862AB0BB1583A29E52D8F4" colspan="1" pageId="0" pageNumber="257" rowspan="1">
2-13 (5.70
<normalizedToken id="C576C59DD44A1CFB7E452E29B26C7BF6" originalValue="±">+/-</normalizedToken>
2.19)
</td>
<td id="363C0DBBFD281DF95E09AAA44D08C21B" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
<td id="4A04B9352F3E0499556C1BA2E1ADF104" colspan="1" pageId="0" pageNumber="257" rowspan="1">
2-13 (5.55
<normalizedToken id="BB532C52BC5E966453172324D36C9BE9" originalValue="±">+/-</normalizedToken>
2.30)
</td>
<td id="912427C7DA8525866B5FE183198D5EB2" colspan="1" pageId="0" pageNumber="257" rowspan="1">
2-9 (6
<normalizedToken id="461D8C9763E14FEE4B1395187745A324" originalValue="±">+/-</normalizedToken>
2.62)
</td>
</tr>
<tr id="378784AB6C5C5A0D56D394ED44A39474" pageId="0" pageNumber="257">
<td id="5E9D54531052B5D6E55E47E844409613" colspan="1" pageId="0" pageNumber="257" rowspan="1">PD (ms)</td>
<td id="061DED04622738231CA6BBC43BB1BE54" colspan="1" pageId="0" pageNumber="257" rowspan="1">
1-8 (2.95
<normalizedToken id="2D5C8C5C58EDD33E991469E93A63B45E" originalValue="±">+/-</normalizedToken>
1.07)
</td>
<td id="6F234BE6FCF8A5BE23FCDB5E32DCCCFB" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
<td id="015F794803B1449286FECF4544EA52D0" colspan="1" pageId="0" pageNumber="257" rowspan="1">
1-8 (3
<normalizedToken id="46BE51A7AE651DDC6B6F4B8CF70F7400" originalValue="±">+/-</normalizedToken>
1.07)
</td>
<td id="3E477E247CE8B2F1EA4784555760FD64" colspan="1" pageId="0" pageNumber="257" rowspan="1">
1-7 (2.47
<normalizedToken id="0BED980FAC673B75028ABCFFF66F26AB" originalValue="±">+/-</normalizedToken>
1.06)
</td>
</tr>
<tr id="1A7C642FA2BF7DE79B8128BDC051017E" pageId="0" pageNumber="257">
<td id="29D4D90215776B3E7E78F693B47E0882" colspan="1" pageId="0" pageNumber="257" rowspan="1">IP (ms)</td>
<td id="66C1D41735D1A500B8ED9E6ED096923B" colspan="1" pageId="0" pageNumber="257" rowspan="1">
0.6-13 (4.19
<normalizedToken id="A4789294A72552457466F29ABF9671A8" originalValue="±">+/-</normalizedToken>
2.80)
</td>
<td id="01FA0EB9CA5C15972747C79492E9B1C7" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
<td id="93A9342379594CE8147566CD253869A1" colspan="1" pageId="0" pageNumber="257" rowspan="1">
0.6-13 (4.32
<normalizedToken id="F181DAA00E25989193C4B11D74E087D1" originalValue="±">+/-</normalizedToken>
2.81)
</td>
<td id="3B7F6F5F8ED237F8E7861FDD6887B003" colspan="1" pageId="0" pageNumber="257" rowspan="1">
0.6-8 (2.02
<normalizedToken id="6D0F3C394429EB48BA1FBA36D1E9A29C" originalValue="±">+/-</normalizedToken>
1.71)
</td>
</tr>
<tr id="B96E63A0FD09CE0279DB812FCF075210" pageId="0" pageNumber="257">
<td id="4CF1AB41835E971490F9AC47328A8196" colspan="1" pageId="0" pageNumber="257" rowspan="1">PR (/s)</td>
<td id="351DB08A6E4F9984874CE8EE8012D646" colspan="1" pageId="0" pageNumber="257" rowspan="1">
66.77-769.23 (202.15
<normalizedToken id="3B5666AAD63FE72243AEB85706C92452" originalValue="±">+/-</normalizedToken>
119.72)
</td>
<td id="FFE112E97139CCE5877AB43FC36BF0B6" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
<td id="7334D252B0EFAA4DDE041800D7EEBA92" colspan="1" pageId="0" pageNumber="257" rowspan="1">
66.77-769.23 (190.05
<normalizedToken id="A044683436C176D365192F10DF76A59E" originalValue="±">+/-</normalizedToken>
108.22)
</td>
<td id="BF1A29DA345DD306CD5D79F2C5C774C8" colspan="1" pageId="0" pageNumber="257" rowspan="1">
69.93-769.23 (345.21
<normalizedToken id="2823628F3742C23AA299B057E06F7DFB" originalValue="±">+/-</normalizedToken>
157.84)
</td>
</tr>
<tr id="D3B759D696C3100A00B49E009D6D6196" pageId="0" pageNumber="257">
<td id="2AE2A102053B1DE2321849767523DDCC" colspan="1" pageId="0" pageNumber="257" rowspan="1">DF (kHz)</td>
<td id="10DD4B8747A0D1B5D8AFD0B106AE29A0" colspan="1" pageId="0" pageNumber="257" rowspan="1">
3-3.38 (3.19
<normalizedToken id="362F153D1AAD4B9898B27A64440A0BAE" originalValue="±">+/-</normalizedToken>
0.05)
</td>
<td id="7C400DDBD30A25BCBC2FDBE44D95F6E5" colspan="1" pageId="0" pageNumber="257" rowspan="1">
3-3.38 (3.21
<normalizedToken id="5858B20D3F13D12EF6BDAD0A0E1A1D4E" originalValue="±">+/-</normalizedToken>
0.07)
</td>
<td id="D12BD8CDAA60A45B1AD2DAEE709B0F95" colspan="1" pageId="0" pageNumber="257" rowspan="1">
3-3.38 (3.19
<normalizedToken id="B3A3EF390CF23AE283ED78C2BDB26417" originalValue="±">+/-</normalizedToken>
0.04)
</td>
<td id="8042F3471FCA7A87A766F360334C0CE8" colspan="1" pageId="0" pageNumber="257" rowspan="1">
3-3.38 (3.19
<normalizedToken id="18DD720C0FF174B5C78A04BE7F49C376" originalValue="±">+/-</normalizedToken>
0.07)
</td>
</tr>
<tr id="4E246D936007334A0541AF1626C12472" pageId="0" pageNumber="257">
<td id="3447F11C0ED9BF9090365521D3008608" colspan="1" pageId="0" pageNumber="257" rowspan="1">NH</td>
<td id="38758712221CF67E2ED3A1DF1C94A25D" colspan="1" pageId="0" pageNumber="257" rowspan="1">
2-6 (4.09
<normalizedToken id="D58C7BFEDB51983975844CC62382A794" originalValue="±">+/-</normalizedToken>
1.04)
</td>
<td id="4C512F60973D2C03A58286633C47F797" colspan="1" pageId="0" pageNumber="257" rowspan="1">
3-7 (5.32
<normalizedToken id="215A3B9BA29F305310B0E2B3894E49D3" originalValue="±">+/-</normalizedToken>
0.91)
</td>
<td id="42A408CA9518D4726154B2AFB915AE51" colspan="1" pageId="0" pageNumber="257" rowspan="1">
3-6 (4.02
<normalizedToken id="1C5C41C93D5EB3DD4C3152C39C43EF7F" originalValue="±">+/-</normalizedToken>
1.01)
</td>
<td id="2C565A420A846D3EF998F150F6CA66D2" colspan="1" pageId="0" pageNumber="257" rowspan="1">
3-6 (4.07
<normalizedToken id="AEFF631FA2B08D1F95F6E5258248BC1D" originalValue="±">+/-</normalizedToken>
1.08)
</td>
</tr>
<tr id="FB4B04B0DFFF0C3BB77FC65915D91357" pageId="0" pageNumber="257">
<td id="1D9BC3C2615107F74B6E56DD1361A1CD" colspan="1" pageId="0" pageNumber="257" rowspan="1">2f0 (kHz)</td>
<td id="DFCC2B8E9A1A250077C3595552758456" colspan="1" pageId="0" pageNumber="257" rowspan="1">
6-6.75 (6.38
<normalizedToken id="4072BC6EC841250FDC489A6915FDA624" originalValue="±">+/-</normalizedToken>
0.09)
</td>
<td id="C90C800AFE8B953FE12AB29DFA10DD68" colspan="1" pageId="0" pageNumber="257" rowspan="1">
6-6.75 (6.46
<normalizedToken id="24D3A8BD35F472D910CF8BC755827A13" originalValue="±">+/-</normalizedToken>
0.16)
</td>
<td id="D0925AA0DFDBDFB964CA6E6FDD533176" colspan="1" pageId="0" pageNumber="257" rowspan="1">
6-6.75 (6.38
<normalizedToken id="D88B36388E168132238E4B57FF709D16" originalValue="±">+/-</normalizedToken>
0.08)
</td>
<td id="3F9E9180DA2C784D4CE7F587F5EF055F" colspan="1" pageId="0" pageNumber="257" rowspan="1">
6-6.75 (6.39
<normalizedToken id="9A840EF4C012345430B57BB332406DC6" originalValue="±">+/-</normalizedToken>
0.11)
</td>
</tr>
<tr id="4FD0FB579FE168527CC0BEE1D2E4BE1B" pageId="0" pageNumber="257">
<td id="19B2798E44B9EA2634C43CACBFEB6B6B" colspan="1" pageId="0" pageNumber="257" rowspan="1">3f0 (kHz)</td>
<td id="33C92A5A71194312606670EBD43AE46F" colspan="1" pageId="0" pageNumber="257" rowspan="1">
9-10.50 (9.56
<normalizedToken id="BB38D97999F55CC5645599A48467B6A7" originalValue="±">+/-</normalizedToken>
0.13)
</td>
<td id="767CC313B45C7814773368E78976C279" colspan="1" pageId="0" pageNumber="257" rowspan="1">
9-10.13 (9.64
<normalizedToken id="02BECC6BF339CC36A747DDCAB5DFA375" originalValue="±">+/-</normalizedToken>
0.22)
</td>
<td id="B59ADABA8315B367DAE3D9D8F6045380" colspan="1" pageId="0" pageNumber="257" rowspan="1">
9-10.50 (9.55
<normalizedToken id="E4528FD6B99BB32480DD2F5FCDF20F69" originalValue="±">+/-</normalizedToken>
0.12)
</td>
<td id="E46D5B964CA351F544903C7C481192F4" colspan="1" pageId="0" pageNumber="257" rowspan="1">
9-10.13 (9.56
<normalizedToken id="D2028DF0BF020707B92B5C8F326ACFFA" originalValue="±">+/-</normalizedToken>
0.18)
</td>
</tr>
<tr id="05836A56797498F343581077A8F294F6" pageId="0" pageNumber="257">
<td id="7BA8D02211DE055906F863B142C6951F" colspan="1" pageId="0" pageNumber="257" rowspan="1">4f0 (kHz)</td>
<td id="CDBF5C794376FFFEB13F5512BC76075A" colspan="1" pageId="0" pageNumber="257" rowspan="1">
12-13.50 (12.75
<normalizedToken id="E777844545B6122D2FFA2FFE078CD4CD" originalValue="±">+/-</normalizedToken>
0.17)
</td>
<td id="B36FC2DDF92C827B31620D5422779E8B" colspan="1" pageId="0" pageNumber="257" rowspan="1">
12-13.50 (12.81
<normalizedToken id="4F087DB5EE6C979F0CB78671F28392D3" originalValue="±">+/-</normalizedToken>
0.27)
</td>
<td id="4CD32FA040BBE2303C8593C8DCF039F4" colspan="1" pageId="0" pageNumber="257" rowspan="1">
12-13.50 (12.74
<normalizedToken id="388B5CB6D9DD26AAE3F93AC32A1435CD" originalValue="±">+/-</normalizedToken>
0.15)
</td>
<td id="BB9F2E4E461E88034F07F83A06210F9E" colspan="1" pageId="0" pageNumber="257" rowspan="1">
12-13.50 (12.80
<normalizedToken id="F59BC4B3860352DBAF5A99899BC080B3" originalValue="±">+/-</normalizedToken>
0.24)
</td>
</tr>
<tr id="F06E83884417D4AAC27186692E3A3737" pageId="0" pageNumber="257">
<td id="632B4B5CE708D2ACEC021AAE5BF5EE26" colspan="1" pageId="0" pageNumber="257" rowspan="1">5f0 (kHz)</td>
<td id="99C2045A4598A064E8473AC4E39B8AC6" colspan="1" pageId="0" pageNumber="257" rowspan="1">
15-16.88 (15.96
<normalizedToken id="D33980812D1F86997CD11118CDBD1AAE" originalValue="±">+/-</normalizedToken>
0.24)
</td>
<td id="D4972EFCC0F4ADAB303648DB805D1A0E" colspan="1" pageId="0" pageNumber="257" rowspan="1">
15.75-16.88 (16.03
<normalizedToken id="95057B4E33260FAC8AC8FDD1105E43B3" originalValue="±">+/-</normalizedToken>
0.27)
</td>
<td id="F466D01BD4A501F154B40260F6171010" colspan="1" pageId="0" pageNumber="257" rowspan="1">
15-16.88 (15.94
<normalizedToken id="0A7D00777C41D6471E8FC5DB83D490D9" originalValue="±">+/-</normalizedToken>
0.21)
</td>
<td id="B18474951C39C0FB45837F1F63E5491C" colspan="1" pageId="0" pageNumber="257" rowspan="1">
15-16.88 (16.04
<normalizedToken id="12F23E265D73E87EC2F2F2C571635E0F" originalValue="±">+/-</normalizedToken>
0.39)
</td>
</tr>
<tr id="88BD4B4A9B4C0D8658FF10B35A2ED673" pageId="0" pageNumber="257">
<td id="562F8A1CA2B03A75E22DB55FB903B6C8" colspan="1" pageId="0" pageNumber="257" rowspan="1">6f0 (kHz)</td>
<td id="72E199AD83A9F46482F54CCA55F6A807" colspan="1" pageId="0" pageNumber="257" rowspan="1">
18-20.25 (19.16
<normalizedToken id="892CF5B2369CA536C50F9A9D797217C0" originalValue="±">+/-</normalizedToken>
0.23)
</td>
<td id="4E0372DDA097AEB8909660416AC8B526" colspan="1" pageId="0" pageNumber="257" rowspan="1">
19.13-20.25 (19.25
<normalizedToken id="FBE3DC19AB5BDAEAB5B8358622195C7E" originalValue="±">+/-</normalizedToken>
0.31)
</td>
<td id="E70C306150782887D0A61B63D89BFFE8" colspan="1" pageId="0" pageNumber="257" rowspan="1">
18-19.13 (19.13
<normalizedToken id="CC896C11DE9E9FE56DF1EFDB640A3BE0" originalValue="±">+/-</normalizedToken>
0.18)
</td>
<td id="422315FA79124EEE8D36056A5D738BE1" colspan="1" pageId="0" pageNumber="257" rowspan="1">
18.94-20.25 (18.13
<normalizedToken id="4A1AD2830ACF02C49565A463EDF98513" originalValue="±">+/-</normalizedToken>
0.08)
</td>
</tr>
<tr id="EC8D360AB9C81BB87EDE267CF4477D22" pageId="0" pageNumber="257">
<td id="EB046DA37F4F506153D7879FB9E75253" colspan="1" pageId="0" pageNumber="257" rowspan="1">7f0 (kHz)</td>
<td id="F83EA64A5ED616E7AB616755DB6A1F2F" colspan="1" pageId="0" pageNumber="257" rowspan="1">22.31</td>
<td id="CDFF2BFFA553A2E335160FE84506B1A4" colspan="1" pageId="0" pageNumber="257" rowspan="1">22.31</td>
<td id="4F7B6D5B1E05E6E143F0F7ED0266D902" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
<td id="CFEEAAC1B16553078D25B89D83F2BC08" colspan="1" pageId="0" pageNumber="257" rowspan="1">-</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection id="5CA2F0EEA4035B55FB1070E646A047AF" pageId="0" pageNumber="257" type="distribution">
<paragraph id="E5CE3A81EB22353974D85F70AE2F460A" pageId="0" pageNumber="257">Distribution and natural history observations.</paragraph>
<paragraph id="5E11BB2F4306D52F3B91D8EA3BF7999F" pageId="0" pageNumber="257">
<taxonomicName id="18B23D04633D614653E16A7A019CF35F" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="6C6026BC7135B69E05EF4362AEA86EF3" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. has been recorded from the Cordillera de Guacamayos,
<normalizedToken id="A1D42D744EF0F361F1D8ED25F8E72CD1" originalValue="Volcán">Volcan</normalizedToken>
Sumaco (Napo Province) and Cerro Mayordomo, Machay Reserve, Naturetrek Vizcaya Reserve, Leito Reserve (Tungurahua Province) at an elevational range of 2190-3600 m a.s.l. (Fig.
<figureCitation id="6C641A1876FA5D60388854C41574B40E" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Distribution of four closely-related species of the Pristimantis myersi species group in the Andes of Ecuador." figureDoi="10.3897/zookeys.1180.107333.figure11" httpUri="https://binary.pensoft.net/fig/912039" pageId="0" pageNumber="257">11</figureCitation>
). Calling males were heard during the day and night. During the night, frogs of
<taxonomicName id="512708AE3797C64B3A903FE7806226F9" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="3D8AC496D025DD8D49E5DFA32BFB9305" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
sp. nov. were found on the forest floor or perched on low vegetation (always below 60 cm from ground level), including shrubs, grasses (
<taxonomicName id="338FA3B8F709BB03F36D062CAE0AC12F" class="Liliopsida" family="Poaceae" genus="Neurolepis" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Neurolepis" order="Cyperales" pageId="0" pageNumber="257" phylum="Magnoliophyta" rank="genus">
<emphasis id="B258008C228583F81F9C354F6B7DC36F" italics="true" pageId="0" pageNumber="257">Neurolepis</emphasis>
</taxonomicName>
sp.), ferns and
<taxonomicName id="9ECF673AB30C91EA258E86F1D2E1A7AA" class="Lycopodiopsida" family="Selaginellaceae" genus="Selaginella" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Selaginella" order="Selaginellales" pageId="0" pageNumber="257" phylum="Lycophyta" rank="genus">
<emphasis id="75950F9B749737A27FD81895B0283AE7" italics="true" pageId="0" pageNumber="257">Selaginella</emphasis>
</taxonomicName>
. The species has been found in primary and disturbed Andean forests, with trees of the genus
<taxonomicName id="C8B4544430BC8B04FB8C994D5CFE94C0" class="Magnoliopsida" family="Cunoniaceae" genus="Weinmannia" higherTaxonomySource="GBIF,IPNI" kingdom="Plantae" lsidName="Weinmannia" order="Oxalidales" pageId="0" pageNumber="257" phylum="Magnoliophyta" rank="genus">
<emphasis id="DD887FDDE4FFBDBB4DDFFD717757373E" italics="true" pageId="0" pageNumber="257">Weinmannia</emphasis>
</taxonomicName>
,
<taxonomicName id="9634B146EA861094F085E7A013E8D1B1" class="Insecta" family="Clusiidae" genus="Clusia" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Clusia" order="Diptera" pageId="0" pageNumber="257" phylum="Arthropoda" rank="genus">
<emphasis id="62958C047A9B8AD60DF28385E7957F07" italics="true" pageId="0" pageNumber="257">Clusia</emphasis>
</taxonomicName>
,
<taxonomicName id="5ADC1B7AB091C38CDA5D16343A4AE154" class="Dicotyledoneae" family="Betulaceae" genus="Alnus" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Alnus" order="Fagales" pageId="0" pageNumber="257" phylum="Angiospermae" rank="genus">
<emphasis id="A5BED34E815E9C6C5EBFD66C1DA5D772" italics="true" pageId="0" pageNumber="257">Alnus</emphasis>
</taxonomicName>
and others, of 10-15 metres (Fig.
<figureCitation id="0F9AFE4030D8626E00D566B9D78E3475" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Habitat vegetation at type localities and etymology of the two new frogs described herein. Photographs by JPRP and Zane Libke A panoramic view of the upper Pastaza Valley and its adjacent mountains, main geographic barriers promoting speciation at a regional scale, at back Tungurahua Volcano and mountains south of the Pastaza River habitat of Pristimantis donnelsoni sp. nov., image from the slopes of Cerro Mayordomo type locality of Pristimantis kayi sp. nov., in the northern mountains to the Pastaza River B Tungurahua Volcano eruption, from Chamana Reserve C Montane forest in Candelaria Reserve D Don Nelson Palacios (+) during an expedition; we tribute his work with the naming of Pristimantis donnelsoni sp. nov. E Sumaco Volcano seen from Wawa Sumaku, habitat of P. kayi sp. nov. F Montane forest and subparamo in Leito, Patate G Andreas Kay (+); we honour his devotion to nature photography and research in the upper Pastaza Valley and through Ecuador by naming Pristimantis kayi sp. nov." figureDoi="10.3897/zookeys.1180.107333.figure12" httpUri="https://binary.pensoft.net/fig/912040" pageId="0" pageNumber="257">12</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="DE33213E95E4490F83427838C3833A6D" pageId="0" pageNumber="257" type="etymology">
<paragraph id="7DE8B1D2ADBCA7661AE6DBFFDBD02162" pageId="0" pageNumber="257">Etymology.</paragraph>
<paragraph id="66AD52C0ECA308B9388DD309B23C322F" pageId="0" pageNumber="257">
The specific epithet &quot;
<taxonomicName id="062B83110E86A59350AFA33313013787" authorityName="Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig" authorityYear="2023" class="Amphibia" family="Strabomantidae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis kayi" order="Anura" pageId="0" pageNumber="257" phylum="Chordata" rank="species" species="kayi">
<emphasis id="A61F65C2A2F76360E365F4764C5D2572" italics="true" pageId="0" pageNumber="257">Pristimantis kayi</emphasis>
</taxonomicName>
&quot; is a noun in the genitive case and a patronym for Andreas Kay, a German physicist, biologist and friend who spent much of his life documenting and exploring biodiversity, contributing substantially to the conservation of Ecuadorian forests (Fig.
<figureCitation id="62B485E0253E19B561E6A86DFB563325" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Habitat vegetation at type localities and etymology of the two new frogs described herein. Photographs by JPRP and Zane Libke A panoramic view of the upper Pastaza Valley and its adjacent mountains, main geographic barriers promoting speciation at a regional scale, at back Tungurahua Volcano and mountains south of the Pastaza River habitat of Pristimantis donnelsoni sp. nov., image from the slopes of Cerro Mayordomo type locality of Pristimantis kayi sp. nov., in the northern mountains to the Pastaza River B Tungurahua Volcano eruption, from Chamana Reserve C Montane forest in Candelaria Reserve D Don Nelson Palacios (+) during an expedition; we tribute his work with the naming of Pristimantis donnelsoni sp. nov. E Sumaco Volcano seen from Wawa Sumaku, habitat of P. kayi sp. nov. F Montane forest and subparamo in Leito, Patate G Andreas Kay (+); we honour his devotion to nature photography and research in the upper Pastaza Valley and through Ecuador by naming Pristimantis kayi sp. nov." figureDoi="10.3897/zookeys.1180.107333.figure12" httpUri="https://binary.pensoft.net/fig/912040" pageId="0" pageNumber="257">12</figureCitation>
). Andreas was part of the team that created
<normalizedToken id="D38A9B62B4EEC2C0ECFB512A2A13355E" originalValue="Fundación">Fundacion</normalizedToken>
<normalizedToken id="55BB3ECED3B5B6071E90C221D6C66D02" originalValue="EcoMinga’s">EcoMinga's</normalizedToken>
Dracula Reserva in north-western Ecuador. Additionally, his work shed light on understanding the diversity and endemism patterns of orchids in the Llanganates-Sangay Ecological Corridor, where he passed away in 2019. Some of his amazing photographs can be seen here: https://www.flickr.com/photos/andreaskay/albums/.
</paragraph>
</subSubSection>
</treatment>
</subSection>
</document>
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