treatments-xml/data/5D/43/87/5D4387F0C01A5522BCE2FC1CBF59FE44.xml

<document id="A923656FB891849A58FDA8EC7D120BF8" ID-DOI="10.5281/zenodo.197710" ID-GBIF-Dataset="1a01882e-4146-4bd1-a66f-28fe965e715d" ID-ISSN="1175-5326" ID-Zenodo-Dep="197710" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460275878432" checkinUser="plazi" docAuthor="Ron, Santiago R., Toral, Eduardo, Rivera, Myrian &amp; Terán-Valdez, Andrea" docDate="2010" docId="5D4387F0C01A5522BCE2FC1CBF59FE44" docLanguage="en" docName="zt02606p049.pdf" docOrigin="Zootaxa 2606" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Engystomops puyango Ron, Toral, Rivera &amp; Terán-Valdez, 2010, sp. nov." docType="treatment" docVersion="8" lastPageNumber="43" masterDocId="A17AFF88C0195530BC75FFBEBC48FFD6" masterDocTitle="A new species of Engystomops (Anura: Leiuperidae) from southwestern Ecuador" masterLastPageNumber="49" masterPageNumber="25" pageNumber="28" updateTime="1698248398553" updateUser="plazi">
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<mods:title id="BFD409174E9AC0E29DEEB40F912B1CEB">A new species of Engystomops (Anura: Leiuperidae) from southwestern Ecuador</mods:title>
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<mods:namePart id="6C45C3B1A65066CFBD4283BE6E966867">Ron, Santiago R.</mods:namePart>
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<mods:namePart id="6BE4E02674FA5D14763FC0DF4C277291">Toral, Eduardo</mods:namePart>
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<mods:namePart id="B90150B8A68CD500A5226196DB56D701">Terán-Valdez, Andrea</mods:namePart>
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<paragraph id="D55536E6C01A5533BCE2FC1CBE44FC6A" blockId="3.[151,524,930,956]" box="[151,524,930,956]" pageId="3" pageNumber="28">
<heading id="8E1D818AC01A5533BCE2FC1CBE44FC6A" bold="true" box="[151,524,930,956]" fontSize="11" level="1" pageId="3" pageNumber="28" reason="1">
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<taxonomicName id="12EA4D65C01A5533BCE2FC1CBDEFFC6A" box="[151,423,930,956]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="3" pageNumber="28" phylum="Chordata" rank="species" species="puyango" status="sp. nov.">
<emphasis id="E79EEAF4C01A5533BCE2FC1CBDEFFC6A" bold="true" box="[151,423,930,956]" italics="true" pageId="3" pageNumber="28">Engystomops puyango</emphasis>
</taxonomicName>
<taxonomicNameLabel id="FCAD578FC01A5533BDDAFC1CBE44FC6A" box="[431,524,930,956]" pageId="3" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
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<subSubSection id="9DF0656DC01A5533BCE2FC54BE4CF9D9" pageId="3" pageNumber="28" type="materials_examined">
<paragraph id="D55536E6C01A5533BCE2FC54B9CAFB99" blockId="3.[151,1438,1002,2036]" pageId="3" pageNumber="28">
<emphasis id="E79EEAF4C01A5533BCE2FC54BD58FBD2" bold="true" box="[151,272,1002,1028]" pageId="3" pageNumber="28">
<typeStatus id="0A518844C01A5533BCE2FC54BD43FBD2" box="[151,267,1002,1028]" pageId="3" pageNumber="28" type="holotype">Holotype</typeStatus>
.
</emphasis>
(
<figureCitation id="4DD12A63C01A5533BD55FC54BD2BFBD2" box="[288,355,1002,1028]" captionStart="FIGURE 1" captionStartId="4.[151,258,1882,1906]" captionTargetBox="[153,1435,217,1850]" captionTargetId="figure@4.[151,1436,194,1861]" captionTargetPageId="4" captionText="FIGURE 1. Lateral (top), ventral (left), and dorsal (right) views of the holotype of Engystomops puyango (QCAZ 26978). Not drawn to scale." httpUri="https://zenodo.org/record/197711/files/figure.png" pageId="3" pageNumber="28">Fig. 1</figureCitation>
) 
<collectionCode id="B3FBAE23C01A5533BD00FC54BD8DFBD2" box="[373,453,1002,1028]" httpUri="http://grbio.org/cool/kwpg-nhbg" name="Museo de Zoologia, Pontificia Universidad Catolica del Ecuador" pageId="3" pageNumber="28">QCAZ</collectionCode>
26978 (field no. 
<collectionCode id="B3FBAE23C01A5533BEFBFC54BE91FBD2" box="[654,729,1002,1028]" pageId="3" pageNumber="28">PUCE</collectionCode>
14586), adult male from 
<collectingCountry id="ADFD7676C01A5533B877FC54B82DFBD2" box="[1026,1125,1002,1028]" name="Ecuador" pageId="3" pageNumber="28">Ecuador</collectingCountry>
, Provincia El Oro, Bosque Protector 
<taxonomicName id="12EA4D65C01A5533BD67FBAEBDC9FBFC" box="[274,385,1040,1066]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="3" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
, near the bridge over the 
<taxonomicName id="12EA4D65C01A5533BEB7FBAEBF66FBFC" box="[706,814,1040,1066]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="3" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
River on the road Alamor-Arenillas (
<geoCoordinate id="B0DE5021C01A5533B972FBAEB9DEFBFC" box="[1287,1430,1040,1066]" direction="south" orientation="latitude" pageId="3" pageNumber="28" precision="1" value="-3.87159">3.87159° S</geoCoordinate>
, 
<geoCoordinate id="B0DE5021C01A5533BCE2FB8BBD7FFB99" box="[151,311,1077,1103]" direction="west" orientation="longitude" pageId="3" pageNumber="28" precision="1" value="-80.0377">80.03770° W</geoCoordinate>
), 
<quantity id="12129B03C01A5533BD39FB8BBDDEFB99" box="[332,406,1077,1103]" metricMagnitude="2" metricUnit="m" metricValue="3.21" pageId="3" pageNumber="28" unit="m" value="321.0">321 m</quantity>
above sea level, collected by S. R. Ron, E. E. Tapia, I. G. Tapia on 
<date id="A1541026C01A5533B8DEFB8BB936FB99" box="[1195,1406,1077,1103]" pageId="3" pageNumber="28" value="2004-02-19">19 February 2004</date>
.
</paragraph>
<paragraph id="D55536E6C01A5533BCB0FBE4B83CFADC" blockId="3.[151,1438,1002,2036]" pageId="3" pageNumber="28">
<emphasis id="E79EEAF4C01A5533BCB0FBE4BDCEFBA2" bold="true" box="[197,390,1114,1140]" pageId="3" pageNumber="28">Paratopotypes.</emphasis>
<collectionCode id="B3FBAE23C01A5533BDFBFBE4BD97FBA2" box="[398,479,1114,1140]" httpUri="http://grbio.org/cool/kwpg-nhbg" name="Museo de Zoologia, Pontificia Universidad Catolica del Ecuador" pageId="3" pageNumber="28">QCAZ</collectionCode>
26983–84, 26986, 26991, 26996, 26999, 27000–01, 27004, 27006, 27008–14, 27016, adult males, 26987–88, 26990, 27005, adult females, collected by S. R. Ron, I. G. Tapia, and E. E. Tapia on 
<date id="A1541026C01A5533BD76FB1BBD9FFB69" box="[259,471,1189,1215]" pageId="3" pageNumber="28" value="2004-02-19">19 February 2004</date>
; 
<collectionCode id="B3FBAE23C01A5533BD96FB1BBE7BFB69" box="[483,563,1189,1215]" httpUri="http://grbio.org/cool/kwpg-nhbg" name="Museo de Zoologia, Pontificia Universidad Catolica del Ecuador" pageId="3" pageNumber="28">QCAZ</collectionCode>
26969, 26971–72, 26974–75, 26977–78, adult males, 26967, adult female collected by G. Romero, E. E. Tapia, and S. R. Ron on 
<date id="A1541026C01A5533BF66FB74BF80FB32" box="[787,968,1226,1252]" pageId="3" pageNumber="28" value="2005-01-07">7 January 2005</date>
; 28717, adult male, 28716, adult female collected by C. Proaño, M. Guerra and S. R. Ron between 
<date id="A1541026C01A5533BF30FB4EB827FADC" box="[837,1135,1264,1290]" pageId="3" pageNumber="28" value="2005-02-13" valueMax="2005-02-20" valueMin="2005-02-13">13 and 20 February 2005</date>
.
</paragraph>
<paragraph id="D55536E6C01A5533BCB0FAABBE4CF9D9" blockId="3.[151,1438,1002,2036]" pageId="3" pageNumber="28">
<emphasis id="E79EEAF4C01A5533BCB0FAABBD1DFAF9" bold="true" box="[197,341,1301,1327]" pageId="3" pageNumber="28">
<typeStatus id="0A518844C01A5533BCB0FAABBD18FAF9" box="[197,336,1301,1327]" pageId="3" pageNumber="28" type="paratype">Paratypes</typeStatus>
.
</emphasis>
Provincia Loja: 
<quantity id="12129B03C01A5533BE5BFAABBE36FAF9" box="[558,638,1301,1327]" metricMagnitude="4" metricUnit="m" metricValue="1.0" pageId="3" pageNumber="28" unit="km" value="10.0">10 km</quantity>
north from Zapotillo along the road to Arenillas (
<geoCoordinate id="B0DE5021C01A5533B971FAABB9DFFAF9" box="[1284,1431,1301,1327]" direction="south" orientation="latitude" pageId="3" pageNumber="28" precision="1" value="-4.31198">4.31198° S</geoCoordinate>
, 
<geoCoordinate id="B0DE5021C01A5533BCE2FA84BD73FA82" box="[151,315,1338,1364]" direction="west" orientation="longitude" pageId="3" pageNumber="28" precision="1" value="-80.2169">80.21690° W</geoCoordinate>
), 
<quantity id="12129B03C01A5533BD24FA84BDD5FA82" box="[337,413,1338,1364]" metricMagnitude="2" metricUnit="m" metricValue="2.31" pageId="3" pageNumber="28" unit="m" value="231.0">231 m</quantity>
above sea level, 
<collectionCode id="B3FBAE23C01A5533BE1AFA84BE88FA82" box="[623,704,1338,1364]" httpUri="http://grbio.org/cool/kwpg-nhbg" name="Museo de Zoologia, Pontificia Universidad Catolica del Ecuador" pageId="3" pageNumber="28">QCAZ</collectionCode>
26959, adult male, collected by S. R. Ron, I. G. Tapia, E. E. Tapia on 
<date id="A1541026C01A5533BD72FADEBDA8FAAC" box="[263,480,1376,1402]" pageId="3" pageNumber="28" value="2004-02-18">18 February 2004</date>
; old road Alamor-Puyango (
<geoCoordinate id="B0DE5021C01A5533BF42FADEBF88FAAC" box="[823,960,1376,1402]" direction="south" orientation="latitude" pageId="3" pageNumber="28" precision="1" value="-3.93548">3.93548° S</geoCoordinate>
, 
<geoCoordinate id="B0DE5021C01A5533BFBBFADEB83AFAAC" box="[974,1138,1376,1402]" direction="west" orientation="longitude" pageId="3" pageNumber="28" precision="1" value="-80.10865">80.10865° W</geoCoordinate>
), 
<quantity id="12129B03C01A5533B8FDFADEB89CFAAC" box="[1160,1236,1376,1402]" metricMagnitude="2" metricUnit="m" metricValue="6.5" pageId="3" pageNumber="28" unit="m" value="650.0">650 m</quantity>
above sea level, 
<collectionCode id="B3FBAE23C01A5533BCE2FA3BBCAFFA49" box="[151,231,1413,1439]" httpUri="http://grbio.org/cool/kwpg-nhbg" name="Museo de Zoologia, Pontificia Universidad Catolica del Ecuador" pageId="3" pageNumber="28">QCAZ</collectionCode>
28758–62, adult males, collected by C. Proaño, M. Guerra and S. R. Ron between 
<date id="A1541026C01A5533B8C2FA3BBC92FA12" pageId="3" pageNumber="28" value="2005-02-20" valueMax="2005-02-23" valueMin="2005-02-20">20 and 23 February 2005</date>
; Catamayo (
<geoCoordinate id="B0DE5021C01A5533BD02FA14BE41FA12" box="[375,521,1450,1476]" direction="south" orientation="latitude" pageId="3" pageNumber="28" precision="1" value="-3.97004">3.97004° S</geoCoordinate>
, 
<geoCoordinate id="B0DE5021C01A5533BE6DFA14BE8CFA12" box="[536,708,1450,1476]" direction="west" orientation="longitude" pageId="3" pageNumber="28" precision="1" value="-79.36876">79.36876° W</geoCoordinate>
), 
<quantity id="12129B03C01A5533BEA9FA14BF73FA12" box="[732,827,1450,1476]" metricMagnitude="3" metricUnit="m" metricValue="1.291" pageId="3" pageNumber="28" unit="m" value="1291.0">1291 m</quantity>
above sea level, 
<collectionCode id="B3FBAE23C01A5533B86DFA14B824FA12" box="[1048,1132,1450,1476]" httpUri="http://grbio.org/cool/kwpg-nhbg" name="Museo de Zoologia, Pontificia Universidad Catolica del Ecuador" pageId="3" pageNumber="28">QCAZ</collectionCode>
31506–08, adult males, collected by I. G. Tapia and G. Onore on 
<date id="A1541026C01A5533BEFDFA6EBF3AFA3C" box="[648,882,1488,1514]" pageId="3" pageNumber="28" value="2005-12-30">30 December 2005</date>
; Mangaurquillo, 
<collectionCode id="B3FBAE23C01A5533B834FA6EB8DAFA3C" box="[1089,1170,1488,1514]" pageId="3" pageNumber="28">FHGO</collectionCode>
3372, 3374–76, adult males, collected by F. Nogales.
</paragraph>
</subSubSection>
<subSubSection id="9DF0656DC01A5536BCB0F9A4BE93FAAE" lastPageId="6" lastPageNumber="31" pageId="3" pageNumber="28" type="diagnosis">
<paragraph id="D55536E6C01A5533BCB0F9A4BE8FF91C" blockId="3.[151,1438,1002,2036]" pageId="3" pageNumber="28">
<emphasis id="E79EEAF4C01A5533BCB0F9A4BD0FF9E2" bold="true" box="[197,327,1562,1588]" pageId="3" pageNumber="28">Diagnosis.</emphasis>
A member of 
<taxonomicName id="12EA4D65C01A5533BD8DF9A5BEDFF9E2" box="[504,663,1563,1588]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="3" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C01A5533BD8DF9A5BEDFF9E2" box="[504,663,1563,1588]" italics="true" pageId="3" pageNumber="28">Engystomops</emphasis>
</taxonomicName>
, clade Duovox. The assignment to 
<taxonomicName id="12EA4D65C01A5533B837F9A5B8A9F9E2" box="[1090,1249,1563,1588]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="3" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C01A5533B837F9A5B8A9F9E2" box="[1090,1249,1563,1588]" italics="true" pageId="3" pageNumber="28">Engystomops</emphasis>
</taxonomicName>
is based on the molecular phylogeny published by 
<bibRefCitation id="B17B4B17C01A5533BE26F9FEBF64F98C" box="[595,812,1600,1626]" pageId="3" pageNumber="28" refString="Ron, S. R., Santos, J. C. &amp; Cannatella, D. C. (2006) Phylogeny of the tungara frog genus Engystomops (= Physalaemus pustulosus species group; Anura; Leptodactylidae). Molecular Phylogenetics and Evolution, 39, 392 - 403." type="journal article">
Ron 
<emphasis id="E79EEAF4C01A5533BEFAF9FEBE8FF98F" box="[655,711,1600,1625]" italics="true" pageId="3" pageNumber="28">et al</emphasis>
. (2006)
</bibRefCitation>
which shows high support for the inclusion of 
<taxonomicName id="12EA4D65C01A5533B9F7F9FEBD4FF9A8" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="3" pageNumber="28" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01A5533B9F7F9FEBD4FF9A8" italics="true" pageId="3" pageNumber="28">E. puyango</emphasis>
</taxonomicName>
(referred as “ 
<taxonomicName id="12EA4D65C01A5533BDBDF9DBBE3FF9A8" box="[456,631,1637,1662]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="3" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C01A5533BDBDF9DBBE3FF9A8" box="[456,631,1637,1662]" italics="true" pageId="3" pageNumber="28">Engystomops</emphasis>
</taxonomicName>
sp. D”) within 
<taxonomicName id="12EA4D65C01A5533BF3BF9DBBFB5F9A8" box="[846,1021,1637,1662]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="3" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C01A5533BF3BF9DBBFB5F9A8" box="[846,1021,1637,1662]" italics="true" pageId="3" pageNumber="28">Engystomops</emphasis>
</taxonomicName>
. The following morphological synapomorphies (
<bibRefCitation id="B17B4B17C01A5533BD1AF934BED9F972" author="Cannatella" box="[367,657,1674,1700]" pageId="3" pageNumber="28" refString="Cannatella, D. C., Hillis, D. M., Chippindale, P. T., Weigt, L., Rand, A. S. &amp; Ryan, M. J. (1998) Phylogeny of frogs of the Physalaemus pustulosus species group, with an examination of data incongruence. Systematic Biology, 47, 311 - 335." type="journal article" year="1998">
Cannatella, 
<emphasis id="E79EEAF4C01A5533BE71F935BE0AF972" box="[516,578,1675,1700]" italics="true" pageId="3" pageNumber="28">et al.</emphasis>
, 1998
</bibRefCitation>
) also support this assignment: (1) presence of flank glands; (2) presence of parotoid glands; and (3) warty skin.
</paragraph>
<paragraph id="D55536E6C01A5533BCB0F96BBD69F822" blockId="3.[151,1438,1002,2036]" pageId="3" pageNumber="28">
<taxonomicName id="12EA4D65C01A5533BCB0F96BBD85F938" box="[197,461,1749,1774]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="3" pageNumber="28" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01A5533BCB0F96BBD85F938" box="[197,461,1749,1774]" italics="true" pageId="3" pageNumber="28">Engystomops puyango</emphasis>
</taxonomicName>
(
<figureCitation id="4DD12A63C01A5533BDA8F96BBE6AF939" box="[477,546,1749,1775]" captionStart="FIGURE 2" captionStartId="5.[151,255,1299,1323]" captionTargetBox="[318,1268,184,1277]" captionTargetId="figure@5.[318,1269,184,1278]" captionTargetPageId="5" captionText="FIGURE 2. Dorsolateral and ventral views of (A) Engystomops puyango, QCAZ 26988, adult female, SVL = 26.54; (B) E. puyango, QCAZ 26987, adult male, SVL = 28.19, and (C) E. pustulatus, QCAZ 26114, adult male, SVL = 28.5. Engystomops puyango from Bosque Protector Puyango (Provincia El Oro, Ecuador); E. pustulatus, from La Maná (Provincia Cotopaxi, Ecuador)." httpUri="https://zenodo.org/record/197712/files/figure.png" pageId="3" pageNumber="28">Fig. 2</figureCitation>
) is characterized by: (1) mean SVL 
<quantity id="12129B03C01A5533BFBCF96BB808F939" box="[969,1088,1749,1775]" metricMagnitude="-2" metricUnit="m" metricValue="2.759" pageId="3" pageNumber="28" unit="mm" value="27.59">27.59 mm</quantity>
in males (range 23.78–30.48; 
<emphasis id="E79EEAF4C01A5533BCE2F945BCEEF8C2" box="[151,166,1787,1812]" italics="true" pageId="3" pageNumber="28">n</emphasis>
= 45), 
<quantity id="12129B03C01A5533BC8EF944BD3BF8C2" box="[251,371,1786,1812]" metricMagnitude="-2" metricUnit="m" metricValue="2.868" pageId="3" pageNumber="28" unit="mm" value="28.68">28.68 mm</quantity>
in females (range 25.41–32.68; 
<emphasis id="E79EEAF4C01A5533BE85F945BEB7F8C2" box="[752,767,1787,1812]" italics="true" pageId="3" pageNumber="28">n</emphasis>
= 72); (2) skin on dorsum bearing a mixture of scattered pustules and minute tubercles; (3) snout varying between rounded and protruding in dorsal view and rounded to subacuminate in lateral view; (4) vomerine teeth and odontophores absent; (5) maxillary and premaxillary teeth present; (6) parotoid and flank glands present, usually fused, mean length = 
<quantity id="12129B03C01A5533B815F8D4B891F852" box="[1120,1241,1898,1924]" metricMagnitude="-2" metricUnit="m" metricValue="1.6629999999999998" pageId="3" pageNumber="28" unit="mm" value="16.63">16.63 mm</quantity>
(SD = 1.05; 
<emphasis id="E79EEAF4C01A5533B901F8D5B9CBF852" box="[1396,1411,1899,1924]" italics="true" pageId="3" pageNumber="28">n</emphasis>
= 28; 56.9–66.2% of SVL); (7) tarsal tubercle absent; (8) nuptial pads present; (9) Finger I shorter than II; (10) tympanic annulus evident, concealed dorsally and posteriorly; (11) tympanic membrane almost always not tuberculate.
</paragraph>
<caption id="8195666EC01D5534BCE2F8E4BD88F847" httpUri="https://zenodo.org/record/197711/files/figure.png" pageId="4" pageNumber="29" targetBox="[153,1435,217,1850]" targetPageId="4">
<paragraph id="D55536E6C01D5534BCE2F8E4BD88F847" blockId="4.[151,1436,1882,1937]" pageId="4" pageNumber="29">
<emphasis id="E79EEAF4C01D5534BCE2F8E4BD57F8A4" bold="true" box="[151,287,1882,1906]" pageId="4" pageNumber="29">FIGURE 1.</emphasis>
Lateral (top), ventral (left), and dorsal (right) views of the holotype of 
<taxonomicName id="12EA4D65C01D5534B830F8E4B908F8A7" box="[1093,1344,1882,1905]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="4" pageNumber="29" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01D5534B830F8E4B908F8A7" box="[1093,1344,1882,1905]" italics="true" pageId="4" pageNumber="29">Engystomops puyango</emphasis>
</taxonomicName>
(QCAZ 26978). Not drawn to scale.
</paragraph>
</caption>
<caption id="8195666EC01C5535BCE2FAADBDADFA5C" httpUri="https://zenodo.org/record/197712/files/figure.png" pageId="5" pageNumber="30" targetBox="[318,1268,184,1277]" targetPageId="5">
<paragraph id="D55536E6C01C5535BCE2FAADBDADFA5C" blockId="5.[151,1437,1299,1418]" pageId="5" pageNumber="30">
<emphasis id="E79EEAF4C01C5535BCE2FAADBD51FAFD" bold="true" box="[151,281,1299,1323]" pageId="5" pageNumber="30">FIGURE 2.</emphasis>
Dorsolateral and ventral views of (A) 
<taxonomicName id="12EA4D65C01C5535BEC6FAADBFECFAFC" box="[691,932,1299,1322]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="5" pageNumber="30" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01C5535BEC6FAADBFECFAFC" box="[691,932,1299,1322]" italics="true" pageId="5" pageNumber="30">Engystomops puyango</emphasis>
</taxonomicName>
, QCAZ 26988, adult female, SVL = 26.54; (B) 
<taxonomicName id="12EA4D65C01C5535BCE2FA8DBD5DFA9C" box="[151,277,1331,1354]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="5" pageNumber="30" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01C5535BCE2FA8DBD5DFA9C" box="[151,277,1331,1354]" italics="true" pageId="5" pageNumber="30">E. puyango</emphasis>
</taxonomicName>
, QCAZ 26987, adult male, SVL = 28.19, and (C) 
<taxonomicName id="12EA4D65C01C5535BF3CFA8DBF93FA9C" box="[841,987,1331,1354]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="5" pageNumber="30" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C01C5535BF3CFA8DBF93FA9C" box="[841,987,1331,1354]" italics="true" pageId="5" pageNumber="30">E. pustulatus</emphasis>
</taxonomicName>
, QCAZ 26114, adult male, SVL = 28.5. 
<taxonomicName id="12EA4D65C01C5535BCE2FAEDBDDBFABC" box="[151,403,1363,1386]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="5" pageNumber="30" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01C5535BCE2FAEDBDDBFABC" box="[151,403,1363,1386]" italics="true" pageId="5" pageNumber="30">Engystomops puyango</emphasis>
</taxonomicName>
from Bosque Protector 
<taxonomicName id="12EA4D65C01C5535BED7FAEDBF4BFABC" box="[674,771,1363,1386]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="5" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
(Provincia El Oro, Ecuador); 
<taxonomicName id="12EA4D65C01C5535B82DFAEDB8A3FABC" box="[1112,1259,1363,1386]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="5" pageNumber="30" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C01C5535B82DFAEDB8A3FABC" box="[1112,1259,1363,1386]" italics="true" pageId="5" pageNumber="30">E. pustulatus</emphasis>
</taxonomicName>
, from La Maná (Provincia Cotopaxi, Ecuador).
</paragraph>
</caption>
<caption id="8195666EC01C5535BCE2F82EBD1BF839" httpUri="https://zenodo.org/record/197713/files/figure.png" pageId="5" pageNumber="30" targetBox="[152,1435,1462,1913]" targetPageId="5">
<paragraph id="D55536E6C01C5535BCE2F82EBD1BF839" blockId="5.[151,1436,1936,2031]" pageId="5" pageNumber="30">
<emphasis id="E79EEAF4C01C5535BCE2F82EBD51F87E" bold="true" box="[151,281,1936,1960]" pageId="5" pageNumber="30">FIGURE 3.</emphasis>
Dorsal photographs of adult 
<taxonomicName id="12EA4D65C01C5535BE25F82FBF08F87E" box="[592,832,1937,1960]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="5" pageNumber="30" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01C5535BE25F82FBF08F87E" box="[592,832,1937,1960]" italics="true" pageId="5" pageNumber="30">Engystomops puyango</emphasis>
</taxonomicName>
(A) QCAZ 26980, (B) QCAZ 31506, (C) QCAZ 26998, (D) QCAZ 2880; and 
<taxonomicName id="12EA4D65C01C5535BDFEF80DBE53F81C" box="[395,539,1971,1994]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="5" pageNumber="30" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C01C5535BDFEF80DBE53F81C" box="[395,539,1971,1994]" italics="true" pageId="5" pageNumber="30">E. pustulatus</emphasis>
</taxonomicName>
(E) QCAZ 26736, showing differences in skin texture. Note that 
<taxonomicName id="12EA4D65C01C5535B886F80DB938F81C" box="[1267,1392,1971,1994]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="5" pageNumber="30" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01C5535B886F80DB938F81C" box="[1267,1392,1971,1994]" italics="true" pageId="5" pageNumber="30">E. puyango</emphasis>
</taxonomicName>
has smaller tubercles.
</paragraph>
</caption>
<caption id="8195666EC01F5536BCE2FF27B8D9FEC0" ID-Table-UUID="8195666EC01F5536BCE2FF27B8D9FEC0" httpUri="http://table.plazi.org/id/8195666EC01F5536BCE2FF27B8D9FEC0" pageId="6" pageNumber="31" targetBox="[151,1436,423,863]" targetIsTable="true" targetPageId="6">
<paragraph id="D55536E6C01F5536BCE2FF27B8D9FEC0" blockId="6.[151,1436,152,404]" pageId="6" pageNumber="31">
<emphasis id="E79EEAF4C01F5536BCE2FF27BD48FF78" bold="true" box="[151,256,153,174]" pageId="6" pageNumber="31">TABLE 3.</emphasis>
Morphometry of karyotypes of 
<taxonomicName id="12EA4D65C01F5536BE2BFF26BF1AFF79" box="[606,850,152,175]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BE2BFF26BF1AFF79" box="[606,850,152,175]" italics="true" pageId="6" pageNumber="31">Engystomops puyango</emphasis>
</taxonomicName>
and 
<emphasis id="E79EEAF4C01F5536BFFDFF26B855FF79" box="[904,1053,152,175]" italics="true" pageId="6" pageNumber="31">
<taxonomicName id="12EA4D65C01F5536BFFDFF26B851FF79" box="[904,1049,152,175]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="pustulatus">E. pustulatus</taxonomicName>
.
</emphasis>
Means and standard deviations are from 25 metaphases from three males (QCAZ 47296, 46932, 47295) and five females (QCAZ 35690–91, 46934, 47297, SC 26360) of 
<taxonomicName id="12EA4D65C01F5536BD40FF63BDFBFF22" box="[309,435,221,244]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BD40FF63BDFBFF22" box="[309,435,221,244]" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
from Bosque Protector 
<taxonomicName id="12EA4D65C01F5536BEB4FF63BF6CFF22" box="[705,804,221,244]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
, Ecuador; and four of 
<taxonomicName id="12EA4D65C01F5536B855FF63B8FAFF22" box="[1056,1202,221,244]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C01F5536B855FF63B8FAFF22" box="[1056,1202,221,244]" italics="true" pageId="6" pageNumber="31">E. pustulatus</emphasis>
</taxonomicName>
(from Cerro Blanco, Ecuador; QCAZ 47031–33, QCAZ 46931). Abbreviations are: m = metacentric; sm = submetacentric.
</paragraph>
</caption>
<paragraph id="D55536E6C01F5536BDBEFE8CB8B4FE9F" blockId="6.[151,1436,152,404]" box="[459,1276,306,329]" pageId="6" pageNumber="31">
<emphasis id="E79EEAF4C01F5536BDBEFE8CB8B4FE9F" box="[459,1276,306,329]" italics="true" pageId="6" pageNumber="31">
<taxonomicName id="12EA4D65C01F5536BDBEFE8CBEF4FE9F" box="[459,700,306,329]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">Engystomops puyango</taxonomicName>
<taxonomicName id="12EA4D65C01F5536BF8FFE8CB8B4FE9F" box="[1018,1276,306,329]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="pustulatus">Engystomops pustulatus</taxonomicName>
</emphasis>
</paragraph>
<paragraph id="D55536E6C01F5536BCEAFEE3B94CFE42" blockId="6.[151,1436,152,404]" pageId="6" pageNumber="31">
Chromosome Relative Arm Centromeric 
<typeStatus id="0A518844C01F5536BF21FEE3BFCFFEA2" box="[852,903,349,372]" pageId="6" pageNumber="31">Type</typeStatus>
Relative Centromeric 
<typeStatus id="0A518844C01F5536B939FEE3B9C9FEA2" box="[1356,1409,349,372]" pageId="6" pageNumber="31">Type</typeStatus>
number Length 
<emphasis id="E79EEAF4C01F5536BDE0FEC2BDECFE42" bold="true" box="[405,420,380,404]" pageId="6" pageNumber="31">±</emphasis>
SD Radio 
<emphasis id="E79EEAF4C01F5536BE4FFEC2BE01FE42" bold="true" box="[570,585,380,404]" pageId="6" pageNumber="31">±</emphasis>
SD Index 
<emphasis id="E79EEAF4C01F5536BE92FEC2BEBEFE42" bold="true" box="[743,758,380,404]" pageId="6" pageNumber="31">±</emphasis>
SD Length 
<emphasis id="E79EEAF4C01F5536B878FEC2B854FE42" bold="true" box="[1037,1052,380,404]" pageId="6" pageNumber="31">±</emphasis>
SD Index 
<emphasis id="E79EEAF4C01F5536B8B9FEC2B893FE42" bold="true" box="[1228,1243,380,404]" pageId="6" pageNumber="31">±</emphasis>
SD
</paragraph>
<paragraph id="D55536E6C01F5536BCEAFE16B929FC89" pageId="6" pageNumber="31">
<table id="A7EAC446C01FAACFBCE2FE19B9D4FC89" box="[151,1436,423,863]" gridcols="7" gridrows="10" pageId="6" pageNumber="31">
<tr id="6BDA34A4C01FAACFBCE2FE19B9D4FE69" box="[151,1436,423,447]" gridrow="0" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FE19BCA8FE69" box="[151,224,423,447]" gridcol="0" gridrow="0" pageId="6" pageNumber="31">1</th>
<th id="280B5DD8C01FAACFBD31FE19BE30FE69" box="[324,632,423,447]" gridcol="1" gridrow="0" pageId="6" pageNumber="31">
14.08 (
<emphasis id="E79EEAF4C01F5536BDFAFE19BDD6FE69" bold="true" box="[399,414,423,447]" pageId="6" pageNumber="31">±</emphasis>
0.08) 1.36 (
<emphasis id="E79EEAF4C01F5536BE47FE19BE09FE69" bold="true" box="[562,577,423,447]" pageId="6" pageNumber="31">±</emphasis>
0.02)
</th>
<th id="280B5DD8C01FAACFBED1FE19BF60FE69" box="[676,808,423,447]" gridcol="2" gridrow="0" pageId="6" pageNumber="31">
0.42 (
<emphasis id="E79EEAF4C01F5536BE97FE19BEB9FE69" bold="true" box="[738,753,423,447]" pageId="6" pageNumber="31">±</emphasis>
0.01)
</th>
<th id="280B5DD8C01FAACFBF21FE19BF3BFE69" box="[852,883,423,447]" gridcol="3" gridrow="0" pageId="6" pageNumber="31">m</th>
<th id="280B5DD8C01FAACFBFCEFE19B804FE69" box="[955,1100,423,447]" gridcol="4" gridrow="0" pageId="6" pageNumber="31">
15.45 (
<emphasis id="E79EEAF4C01F5536B873FE19B85DFE69" bold="true" box="[1030,1045,423,447]" pageId="6" pageNumber="31">±</emphasis>
0.10)
</th>
<th id="280B5DD8C01FAACFB8FCFE19B945FE69" box="[1161,1293,423,447]" gridcol="5" gridrow="0" pageId="6" pageNumber="31">
0.43 (
<emphasis id="E79EEAF4C01F5536B8B2FE19B89EFE69" bold="true" box="[1223,1238,423,447]" pageId="6" pageNumber="31">±</emphasis>
0.03)
</th>
<th id="280B5DD8C01FAACFB939FE19B9D4FE69" box="[1356,1436,423,447]" gridcol="6" gridrow="0" pageId="6" pageNumber="31">m</th>
</tr>
<tr id="6BDA34A4C01FAACFBCE2FE6CB9D4FE3C" box="[151,1436,466,490]" gridrow="1" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FE6CBCA8FE3C" box="[151,224,466,490]" gridcol="0" gridrow="1" pageId="6" pageNumber="31">2</th>
<td id="280B5DD8C01FAACFBD31FE6CBE30FE3C" box="[324,632,466,490]" gridcol="1" gridrow="1" pageId="6" pageNumber="31">
12.80 (
<emphasis id="E79EEAF4C01F5536BDFAFE6CBDD6FE3C" bold="true" box="[399,414,466,490]" pageId="6" pageNumber="31">±</emphasis>
0.12) 1.81 (
<emphasis id="E79EEAF4C01F5536BE47FE6CBE09FE3C" bold="true" box="[562,577,466,490]" pageId="6" pageNumber="31">±</emphasis>
0.05)
</td>
<td id="280B5DD8C01FAACFBED1FE6CBF60FE3C" box="[676,808,466,490]" gridcol="2" gridrow="1" pageId="6" pageNumber="31">
0.36 (
<emphasis id="E79EEAF4C01F5536BE97FE6CBEB9FE3C" bold="true" box="[738,753,466,490]" pageId="6" pageNumber="31">±</emphasis>
0.06)
</td>
<td id="280B5DD8C01FAACFBF21FE6CBF3BFE3C" box="[852,883,466,490]" gridcol="3" gridrow="1" pageId="6" pageNumber="31">sm</td>
<td id="280B5DD8C01FAACFBFCEFE6CB804FE3C" box="[955,1100,466,490]" gridcol="4" gridrow="1" pageId="6" pageNumber="31">
12.54 (
<emphasis id="E79EEAF4C01F5536B873FE6CB85DFE3C" bold="true" box="[1030,1045,466,490]" pageId="6" pageNumber="31">±</emphasis>
0.09)
</td>
<td id="280B5DD8C01FAACFB8FCFE6CB945FE3C" box="[1161,1293,466,490]" gridcol="5" gridrow="1" pageId="6" pageNumber="31">
0.37 (
<emphasis id="E79EEAF4C01F5536B8B2FE6CB89EFE3C" bold="true" box="[1223,1238,466,490]" pageId="6" pageNumber="31">±</emphasis>
0.02)
</td>
<td id="280B5DD8C01FAACFB939FE6CB9D4FE3C" box="[1356,1436,466,490]" gridcol="6" gridrow="1" pageId="6" pageNumber="31">sm</td>
</tr>
<tr id="6BDA34A4C01FAACFBCE2FE42B9D4FDC2" box="[151,1436,508,532]" gridrow="2" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FE42BCA8FDC2" box="[151,224,508,532]" gridcol="0" gridrow="2" pageId="6" pageNumber="31">3</th>
<td id="280B5DD8C01FAACFBD31FE42BE30FDC2" box="[324,632,508,532]" gridcol="1" gridrow="2" pageId="6" pageNumber="31">
11.66 (
<emphasis id="E79EEAF4C01F5536BDFBFE42BDD5FDC2" bold="true" box="[398,413,508,532]" pageId="6" pageNumber="31">±</emphasis>
0.01) 1.93 (
<emphasis id="E79EEAF4C01F5536BE47FE42BE09FDC2" bold="true" box="[562,577,508,532]" pageId="6" pageNumber="31">±</emphasis>
0.05)
</td>
<td id="280B5DD8C01FAACFBED1FE42BF60FDC2" box="[676,808,508,532]" gridcol="2" gridrow="2" pageId="6" pageNumber="31">
0.34 (
<emphasis id="E79EEAF4C01F5536BE97FE42BEB9FDC2" bold="true" box="[738,753,508,532]" pageId="6" pageNumber="31">±</emphasis>
0.04)
</td>
<td id="280B5DD8C01FAACFBF21FE42BF3BFDC2" box="[852,883,508,532]" gridcol="3" gridrow="2" pageId="6" pageNumber="31">sm</td>
<td id="280B5DD8C01FAACFBFCEFE42B804FDC2" box="[955,1100,508,532]" gridcol="4" gridrow="2" pageId="6" pageNumber="31">
10.9 (
<emphasis id="E79EEAF4C01F5536BF8CFE42B840FDC2" bold="true" box="[1017,1032,508,532]" pageId="6" pageNumber="31">±</emphasis>
0.04)
</td>
<td id="280B5DD8C01FAACFB8FCFE42B945FDC2" box="[1161,1293,508,532]" gridcol="5" gridrow="2" pageId="6" pageNumber="31">
0.34 (
<emphasis id="E79EEAF4C01F5536B8B2FE42B89EFDC2" bold="true" box="[1223,1238,508,532]" pageId="6" pageNumber="31">±</emphasis>
0.04)
</td>
<td id="280B5DD8C01FAACFB939FE42B9D4FDC2" box="[1356,1436,508,532]" gridcol="6" gridrow="2" pageId="6" pageNumber="31">sm</td>
</tr>
<tr id="6BDA34A4C01FAACFBCE2FD99B9D4FDE9" box="[151,1436,551,575]" gridrow="3" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FD99BCA8FDE9" box="[151,224,551,575]" gridcol="0" gridrow="3" pageId="6" pageNumber="31">4</th>
<td id="280B5DD8C01FAACFBD31FD99BE30FDE9" box="[324,632,551,575]" gridcol="1" gridrow="3" pageId="6" pageNumber="31">
10.53 (
<emphasis id="E79EEAF4C01F5536BDFAFD99BDD6FDE9" bold="true" box="[399,414,551,575]" pageId="6" pageNumber="31">±</emphasis>
0.17) 2.70 (
<emphasis id="E79EEAF4C01F5536BE47FD99BE09FDE9" bold="true" box="[562,577,551,575]" pageId="6" pageNumber="31">±</emphasis>
0.06)
</td>
<td id="280B5DD8C01FAACFBED1FD99BF60FDE9" box="[676,808,551,575]" gridcol="2" gridrow="3" pageId="6" pageNumber="31">
0.27 (
<emphasis id="E79EEAF4C01F5536BE97FD99BEB9FDE9" bold="true" box="[738,753,551,575]" pageId="6" pageNumber="31">±</emphasis>
0.07)
</td>
<td id="280B5DD8C01FAACFBF21FD99BF3BFDE9" box="[852,883,551,575]" gridcol="3" gridrow="3" pageId="6" pageNumber="31">sm</td>
<td id="280B5DD8C01FAACFBFCEFD99B804FDE9" box="[955,1100,551,575]" gridcol="4" gridrow="3" pageId="6" pageNumber="31">
10.77 (
<emphasis id="E79EEAF4C01F5536B873FD99B85DFDE9" bold="true" box="[1030,1045,551,575]" pageId="6" pageNumber="31">±</emphasis>
0.11)
</td>
<td id="280B5DD8C01FAACFB8FCFD99B945FDE9" box="[1161,1293,551,575]" gridcol="5" gridrow="3" pageId="6" pageNumber="31">
0.45 (
<emphasis id="E79EEAF4C01F5536B8B2FD99B89EFDE9" bold="true" box="[1223,1238,551,575]" pageId="6" pageNumber="31">±</emphasis>
0.05)
</td>
<td id="280B5DD8C01FAACFB939FD99B9D4FDE9" box="[1356,1436,551,575]" gridcol="6" gridrow="3" pageId="6" pageNumber="31">m</td>
</tr>
<tr id="6BDA34A4C01FAACFBCE2FDECB9D4FDBC" box="[151,1436,594,618]" gridrow="4" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FDECBCA8FDBC" box="[151,224,594,618]" gridcol="0" gridrow="4" pageId="6" pageNumber="31">5</th>
<td id="280B5DD8C01FAACFBD31FDECBE30FDBC" box="[324,632,594,618]" gridcol="1" gridrow="4" pageId="6" pageNumber="31">
9.25 (
<emphasis id="E79EEAF4C01F5536BDF7FDECBDD9FDBC" bold="true" box="[386,401,594,618]" pageId="6" pageNumber="31">±</emphasis>
0.22) 1.32 (
<emphasis id="E79EEAF4C01F5536BE47FDECBE09FDBC" bold="true" box="[562,577,594,618]" pageId="6" pageNumber="31">±</emphasis>
0.04)
</td>
<td id="280B5DD8C01FAACFBED1FDECBF60FDBC" box="[676,808,594,618]" gridcol="2" gridrow="4" pageId="6" pageNumber="31">
0.43 (
<emphasis id="E79EEAF4C01F5536BE97FDECBEB9FDBC" bold="true" box="[738,753,594,618]" pageId="6" pageNumber="31">±</emphasis>
0.06)
</td>
<td id="280B5DD8C01FAACFBF21FDECBF3BFDBC" box="[852,883,594,618]" gridcol="3" gridrow="4" pageId="6" pageNumber="31">m</td>
<td id="280B5DD8C01FAACFBFCEFDECB804FDBC" box="[955,1100,594,618]" gridcol="4" gridrow="4" pageId="6" pageNumber="31">
10.44 (
<emphasis id="E79EEAF4C01F5536B873FDECB85DFDBC" bold="true" box="[1030,1045,594,618]" pageId="6" pageNumber="31">±</emphasis>
0.28)
</td>
<td id="280B5DD8C01FAACFB8FCFDECB945FDBC" box="[1161,1293,594,618]" gridcol="5" gridrow="4" pageId="6" pageNumber="31">
0.34 (
<emphasis id="E79EEAF4C01F5536B8B2FDECB89EFDBC" bold="true" box="[1223,1238,594,618]" pageId="6" pageNumber="31">±</emphasis>
0.03)
</td>
<td id="280B5DD8C01FAACFB939FDECB9D4FDBC" box="[1356,1436,594,618]" gridcol="6" gridrow="4" pageId="6" pageNumber="31">sm</td>
</tr>
<tr id="6BDA34A4C01FAACFBCE2FDC2B9D4FD42" box="[151,1436,636,660]" gridrow="5" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FDC2BCA8FD42" box="[151,224,636,660]" gridcol="0" gridrow="5" pageId="6" pageNumber="31">6</th>
<td id="280B5DD8C01FAACFBD31FDC2BE30FD42" box="[324,632,636,660]" gridcol="1" gridrow="5" pageId="6" pageNumber="31">
9.25 (
<emphasis id="E79EEAF4C01F5536BDF7FDC2BDD9FD42" bold="true" box="[386,401,636,660]" pageId="6" pageNumber="31">±</emphasis>
0.09) 1.60 (
<emphasis id="E79EEAF4C01F5536BE47FDC2BE09FD42" bold="true" box="[562,577,636,660]" pageId="6" pageNumber="31">±</emphasis>
0.03)
</td>
<td id="280B5DD8C01FAACFBED1FDC2BF60FD42" box="[676,808,636,660]" gridcol="2" gridrow="5" pageId="6" pageNumber="31">
0.38 (
<emphasis id="E79EEAF4C01F5536BE97FDC2BEB9FD42" bold="true" box="[738,753,636,660]" pageId="6" pageNumber="31">±</emphasis>
0.03)
</td>
<td id="280B5DD8C01FAACFBF21FDC2BF3BFD42" box="[852,883,636,660]" gridcol="3" gridrow="5" pageId="6" pageNumber="31">m</td>
<td id="280B5DD8C01FAACFBFCEFDC2B804FD42" box="[955,1100,636,660]" gridcol="4" gridrow="5" pageId="6" pageNumber="31">
10.03 (
<emphasis id="E79EEAF4C01F5536B873FDC2B85DFD42" bold="true" box="[1030,1045,636,660]" pageId="6" pageNumber="31">±</emphasis>
0.17)
</td>
<td id="280B5DD8C01FAACFB8FCFDC2B945FD42" box="[1161,1293,636,660]" gridcol="5" gridrow="5" pageId="6" pageNumber="31">
0.31 (
<emphasis id="E79EEAF4C01F5536B8B2FDC2B89EFD42" bold="true" box="[1223,1238,636,660]" pageId="6" pageNumber="31">±</emphasis>
0.03)
</td>
<td id="280B5DD8C01FAACFB939FDC2B9D4FD42" box="[1356,1436,636,660]" gridcol="6" gridrow="5" pageId="6" pageNumber="31">sm</td>
</tr>
<tr id="6BDA34A4C01FAACFBCE2FD19B9D4FD69" box="[151,1436,679,703]" gridrow="6" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FD19BCA8FD69" box="[151,224,679,703]" gridcol="0" gridrow="6" pageId="6" pageNumber="31">7</th>
<td id="280B5DD8C01FAACFBD31FD19BE30FD69" box="[324,632,679,703]" gridcol="1" gridrow="6" pageId="6" pageNumber="31">
9.10 (
<emphasis id="E79EEAF4C01F5536BDF7FD19BDD9FD69" bold="true" box="[386,401,679,703]" pageId="6" pageNumber="31">±</emphasis>
0.07) 1.29 (
<emphasis id="E79EEAF4C01F5536BE47FD19BE09FD69" bold="true" box="[562,577,679,703]" pageId="6" pageNumber="31">±</emphasis>
0.04)
</td>
<td id="280B5DD8C01FAACFBED1FD19BF60FD69" box="[676,808,679,703]" gridcol="2" gridrow="6" pageId="6" pageNumber="31">
0.44 (
<emphasis id="E79EEAF4C01F5536BE97FD19BEB9FD69" bold="true" box="[738,753,679,703]" pageId="6" pageNumber="31">±</emphasis>
0.05)
</td>
<td id="280B5DD8C01FAACFBF21FD19BF3BFD69" box="[852,883,679,703]" gridcol="3" gridrow="6" pageId="6" pageNumber="31">m</td>
<td id="280B5DD8C01FAACFBFCEFD19B804FD69" box="[955,1100,679,703]" gridcol="4" gridrow="6" pageId="6" pageNumber="31">
9.02 (
<emphasis id="E79EEAF4C01F5536BF8CFD19B840FD69" bold="true" box="[1017,1032,679,703]" pageId="6" pageNumber="31">±</emphasis>
0.12)
</td>
<td id="280B5DD8C01FAACFB8FCFD19B945FD69" box="[1161,1293,679,703]" gridcol="5" gridrow="6" pageId="6" pageNumber="31">
0.42 (
<emphasis id="E79EEAF4C01F5536B8B2FD19B89EFD69" bold="true" box="[1223,1238,679,703]" pageId="6" pageNumber="31">±</emphasis>
0.07)
</td>
<td id="280B5DD8C01FAACFB939FD19B9D4FD69" box="[1356,1436,679,703]" gridcol="6" gridrow="6" pageId="6" pageNumber="31">m</td>
</tr>
<tr id="6BDA34A4C01FAACFBCE2FD6CB9D4FD3C" box="[151,1436,722,746]" gridrow="7" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FD6CBCA8FD3C" box="[151,224,722,746]" gridcol="0" gridrow="7" pageId="6" pageNumber="31">8</th>
<td id="280B5DD8C01FAACFBD31FD6CBE30FD3C" box="[324,632,722,746]" gridcol="1" gridrow="7" pageId="6" pageNumber="31">
8.82 (
<emphasis id="E79EEAF4C01F5536BDF7FD6CBDD9FD3C" bold="true" box="[386,401,722,746]" pageId="6" pageNumber="31">±</emphasis>
0.026) 2.10 (
<emphasis id="E79EEAF4C01F5536BE47FD6CBE09FD3C" bold="true" box="[562,577,722,746]" pageId="6" pageNumber="31">±</emphasis>
0.03)
</td>
<td id="280B5DD8C01FAACFBED1FD6CBF60FD3C" box="[676,808,722,746]" gridcol="2" gridrow="7" pageId="6" pageNumber="31">
0.32 (
<emphasis id="E79EEAF4C01F5536BE97FD6CBEB9FD3C" bold="true" box="[738,753,722,746]" pageId="6" pageNumber="31">±</emphasis>
0.02)
</td>
<td id="280B5DD8C01FAACFBF21FD6CBF3BFD3C" box="[852,883,722,746]" gridcol="3" gridrow="7" pageId="6" pageNumber="31">sm</td>
<td id="280B5DD8C01FAACFBFCEFD6CB804FD3C" box="[955,1100,722,746]" gridcol="4" gridrow="7" pageId="6" pageNumber="31">
8.62 (
<emphasis id="E79EEAF4C01F5536BF8CFD6CB840FD3C" bold="true" box="[1017,1032,722,746]" pageId="6" pageNumber="31">±</emphasis>
0.015)
</td>
<td id="280B5DD8C01FAACFB8FCFD6CB945FD3C" box="[1161,1293,722,746]" gridcol="5" gridrow="7" pageId="6" pageNumber="31">
0.37 (
<emphasis id="E79EEAF4C01F5536B8B2FD6CB89EFD3C" bold="true" box="[1223,1238,722,746]" pageId="6" pageNumber="31">±</emphasis>
0.04)
</td>
<td id="280B5DD8C01FAACFB939FD6CB9D4FD3C" box="[1356,1436,722,746]" gridcol="6" gridrow="7" pageId="6" pageNumber="31">sm</td>
</tr>
<tr id="6BDA34A4C01FAACFBCE2FD42B9D4FCE2" box="[151,1436,764,820]" gridrow="8" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FD42BCA8FCE2" box="[151,224,764,820]" gridcol="0" gridrow="8" pageId="6" pageNumber="31">9a, 9b</th>
<td id="280B5DD8C01FAACFBD31FD42BE30FCE2" box="[324,632,764,820]" gridcol="1" gridrow="8" pageId="6" pageNumber="31">
8.61 (
<emphasis id="E79EEAF4C01F5536BDF7FD42BDD9FCC2" bold="true" box="[386,401,764,788]" pageId="6" pageNumber="31">±</emphasis>
0.32) 1.63 (
<emphasis id="E79EEAF4C01F5536BE47FD42BE09FCC2" bold="true" box="[562,577,764,788]" pageId="6" pageNumber="31">±</emphasis>
0.07) 1.77 (
<emphasis id="E79EEAF4C01F5536BE47FCA2BE09FCE2" bold="true" box="[562,577,796,820]" pageId="6" pageNumber="31">±</emphasis>
0.06)
</td>
<td id="280B5DD8C01FAACFBED1FD42BF60FCE2" box="[676,808,764,820]" gridcol="2" gridrow="8" pageId="6" pageNumber="31">
0.38 (
<emphasis id="E79EEAF4C01F5536BE97FD42BEB9FCC2" bold="true" box="[738,753,764,788]" pageId="6" pageNumber="31">±</emphasis>
0.04) 0.36 (
<emphasis id="E79EEAF4C01F5536BE97FCA2BEB9FCE2" bold="true" box="[738,753,796,820]" pageId="6" pageNumber="31">±</emphasis>
0.03)
</td>
<td id="280B5DD8C01FAACFBF21FD42BF3BFCE2" box="[852,883,764,820]" gridcol="3" gridrow="8" pageId="6" pageNumber="31">m sm</td>
<td id="280B5DD8C01FAACFBFCEFD42B804FCE2" box="[955,1100,764,820]" gridcol="4" gridrow="8" pageId="6" pageNumber="31">
6.52 (
<emphasis id="E79EEAF4C01F5536BF8CFD42B840FCC2" bold="true" box="[1017,1032,764,788]" pageId="6" pageNumber="31">±</emphasis>
0.23)
</td>
<td id="280B5DD8C01FAACFB8FCFD42B945FCE2" box="[1161,1293,764,820]" gridcol="5" gridrow="8" pageId="6" pageNumber="31">
0.44 (
<emphasis id="E79EEAF4C01F5536B8B2FD42B89EFCC2" bold="true" box="[1223,1238,764,788]" pageId="6" pageNumber="31">±</emphasis>
0.08)
</td>
<td id="280B5DD8C01FAACFB939FD42B9D4FCE2" box="[1356,1436,764,820]" gridcol="6" gridrow="8" pageId="6" pageNumber="31">m</td>
</tr>
<tr id="6BDA34A4C01FAACFBCE2FCF9B9D4FC89" box="[151,1436,839,863]" gridrow="9" pageId="6" pageNumber="31">
<th id="280B5DD8C01FAACFBCE2FCF9BCA8FC89" box="[151,224,839,863]" gridcol="0" gridrow="9" pageId="6" pageNumber="31">10</th>
<td id="280B5DD8C01FAACFBD31FCF9BE30FC89" box="[324,632,839,863]" gridcol="1" gridrow="9" pageId="6" pageNumber="31">
5.90 (
<emphasis id="E79EEAF4C01F5536BDF7FCF9BDD9FC89" bold="true" box="[386,401,839,863]" pageId="6" pageNumber="31">±</emphasis>
0.19) 1.37 (
<emphasis id="E79EEAF4C01F5536BE47FCF9BE09FC89" bold="true" box="[562,577,839,863]" pageId="6" pageNumber="31">±</emphasis>
0.02)
</td>
<td id="280B5DD8C01FAACFBED1FCF9BF60FC89" box="[676,808,839,863]" gridcol="2" gridrow="9" pageId="6" pageNumber="31">
0.42 (
<emphasis id="E79EEAF4C01F5536BE97FCF9BEB9FC89" bold="true" box="[738,753,839,863]" pageId="6" pageNumber="31">±</emphasis>
0.01)
</td>
<td id="280B5DD8C01FAACFBF21FCF9BF3BFC89" box="[852,883,839,863]" gridcol="3" gridrow="9" pageId="6" pageNumber="31">m</td>
<td id="280B5DD8C01FAACFBFCEFCF9B804FC89" box="[955,1100,839,863]" gridcol="4" gridrow="9" pageId="6" pageNumber="31">
5.69 (
<emphasis id="E79EEAF4C01F5536BF8CFCF9B840FC89" bold="true" box="[1017,1032,839,863]" pageId="6" pageNumber="31">±</emphasis>
0.22)
</td>
<td id="280B5DD8C01FAACFB8FCFCF9B945FC89" box="[1161,1293,839,863]" gridcol="5" gridrow="9" pageId="6" pageNumber="31">
0.43 (
<emphasis id="E79EEAF4C01F5536B8B2FCF9B89EFC89" bold="true" box="[1223,1238,839,863]" pageId="6" pageNumber="31">±</emphasis>
0.07)
</td>
<td id="280B5DD8C01FAACFB939FCF9B9D4FC89" box="[1356,1436,839,863]" gridcol="6" gridrow="9" pageId="6" pageNumber="31">m</td>
</tr>
</table>
</paragraph>
<paragraph id="D55536E6C01F5536BCB3FC26BE93FAAE" blockId="6.[151,1437,920,2034]" pageId="6" pageNumber="31">
<taxonomicName id="12EA4D65C01F5536BCB3FC26BD87FC67" box="[198,463,920,945]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BCB3FC26BD87FC67" box="[198,463,920,945]" italics="true" pageId="6" pageNumber="31">Engystomops puyango</emphasis>
</taxonomicName>
is most similar to 
<taxonomicName id="12EA4D65C01F5536BEDFFC26BF0AFC67" box="[682,834,920,945]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C01F5536BEDFFC26BF0AFC67" box="[682,834,920,945]" italics="true" pageId="6" pageNumber="31">E. pustulatus</emphasis>
</taxonomicName>
. They differ in dorsal skin texture (less tuberculate in 
<taxonomicName id="12EA4D65C01F5536BCC1FC03BD72FC00" box="[180,314,957,982]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BCC1FC03BD72FC00" box="[180,314,957,982]" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
; 
<figureCitation id="4DD12A63C01F5536BD3DFC03BDC5FC01" box="[328,397,957,983]" captionStart="FIGURE 3" captionStartId="5.[151,255,1936,1960]" captionTargetBox="[152,1435,1462,1913]" captionTargetId="figure@5.[151,1436,1462,1915]" captionTargetPageId="5" captionText="FIGURE 3. Dorsal photographs of adult Engystomops puyango (A) QCAZ 26980, (B) QCAZ 31506, (C) QCAZ 26998, (D) QCAZ 2880; and E. pustulatus (E) QCAZ 26736, showing differences in skin texture. Note that E. puyango has smaller tubercles." httpUri="https://zenodo.org/record/197713/files/figure.png" pageId="6" pageNumber="31">Fig. 3</figureCitation>
), chromosomes shape and size (
<tableCitation id="9868035DC01F5536BF70FC03BF16FC01" box="[773,862,957,983]" captionStart="TABLE 3" captionStartId="6.[151,231,153,174]" captionTargetBox="[151,1436,423,863]" captionTargetPageId="6" captionText="TABLE 3. Morphometry of karyotypes of Engystomops puyango and E. pustulatus. Means and standard deviations are from 25 metaphases from three males (QCAZ 47296, 46932, 47295) and five females (QCAZ 35690 – 91, 46934, 47297, SC 26360) of E. puyango from Bosque Protector Puyango, Ecuador; and four of E. pustulatus (from Cerro Blanco, Ecuador; QCAZ 47031 – 33, QCAZ 46931). Abbreviations are: m = metacentric; sm = submetacentric." httpUri="http://table.plazi.org/id/8195666EC01F5536BCE2FF27B8D9FEC0" pageId="6" pageNumber="31" tableUuid="8195666EC01F5536BCE2FF27B8D9FEC0">Table 3</tableCitation>
), and advertisement call (shorter and with higher frequency in 
<taxonomicName id="12EA4D65C01F5536BD4CFC5CBD8CFC2D" box="[313,452,994,1019]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BD4CFC5CBD8CFC2D" box="[313,452,994,1019]" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
; 
<figureCitation id="4DD12A63C01F5536BDA0FC5CBE69FC2A" box="[469,545,994,1020]" captionStart="FIGURE 4" captionStartId="9.[151,255,983,1007]" captionTargetBox="[373,1217,189,958]" captionTargetId="figure@9.[369,1218,180,962]" captionTargetPageId="9" captionText="FIGURE 4. Axes I and II from Principal Components Analysis based on eight acoustic variables from the advertisement calls of Engystomops puyango (10 males) and E. pustulatus (39). See Table 2 for character loadings on each component." httpUri="https://zenodo.org/record/197714/files/figure.png" pageId="6" pageNumber="31">Fig. 4</figureCitation>
). Calls are significantly different in total duration (
<emphasis id="E79EEAF4C01F5536B8E3FC5CB8D6FC2D" box="[1174,1182,994,1019]" italics="true" pageId="6" pageNumber="31">t</emphasis>
= 9.68, df = 26, 
<emphasis id="E79EEAF4C01F5536B905FC5CB9CAFC2D" box="[1392,1410,994,1019]" italics="true" pageId="6" pageNumber="31">P</emphasis>
&lt;0.001), duration of the first component (
<emphasis id="E79EEAF4C01F5536BE1AFBB5BE3FFBF2" box="[623,631,1035,1060]" italics="true" pageId="6" pageNumber="31">t</emphasis>
= 4.01, df = 24, 
<emphasis id="E79EEAF4C01F5536BF48FBB5BF07FBF2" box="[829,847,1035,1060]" italics="true" pageId="6" pageNumber="31">P</emphasis>
&lt;0.001), frequency of the 2nd spectral peak of the call (
<emphasis id="E79EEAF4C01F5536BCA6FB8DBC93FB9A" box="[211,219,1075,1100]" italics="true" pageId="6" pageNumber="31">t</emphasis>
= 910.47, df = 43, 
<emphasis id="E79EEAF4C01F5536BDB4FB8DBD9BFB9A" box="[449,467,1075,1100]" italics="true" pageId="6" pageNumber="31">P</emphasis>
&lt;0.001), and frequency of the 2nd spectral peak of the first component (
<emphasis id="E79EEAF4C01F5536B95FFB8DB97AFB9A" box="[1322,1330,1075,1100]" italics="true" pageId="6" pageNumber="31">t</emphasis>
= 10.65, df = 45, 
<emphasis id="E79EEAF4C01F5536BC8BFBE7BD58FBA4" box="[254,272,1113,1138]" italics="true" pageId="6" pageNumber="31">P</emphasis>
&lt;0.001; Table 4). Genetically, large distances, typical of species pairs in 
<taxonomicName id="12EA4D65C01F5536B8F9FBE7B963FBA4" box="[1164,1323,1113,1138]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C01F5536B8F9FBE7B963FBA4" box="[1164,1323,1113,1138]" italics="true" pageId="6" pageNumber="31">Engystomops</emphasis>
</taxonomicName>
, separate both species. Pairwise uncorrected 
<emphasis id="E79EEAF4C01F5536BE47FBC0BE08FB41" box="[562,576,1150,1175]" italics="true" pageId="6" pageNumber="31">p</emphasis>
-distances between 
<taxonomicName id="12EA4D65C01F5536BF50FBC0BFF5FB41" box="[805,957,1150,1175]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C01F5536BF50FBC0BFF5FB41" box="[805,957,1150,1175]" italics="true" pageId="6" pageNumber="31">E. pustulatus</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C01F5536BF82FBC0B834FB41" box="[1015,1148,1150,1175]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BF82FBC0B834FB41" box="[1015,1148,1150,1175]" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
(range 0.065–0.068) are higher than distances between sister species 
<taxonomicName id="12EA4D65C01F5536BEDBFB1DBF09FB6A" box="[686,833,1187,1212]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="montubio">
<emphasis id="E79EEAF4C01F5536BEDBFB1DBF09FB6A" box="[686,833,1187,1212]" italics="true" pageId="6" pageNumber="31">E. montubio</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C01F5536BF09FB1DBF97FB6A" box="[892,991,1187,1212]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C01F5536BF09FB1DBF97FB6A" box="[892,991,1187,1212]" italics="true" pageId="6" pageNumber="31">E. randi</emphasis>
</taxonomicName>
(0.027–0.032) or even between nonsister species like 
<taxonomicName id="12EA4D65C01F5536BD1BFB77BDBCFB34" box="[366,500,1225,1250]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="guayaco">
<emphasis id="E79EEAF4C01F5536BD1BFB77BDBCFB34" box="[366,500,1225,1250]" italics="true" pageId="6" pageNumber="31">E. guayaco</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C01F5536BE5BFB77BEF7FB34" box="[558,703,1225,1250]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="montubio">
<emphasis id="E79EEAF4C01F5536BE5BFB77BEF7FB34" box="[558,703,1225,1250]" italics="true" pageId="6" pageNumber="31">E. montubio</emphasis>
</taxonomicName>
(0.059), and 
<taxonomicName id="12EA4D65C01F5536BF2AFB77BFB8FB34" box="[863,1008,1225,1250]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="montubio">
<emphasis id="E79EEAF4C01F5536BF2AFB77BFB8FB34" box="[863,1008,1225,1250]" italics="true" pageId="6" pageNumber="31">E. montubio</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C01F5536B85EFB77B8AEFB34" box="[1067,1254,1225,1250]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="coloradorum">
<emphasis id="E79EEAF4C01F5536B85EFB77B8AEFB34" box="[1067,1254,1225,1250]" italics="true" pageId="6" pageNumber="31">E. coloradorum</emphasis>
</taxonomicName>
(0.057–0.060). However, genetic distances should not be used as the only evidence for species delimitation (
<bibRefCitation id="B17B4B17C01F5536B8ADFB50BC90FAFB" author="Vences" pageId="6" pageNumber="31" refString="Vences, M. &amp; Wake, D. B. (2007) Speciation, species boundaries and phylogeography of amphibians. In: Heatwole, H. &amp; Tyler, M. J. (Eds.) Amphibian Biology: Systematics. Surrey Beatty &amp; Sons Pty. Limited, Chipping Norton." type="book" year="2007">
Vences &amp; 
<collectingCountry id="ADFD7676C01F5536B927FB50B9DFFADE" box="[1362,1431,1262,1288]" name="Wake Island" pageId="6" pageNumber="31">Wake</collectingCountry>
, 2007
</bibRefCitation>
). In our dataset, nevertheless, the genetic data is corroborated by the pattern of differentiation in skin texture, advertisement calls, and chromosome morphology and is consistent with the recognition of 
<taxonomicName id="12EA4D65C01F5536B9F7FA87BCB4FAA1" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536B9F7FA87BCB4FAA1" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
as a separate species from 
<taxonomicName id="12EA4D65C01F5536BE4EFAE0BE9BFAA1" box="[571,723,1374,1399]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C01F5536BE4EFAE0BE9BFAA1" box="[571,723,1374,1399]" italics="true" pageId="6" pageNumber="31">E. pustulatus</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="9DF0656DC01F5536BCB0FA3DBF53F91E" pageId="6" pageNumber="31" type="reference_group">
<paragraph id="D55536E6C01F5536BCB0FA3DBF53F91E" blockId="6.[151,1437,920,2034]" pageId="6" pageNumber="31">
<taxonomicName id="12EA4D65C01F5536BCB0FA3DBD85FA4A" box="[197,461,1411,1436]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BCB0FA3DBD85FA4A" box="[197,461,1411,1436]" italics="true" pageId="6" pageNumber="31">Engystomops puyango</emphasis>
</taxonomicName>
is larger than 
<taxonomicName id="12EA4D65C01F5536BE04FA3DBE98FA4A" box="[625,720,1411,1436]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C01F5536BE04FA3DBE98FA4A" box="[625,720,1411,1436]" italics="true" pageId="6" pageNumber="31">E. randi</emphasis>
</taxonomicName>
, 
<taxonomicName id="12EA4D65C01F5536BEAAFA3DBF25FA4A" box="[735,877,1411,1436]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="montubio">
<emphasis id="E79EEAF4C01F5536BEAAFA3DBF25FA4A" box="[735,877,1411,1436]" italics="true" pageId="6" pageNumber="31">E. montubio</emphasis>
</taxonomicName>
, and 
<taxonomicName id="12EA4D65C01F5536BFD9FA3DB878FA4A" box="[940,1072,1411,1436]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="guayaco">
<emphasis id="E79EEAF4C01F5536BFD9FA3DB878FA4A" box="[940,1072,1411,1436]" italics="true" pageId="6" pageNumber="31">E. guayaco</emphasis>
</taxonomicName>
(non overlapping SVL in adult males) and has less extensive to absent lateral fringes on the toes (fringes are prominent in 
<taxonomicName id="12EA4D65C01F5536B97EFA17B939FA14" box="[1291,1393,1449,1474]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C01F5536B97EFA17B939FA14" box="[1291,1393,1449,1474]" italics="true" pageId="6" pageNumber="31">E. randi</emphasis>
</taxonomicName>
, 
<taxonomicName id="12EA4D65C01F5536B9F7FA17BD41FA31" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="montubio">
<emphasis id="E79EEAF4C01F5536B9F7FA17BD41FA31" italics="true" pageId="6" pageNumber="31">E. montubio</emphasis>
</taxonomicName>
, and 
<taxonomicName id="12EA4D65C01F5536BD38FA70BD86FA31" box="[333,462,1486,1511]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="guayaco">
<emphasis id="E79EEAF4C01F5536BD38FA70BD86FA31" box="[333,462,1486,1511]" italics="true" pageId="6" pageNumber="31">E guayaco</emphasis>
</taxonomicName>
; 
<bibRefCitation id="B17B4B17C01F5536BDAAFA70BEF8FA3E" author="Ron" box="[479,688,1486,1512]" pageId="6" pageNumber="31" refString="Ron, S. R., Cannatella, D. C. &amp; Coloma, L. A. (2004) Two new species of Physalaemus (Anura: Leptodactylidae) from western Ecuador. Herpetologica, 60, 261 - 275." type="journal article" year="2004">
Ron, 
<emphasis id="E79EEAF4C01F5536BE57FA70BE28FA31" box="[546,608,1486,1511]" italics="true" pageId="6" pageNumber="31">et al.</emphasis>
, 2004
</bibRefCitation>
; 
<bibRefCitation id="B17B4B17C01F5536BEC8FA70BFD8FA3E" author="Ron" box="[701,912,1486,1512]" pageId="6" pageNumber="31" refString="Ron, S. R., Coloma, L. A. &amp; Cannatella, D. C. (2005) A new, cryptic species of Physalaemus (Anura: Leptodactylidae) from western Ecuador with comments on the call structure of the P. pustulosus species group. Herpetologica, 61, 178 - 198." type="journal article" year="2005">
Ron, 
<emphasis id="E79EEAF4C01F5536BF75FA70BF75FA31" box="[768,829,1486,1511]" italics="true" pageId="6" pageNumber="31">et al.</emphasis>
, 2005
</bibRefCitation>
). Furthermore, the advertisement call of 
<taxonomicName id="12EA4D65C01F5536B9F7FA70BCB4F9DA" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536B9F7FA70BCB4F9DA" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
is longer and lacks well-defined pulses at the beginning as in 
<taxonomicName id="12EA4D65C01F5536BFADFA4DB870F9DA" box="[984,1080,1523,1548]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C01F5536BFADFA4DB870F9DA" box="[984,1080,1523,1548]" italics="true" pageId="6" pageNumber="31">E. randi</emphasis>
</taxonomicName>
, 
<taxonomicName id="12EA4D65C01F5536B832FA4DB89FF9DA" box="[1095,1239,1523,1548]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="montubio">
<emphasis id="E79EEAF4C01F5536B832FA4DB89FF9DA" box="[1095,1239,1523,1548]" italics="true" pageId="6" pageNumber="31">E. montubio</emphasis>
</taxonomicName>
, and 
<taxonomicName id="12EA4D65C01F5536B96DFA4DB9D4F9DA" box="[1304,1436,1523,1548]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="guayaco">
<emphasis id="E79EEAF4C01F5536B96DFA4DB9D4F9DA" box="[1304,1436,1523,1548]" italics="true" pageId="6" pageNumber="31">E. guayaco</emphasis>
</taxonomicName>
(
<bibRefCitation id="B17B4B17C01F5536BCEAF9A6BDADF9E4" author="Ron" box="[159,485,1560,1586]" pageId="6" pageNumber="31" refString="Ron, S. R. (2008) The evolution of female mate choice for complex calls in tungara frogs. Animal Behaviour, 76, 1783 - 1794." type="journal article" year="2008">
Ron, 2008; Table 4, 
<figureCitation id="4DD12A63C01F5536BDEEF9A6BDADF9E4" box="[411,485,1560,1586]" captionStart="FIGURE 5" captionStartId="12.[151,258,1915,1939]" captionTargetBox="[154,1436,198,1899]" captionTargetId="figure@12.[151,1436,194,1903]" captionTargetPageId="12" captionText="FIGURE 5. Calls of Engystomops from western Ecuador. A – E: E. puyango (QCAZ 26968) from Bosque Protector Puyango, Provincia El Oro, Ecuador; F – J: E. pustulatus (QCAZ 26852) from El Empalme, Provincia Guayas, Ecuador. A and F are oscilograms; B and G spectrograms; C and H power spectra of complete call, D and I power spectra of the first component, E and J power spectra of the third component." httpUri="https://zenodo.org/record/197715/files/figure.png" pageId="6" pageNumber="31">Fig. 5</figureCitation>
</bibRefCitation>
). The absence of a tarsal tubercle and the presence of teeth in the maxilla and premaxilla distinguish 
<taxonomicName id="12EA4D65C01F5536BDD6F980BE61F981" box="[419,553,1598,1623]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BDD6F980BE61F981" box="[419,553,1598,1623]" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
from 
<taxonomicName id="12EA4D65C01F5536BE05F980BEA9F981" box="[624,737,1598,1623]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="petersi">
<emphasis id="E79EEAF4C01F5536BE05F980BEA9F981" box="[624,737,1598,1623]" italics="true" pageId="6" pageNumber="31">E. petersi</emphasis>
</taxonomicName>
, 
<taxonomicName id="12EA4D65C01F5536BE9AF980BF3CF981" box="[751,884,1598,1623]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="freibergi">
<emphasis id="E79EEAF4C01F5536BE9AF980BF3CF981" box="[751,884,1598,1623]" italics="true" pageId="6" pageNumber="31">E. freibergi</emphasis>
</taxonomicName>
, and 
<emphasis id="E79EEAF4C01F5536BFC1F980B9D4F981" box="[948,1436,1598,1623]" italics="true" pageId="6" pageNumber="31">
<taxonomicName id="12EA4D65C01F5536BFC1F980B81CF981" box="[948,1108,1598,1623]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="pustulosus">E. pustulosus</taxonomicName>
. 
<taxonomicName id="12EA4D65C01F5536B82AF980B9D4F981" box="[1119,1436,1598,1623]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="coloradorum">Engystomops coloradorum</taxonomicName>
</emphasis>
has a shorter advertisement call (
<bibRefCitation id="B17B4B17C01F5536BE5CF9DDBF57F9AB" author="Ryan" box="[553,799,1635,1661]" pageId="6" pageNumber="31" refString="Ryan, M. J. &amp; Rand, A. S. (2001) Feature weighting in signal recognition and discrimination by tungara frogs. In: Ryan, M. J. (Ed.) Anuran Communication. Smithsonian Institution Press, Washington." type="book" year="2001">Ryan &amp; Rand, 2001</bibRefCitation>
) and more prominent dorsal tubercles (tubercles are smaller and scattered in 
<taxonomicName id="12EA4D65C01F5536BDC8F937BE0EF974" box="[445,582,1673,1698]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BDC8F937BE0EF974" box="[445,582,1673,1698]" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
; 
<figureCitation id="4DD12A63C01F5536BE23F936BED7F974" box="[598,671,1672,1698]" captionStart="FIGURE 3" captionStartId="5.[151,255,1936,1960]" captionTargetBox="[152,1435,1462,1913]" captionTargetId="figure@5.[151,1436,1462,1915]" captionTargetPageId="5" captionText="FIGURE 3. Dorsal photographs of adult Engystomops puyango (A) QCAZ 26980, (B) QCAZ 31506, (C) QCAZ 26998, (D) QCAZ 2880; and E. pustulatus (E) QCAZ 26736, showing differences in skin texture. Note that E. puyango has smaller tubercles." httpUri="https://zenodo.org/record/197713/files/figure.png" pageId="6" pageNumber="31">Fig. 3</figureCitation>
). 
<taxonomicName id="12EA4D65C01F5536BEC3F937BFB2F974" box="[694,1018,1673,1698]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="coloradorum">
<emphasis id="E79EEAF4C01F5536BEC3F937BFB2F974" box="[694,1018,1673,1698]" italics="true" pageId="6" pageNumber="31">Engystomops coloradorum</emphasis>
</taxonomicName>
further differs from 
<taxonomicName id="12EA4D65C01F5536B881F937B935F974" box="[1268,1405,1673,1698]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536B881F937B935F974" box="[1268,1405,1673,1698]" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
in having a vertical loreal region (oblique in 
<taxonomicName id="12EA4D65C01F5536BEF0F910BF42F911" box="[645,778,1710,1735]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="6" pageNumber="31" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C01F5536BEF0F910BF42F911" box="[645,778,1710,1735]" italics="true" pageId="6" pageNumber="31">E. puyango</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="9DF0656DC01F553ABCB3F96CBD35FF2A" lastPageId="10" lastPageNumber="35" pageId="6" pageNumber="31" type="description">
<paragraph id="D55536E6C01F5536BCB3F96CBE27F824" blockId="6.[151,1437,920,2034]" pageId="6" pageNumber="31">
<emphasis id="E79EEAF4C01F5536BCB3F96CBDBDF93A" bold="true" box="[198,501,1746,1772]" pageId="6" pageNumber="31">
Description of 
<typeStatus id="0A518844C01F5536BDF6F96CBDB9F93A" box="[387,497,1746,1772]" pageId="6" pageNumber="31" type="holotype">holotype</typeStatus>
.
</emphasis>
Adult male, 
<quantity id="12129B03C01F5536BEE0F96DBF46F93B" box="[661,782,1747,1773]" metricMagnitude="-2" metricUnit="m" metricValue="2.7329999999999997" pageId="6" pageNumber="31" unit="mm" value="27.33">27.33 mm</quantity>
SVL, tibia length 
<quantity id="12129B03C01F5536BF85F96DB825F93B" box="[1008,1133,1747,1773]" metricMagnitude="-2" metricUnit="m" metricValue="1.2429999999999999" pageId="6" pageNumber="31" unit="mm" value="12.43">12.43 mm</quantity>
, femur length 
<quantity id="12129B03C01F5536B969F96DB9D0F93B" box="[1308,1432,1747,1773]" metricMagnitude="-2" metricUnit="m" metricValue="1.184" pageId="6" pageNumber="31" unit="mm" value="11.84">11.84 mm</quantity>
, arm length 
<quantity id="12129B03C01F5536BD68F946BDC0F8C4" box="[285,392,1784,1810]" metricMagnitude="-3" metricUnit="m" metricValue="6.17" pageId="6" pageNumber="31" unit="mm" value="6.17">6.17 mm</quantity>
, head length 
<quantity id="12129B03C01F5536BE50F946BED8F8C4" box="[549,656,1784,1810]" metricMagnitude="-3" metricUnit="m" metricValue="8.82" pageId="6" pageNumber="31" unit="mm" value="8.82">8.82 mm</quantity>
, head width 
<quantity id="12129B03C01F5536BF53F946BFDAF8C4" box="[806,914,1784,1810]" metricMagnitude="-3" metricUnit="m" metricValue="9.02" pageId="6" pageNumber="31" unit="mm" value="9.02">9.02 mm</quantity>
, eye-nostril distance 
<quantity id="12129B03C01F5536B8FFF946B8AFF8C4" box="[1162,1255,1784,1810]" metricMagnitude="-3" metricUnit="m" metricValue="2.7" pageId="6" pageNumber="31" unit="mm" value="2.7">2.7 mm</quantity>
, head narrower than body except in scapular region; diameter of eye 2.2 times diameter of tympanic annulus; tympanic membrane and tympanic annulus barely evident; tympanic annulus ovoid, longer dorsoventrally; few small tubercles on anterior margin of tympanic annulus, absent from tympanic membrane; supratympanic fold absent; head between the orbits and intercanthal region flat except for scattered tubercles; snout protruding in profile and slightly truncated in dorsal view; nostrils slightly elevated, internarial region convex; canthus rostralis rounded; loreal region concave.
</paragraph>
<caption id="8195666EC0105539BCE2FC69B9D4FBD9" httpUri="https://zenodo.org/record/197714/files/figure.png" pageId="9" pageNumber="34" targetBox="[373,1217,189,958]" targetPageId="9">
<paragraph id="D55536E6C0105539BCE2FC69B9D4FBD9" blockId="9.[151,1436,983,1039]" pageId="9" pageNumber="34">
<emphasis id="E79EEAF4C0105539BCE2FC69BD51FC39" bold="true" box="[151,281,983,1007]" pageId="9" pageNumber="34">FIGURE 4.</emphasis>
Axes I and II from Principal Components Analysis based on eight acoustic variables from the advertisement calls of 
<taxonomicName id="12EA4D65C0105539BC9EFC46BD94FBD9" box="[235,476,1016,1039]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="9" pageNumber="34" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0105539BC9EFC46BD94FBD9" box="[235,476,1016,1039]" italics="true" pageId="9" pageNumber="34">Engystomops puyango</emphasis>
</taxonomicName>
(10 males) and 
<taxonomicName id="12EA4D65C0105539BEF2FC46BF5AFBD9" box="[647,786,1016,1039]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="9" pageNumber="34" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C0105539BEF2FC46BF5AFBD9" box="[647,786,1016,1039]" italics="true" pageId="9" pageNumber="34">E. pustulatus</emphasis>
</taxonomicName>
(39). See Table 2 for character loadings on each component.
</paragraph>
</caption>
<paragraph id="D55536E6C0105539BCB0FBE6BE73F9DA" blockId="9.[151,1437,1112,2034]" pageId="9" pageNumber="34">Fingers without expanded discs; nuptial pad present, keratinized, brown, divided in two portions, one covering posterior half of thenar tubercle, other covering Finger I dorsally and posteriorly along its proximal half and posteriorly to distal edge of first phalanx. Base of thenar tubercle ovoid, that of palmar tubercle nearly round; palmar tubercle less prominent than thenar tubercle; subarticular tubercles with round base, all conical; second subarticular tubercle on Finger IV present; few supernumerary palmar tubercles present. Webbing between fingers absent; relative lengths of adpressed fingers III&gt; IV&gt; II&gt; I. Toes without expanded discs; base of inner metatarsal tubercle ovoid, larger than round base of outer metatarsal tubercle; inner metatarsal tubercle more prominent than outer metatarsal tubercle; subarticular tubercles with round base except for proximal tubercles of Toe III and IV (ovoid base), all conical except for subconical distal subarticular tubercles of Toe V; sparse and minute conical and subconical plantar supernumerary tubercles; tarsal tubercle absent. Lateral fringes on toes absent; webbing between toes absent; relative lengths of adpressed toes IV&gt; III&gt; V&gt; II&gt; I.</paragraph>
<paragraph id="D55536E6C0105539BCB0F9A6B943F974" blockId="9.[151,1437,1112,2034]" pageId="9" pageNumber="34">Skin on dorsum bearing a mixture of pustules and minute, round to subconical tubercles. Some tubercles are arranged in longitudinal rows; skin on venter smooth. Tongue longer than wide; vomerine teeth and odontophores absent. Maxillary and premaxillary teeth present. Vocal slits present, parallel to margins of mandible. Deflated vocal sac forming folds on gular region, extending posteriorly to base of arm.</paragraph>
<paragraph id="D55536E6C010553ABCB0F913BD35FF2A" blockId="9.[151,1437,1112,2034]" lastBlockId="10.[151,1437,152,2007]" lastPageId="10" lastPageNumber="35" pageId="9" pageNumber="34">
<emphasis id="E79EEAF4C0105539BCB0F913BE26F910" box="[197,622,1709,1734]" italics="true" pageId="9" pageNumber="34">
Color of 
<typeStatus id="0A518844C0105539BD42F913BDEAF910" box="[311,418,1709,1734]" pageId="9" pageNumber="34" type="holotype">holotype</typeStatus>
in preservative.
</emphasis>
Dorsum grayish brown with lighter gray on the snout; dark gray transversal bar between orbits and irregular dark marks on posterior half of dorsum; light gray middorsal blotch bordered by irregular dark bands at arm insertion axis; dorsal tubercles and pustules light gray; dorsal surfaces of forearms and hindlimbs light gray with dark gray transversal bands. Venter cream yellowish with few minute light gray marks along posterior one-third of the body; Dark-gray blotches on chest and anterior half of abdomen; minute dark gray spots abundant between arms and on posterior half of gular region; anterior half of gular region dark gray with light gray irregular marks close to midline; ventral surfaces of hindlimbs and forelimbs cream to cream yellowish, becoming light gray or cream towards outer and inner edges; some dark blotches present on shanks; outer half of ventral surfaces of forearms dark gray; sides of head gray with ill-defined dark gray marks on the snout and a cream area below the orbit and tympanum, extending posteriorly above deflated vocal folds and anteriorly as a labial stripe; flanks dark gray dorsally, light gray ventrally.
</paragraph>
</subSubSection>
<subSubSection id="9DF0656DC013553ABCB0FEB9BFF1FEBA" pageId="10" pageNumber="35" type="etymology">
<paragraph id="D55536E6C013553ABCB0FEB9BFF1FEBA" blockId="10.[151,1437,152,2007]" pageId="10" pageNumber="35">
<emphasis id="E79EEAF4C013553ABCB0FEB9BD1FFEF7" bold="true" box="[197,343,263,289]" pageId="10" pageNumber="35">Etymology.</emphasis>
The specific name 
<taxonomicName id="12EA4D65C013553ABE3FFEB6BEFBFEF7" box="[586,691,264,289]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="10" pageNumber="35" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C013553ABE3FFEB6BEFBFEF7" box="[586,691,264,289]" italics="true" pageId="10" pageNumber="35">puyango</emphasis>
</taxonomicName>
is a noun in apposition, in reference to the 
<typeStatus id="0A518844C013553AB8BAFEB6B94CFEF4" box="[1231,1284,264,290]" pageId="10" pageNumber="35">type</typeStatus>
locality, the 
<taxonomicName id="12EA4D65C013553ABCE2FE93BCB6FE91" box="[151,254,301,327]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="10" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
Protected Forest. The 2658 ha reserve has one of the world’s largest deposits of petrified tree trunks. It is also one of the largest reserves of tropical dry forest in 
<collectingCountry id="ADFD7676C013553ABF24FEECBFFDFEBA" box="[849,949,338,364]" name="Ecuador" pageId="10" pageNumber="35">Ecuador</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="9DF0656DC013553FBCB0FEC9B8C8FED7" lastPageId="15" lastPageNumber="40" pageId="10" pageNumber="35" type="description">
<paragraph id="D55536E6C013553ABCB0FEC9BE9FFD34" blockId="10.[151,1437,152,2007]" pageId="10" pageNumber="35">
<emphasis id="E79EEAF4C013553ABCB0FEC9BD08FE47" bold="true" box="[197,320,375,401]" pageId="10" pageNumber="35">Variation.</emphasis>
Variation in dorsal coloration of preserved specimens is extensive (
<figureCitation id="4DD12A63C013553AB82AFEC6B8EFFE44" box="[1119,1191,376,402]" captionStart="FIGURE 6" captionStartId="13.[151,256,1539,1563]" captionTargetBox="[321,1261,185,1519]" captionTargetId="figure@13.[318,1269,162,1523]" captionTargetPageId="13" captionText="FIGURE 6. Dorsal and ventral views of adult Engystomops puyango showing variation in dorsal and ventral patterns. From left to right, above: QCAZ 26976, 26982 – 83, 26986; below: QCAZ 27009, 27013, 37276 (all males). Provincia Loja and El Oro, Ecuador (See Appendix I for locality data)." httpUri="https://zenodo.org/record/197716/files/figure.png" pageId="10" pageNumber="35">Fig. 6</figureCitation>
). Background dorsal coloration varies from light gray (QCAZ 27005, 27009) to dark gray (QCAZ 31508) or dark brown (QCAZ 35691). Irregular dark marks may be present in diverse patterns (
<figureCitation id="4DD12A63C013553ABFFFFE7CBF99FE0A" box="[906,977,450,476]" captionStart="FIGURE 6" captionStartId="13.[151,256,1539,1563]" captionTargetBox="[321,1261,185,1519]" captionTargetId="figure@13.[318,1269,162,1523]" captionTargetPageId="13" captionText="FIGURE 6. Dorsal and ventral views of adult Engystomops puyango showing variation in dorsal and ventral patterns. From left to right, above: QCAZ 26976, 26982 – 83, 26986; below: QCAZ 27009, 27013, 37276 (all males). Provincia Loja and El Oro, Ecuador (See Appendix I for locality data)." httpUri="https://zenodo.org/record/197716/files/figure.png" pageId="10" pageNumber="35">Fig. 6</figureCitation>
). A clear mid-dorsal line extends along the posterior half of the body (QCAZ 36511) but in some specimens (QCAZ 26988, 26997) it only extends along the posterior one fifth. There is variation in the abundance and arrangement of tubercles (all lighter than the background; 
<figureCitation id="4DD12A63C013553ABD16FD8CBDFEFD9A" box="[355,438,562,588]" captionStart="FIGURE 3" captionStartId="5.[151,255,1936,1960]" captionTargetBox="[152,1435,1462,1913]" captionTargetId="figure@5.[151,1436,1462,1915]" captionTargetPageId="5" captionText="FIGURE 3. Dorsal photographs of adult Engystomops puyango (A) QCAZ 26980, (B) QCAZ 31506, (C) QCAZ 26998, (D) QCAZ 2880; and E. pustulatus (E) QCAZ 26736, showing differences in skin texture. Note that E. puyango has smaller tubercles." httpUri="https://zenodo.org/record/197713/files/figure.png" pageId="10" pageNumber="35">Figs. 3</figureCitation>
and 
<figureCitation id="4DD12A63C013553ABD86FD8CBE4DFD9A" box="[499,517,562,588]" captionStart="FIGURE 6" captionStartId="13.[151,256,1539,1563]" captionTargetBox="[321,1261,185,1519]" captionTargetId="figure@13.[318,1269,162,1523]" captionTargetPageId="13" captionText="FIGURE 6. Dorsal and ventral views of adult Engystomops puyango showing variation in dorsal and ventral patterns. From left to right, above: QCAZ 26976, 26982 – 83, 26986; below: QCAZ 27009, 27013, 37276 (all males). Provincia Loja and El Oro, Ecuador (See Appendix I for locality data)." httpUri="https://zenodo.org/record/197716/files/figure.png" pageId="10" pageNumber="35">6</figureCitation>
). Tubercles coalesce into small ridges that partly enclose a mid-dorsal light blotch in QCAZ 26984, 27005, 27012. The parotoid glands are fused with the flank glands in 28 out of 
<specimenCount id="C3ECFD6FC013553AB90AFDE6BD59FD41" pageId="10" pageNumber="35" type="generic">32 specimens</specimenCount>
(average gland length is 
<quantity id="12129B03C013553ABE49FDC3BEFFFD41" box="[572,695,637,663]" metricMagnitude="-2" metricUnit="m" metricValue="1.6629999999999998" pageId="10" pageNumber="35" unit="mm" value="16.63">16.63 mm</quantity>
, SD = 1.05; 56.9–66.2% of SVL). In QCAZ 27013, the glands are fused on the left side but separated on the right side. In QCAZ 26983 and 26989, the flank glands are separated from the parotoid glands on both sides.
</paragraph>
<paragraph id="D55536E6C013553ABCB0FD53BD97FBE4" blockId="10.[151,1437,152,2007]" pageId="10" pageNumber="35">
Ventral surfaces of preserved specimens have a cream to yellowish-cream background color with light gray (QCAZ 26991, 28762) to dark gray markings (QCAZ 26874, 26984). Marks are arranged in diverse patterns (
<figureCitation id="4DD12A63C013553ABD70FC86BD04FC84" box="[261,332,824,850]" captionStart="FIGURE 6" captionStartId="13.[151,256,1539,1563]" captionTargetBox="[321,1261,185,1519]" captionTargetId="figure@13.[318,1269,162,1523]" captionTargetPageId="13" captionText="FIGURE 6. Dorsal and ventral views of adult Engystomops puyango showing variation in dorsal and ventral patterns. From left to right, above: QCAZ 26976, 26982 – 83, 26986; below: QCAZ 27009, 27013, 37276 (all males). Provincia Loja and El Oro, Ecuador (See Appendix I for locality data)." httpUri="https://zenodo.org/record/197716/files/figure.png" pageId="10" pageNumber="35">Fig. 6</figureCitation>
) and vary from being restricted to the anterior half of the body (darker on folded vocal sacs; QCAZ 26968) to being present over the entire venter (less abundant posteriorly, QCAZ 27008). In a few specimens, the ventral marks are arranged in well-defined large spots (QCAZ 27002, 26983). A midventral cream stripe can be present from near the tip of the snout to the gular region (QCAZ 26994) or continue to the mid venter (QCAZ 26988). In most individuals the stripe is absent (QCAZ 27008) or ill defined (QCAZ 26986). The arrangement of dark spots and tubercles on the ventral surfaces of the feet and hands of QCAZ 26975 is shown in 
<figureCitation id="4DD12A63C013553ABD01FBA6BD92FBE4" box="[372,474,1048,1074]" captionStart="FIGURE 7" captionStartId="14.[151,255,1980,2004]" captionTargetBox="[204,1382,674,1959]" captionTargetId="figure@14.[204,1382,674,1959]" captionTargetPageId="14" captionText="FIGURE 7. Ventral views of the right hand and foot of Engystomops puyango, QCAZ 26975 (adult male from Bosque Protector Puyango, Ecuador; SVL = 28.16)." httpUri="https://zenodo.org/record/197717/files/figure.png" pageId="10" pageNumber="35">Figure 7</figureCitation>
.
</paragraph>
<paragraph id="D55536E6C013553ABCB0FB83B910FA8A" blockId="10.[151,1437,152,2007]" pageId="10" pageNumber="35">
Head shape varies between rounded (QCAZ 34511) and subacuminate (QCAZ 28760) in dorsal view; in lateral view it varies between rounded (QCAZ 26972) and protruding (QCAZ 31506, 26974). Lateral head coloration varies between light gray and dark gray. The area below the eye and tympanum is cream in most specimens (e.g., QCAZ 26987–88) but can be restricted to a thin longitudinal light stripe from the jaw articulation to the parotoid gland (QCAZ 26976). Two or three vertical dark bars can be present in the loreal region and below the orbit (e.g., QCAZ 26968, 26973, 26982). The tympanic annulus is concealed dorsally and posteriorly. With few exceptions (e.g., QCAZ 26984, 34511) the tympanic annulus has tubercles; the tympanic membrane has tubercles in only five specimens of the 
<typeStatus id="0A518844C013553ABFFDFAFCBFF3FA8A" box="[904,955,1346,1372]" pageId="10" pageNumber="35">type</typeStatus>
series (e.g., QCAZ 26989, 28758).
</paragraph>
<paragraph id="D55536E6C013553ABCB0FAD6B93CF9EA" blockId="10.[151,1437,152,2007]" pageId="10" pageNumber="35">
The following morphometric data pertain to adults. In the 
<typeStatus id="0A518844C013553ABF1BFAD6BFE9FA54" box="[878,929,1384,1410]" pageId="10" pageNumber="35">type</typeStatus>
series, the largest male has a SVL of 
<quantity id="12129B03C013553AB92FFAD6BC80FA71" metricMagnitude="-2" metricUnit="m" metricValue="3.048" pageId="10" pageNumber="35" unit="mm" value="30.48">30.48 mm</quantity>
, and the largest female 
<quantity id="12129B03C013553ABDAEFA33BE1DFA71" box="[475,597,1421,1447]" metricMagnitude="-2" metricUnit="m" metricValue="3.2619999999999996" pageId="10" pageNumber="35" unit="mm" value="32.62">32.62 mm</quantity>
; mean male SVL = 
<quantity id="12129B03C013553ABF4BFA33BFFDFA71" box="[830,949,1421,1447]" metricMagnitude="-2" metricUnit="m" metricValue="2.759" pageId="10" pageNumber="35" unit="mm" value="27.59">27.59 mm</quantity>
(
<emphasis id="E79EEAF4C013553ABFB0FA33BF9CFA70" box="[965,980,1421,1446]" italics="true" pageId="10" pageNumber="35">n</emphasis>
= 45; SD = 1.17), mean female SVL = 
<quantity id="12129B03C013553ABCE2FA0CBD5BFA1A" box="[151,275,1458,1484]" metricMagnitude="-2" metricUnit="m" metricValue="2.798" pageId="10" pageNumber="35" unit="mm" value="27.98">27.98 mm</quantity>
(
<emphasis id="E79EEAF4C013553ABD50FA0DBD7CFA1A" box="[293,308,1459,1484]" italics="true" pageId="10" pageNumber="35">n</emphasis>
= 6; SD = 2.38). Amplectant pairs measured during a behavioral study at Bosque Protector 
<taxonomicName id="12EA4D65C013553ABCE2FA66BCB6FA24" box="[151,254,1496,1522]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="10" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
(not collected) had mean male SVL = 26.76 (
<emphasis id="E79EEAF4C013553ABF6DFA66BF6FFA27" box="[792,807,1496,1521]" italics="true" pageId="10" pageNumber="35">n</emphasis>
= 72; SD = 1.62) and mean female SVL = 28.68 (
<emphasis id="E79EEAF4C013553AB900FA66B9CCFA27" box="[1397,1412,1496,1521]" italics="true" pageId="10" pageNumber="35">n</emphasis>
= 72; SD = 1.49). Females were significantly larger than males (
<emphasis id="E79EEAF4C013553ABF1EFA43BF3BF9C0" box="[875,883,1533,1558]" italics="true" pageId="10" pageNumber="35">t</emphasis>
= 6.19, df = 100, 
<emphasis id="E79EEAF4C013553AB837FA43B81CF9C0" box="[1090,1108,1533,1558]" italics="true" pageId="10" pageNumber="35">P</emphasis>
&lt;0.001). Snout-vent length was positively correlated between amplectant males and females (ANOVA 
<emphasis id="E79EEAF4C013553AB87EF99DB855F9EA" box="[1035,1053,1571,1596]" italics="true" pageId="10" pageNumber="35">F</emphasis>
= 19.93, df = 71, 
<emphasis id="E79EEAF4C013553AB885F99DB94AF9EA" box="[1264,1282,1571,1596]" italics="true" pageId="10" pageNumber="35">P</emphasis>
&lt;0.001).
</paragraph>
<paragraph id="D55536E6C013553ABCB0F9F6B96AF904" blockId="10.[151,1437,152,2007]" pageId="10" pageNumber="35">
Among the 
<typeStatus id="0A518844C013553ABD3BF9F6BDC8F9B4" box="[334,384,1608,1634]" pageId="10" pageNumber="35">type</typeStatus>
specimens, Catamayo has the highest male SVL (mean = 
<quantity id="12129B03C013553AB852F9F6B8E9F9B4" box="[1063,1185,1608,1634]" metricMagnitude="-2" metricUnit="m" metricValue="2.931" pageId="10" pageNumber="35" unit="mm" value="29.31">29.31 mm</quantity>
, SD = 0.51) followed by 
<taxonomicName id="12EA4D65C013553ABCCBF9D3BD61F951" box="[190,297,1645,1671]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="10" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
(27.50, SD = 0.95) and Mangaurquillo (26.17, SD = 1.75; 
<tableCitation id="9868035DC013553AB872F9D3B82DF951" box="[1031,1125,1645,1671]" captionStart="TABLE 5" captionStartId="11.[151,231,377,398]" captionTargetBox="[151,1436,522,1167]" captionTargetPageId="11" captionText="TABLE 5. Descriptive statistics for morphometric measurements of male Engystomops puyango and E. pustulatus used for Principal Component Analysis. Mean ± SD is given with range below. Bold figures are combined for males of all populations. Abbreviations are: SVL = snout-vent length; TL = tibia length; FL = femur length; AL = arm length; HL = head length; HW = head width; EN = eye-nostril distance. All measurements are in mm." httpUri="http://table.plazi.org/id/8195666EC012553BBCE2FEC7B80DFE39" pageId="10" pageNumber="35" tableUuid="8195666EC012553BBCE2FEC7B80DFE39">Table 5</tableCitation>
). Differences in SVL are significant between Catamayo and Mangaurquillo (Tukey’s HSD 
<emphasis id="E79EEAF4C013553ABFC3F92DBF80F97A" box="[950,968,1683,1708]" italics="true" pageId="10" pageNumber="35">P</emphasis>
&lt;0.001), and Catamayo and 
<taxonomicName id="12EA4D65C013553AB946F92CB9D5F97A" box="[1331,1437,1682,1708]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="10" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
(Tukey’s HSD 
<emphasis id="E79EEAF4C013553ABD32F906BD11F907" box="[327,345,1720,1745]" italics="true" pageId="10" pageNumber="35">P</emphasis>
= 0.020). All other pairwise comparisons were non-significant (
<emphasis id="E79EEAF4C013553AB839F906B816F907" box="[1100,1118,1720,1745]" italics="true" pageId="10" pageNumber="35">P</emphasis>
values&gt; 0.071).
</paragraph>
<paragraph id="D55536E6C013553ABCB3F963B9D4F801" blockId="10.[151,1437,152,2007]" pageId="10" pageNumber="35">
<emphasis id="E79EEAF4C013553ABCB3F963BD49F921" bold="true" box="[198,257,1757,1783]" pageId="10" pageNumber="35">Call.</emphasis>
Males call while floating on standing water. Acoustic parameters of the advertisement call of 
<taxonomicName id="12EA4D65C013553AB9F6F963BD48F8CA" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="10" pageNumber="35" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C013553AB9F6F963BD48F8CA" italics="true" pageId="10" pageNumber="35">E. puyango</emphasis>
</taxonomicName>
are shown in Table 4. The call consists of two obligatory and one facultative components with harmonic structure. The first component is characterized by a slight increase in frequency (mean = 48.0 Hz, SD = 67.24, 
<emphasis id="E79EEAF4C013553ABD40F8F3BD0CF8B0" box="[309,324,1869,1894]" italics="true" pageId="10" pageNumber="35">n</emphasis>
= 10) with decreasing amplitude modulation. Sidebands (from amplitude modulation) are evident in each harmonic but they merge at the end of the component (as a result, amplitude modulation ends; 
<figureCitation id="4DD12A63C013553ABCE2F826BC97F864" box="[151,223,1944,1970]" captionStart="FIGURE 5" captionStartId="12.[151,258,1915,1939]" captionTargetBox="[154,1436,198,1899]" captionTargetId="figure@12.[151,1436,194,1903]" captionTargetPageId="12" captionText="FIGURE 5. Calls of Engystomops from western Ecuador. A – E: E. puyango (QCAZ 26968) from Bosque Protector Puyango, Provincia El Oro, Ecuador; F – J: E. pustulatus (QCAZ 26852) from El Empalme, Provincia Guayas, Ecuador. A and F are oscilograms; B and G spectrograms; C and H power spectra of complete call, D and I power spectra of the first component, E and J power spectra of the third component." httpUri="https://zenodo.org/record/197715/files/figure.png" pageId="10" pageNumber="35">Fig. 5</figureCitation>
). The power spectrum along all the call shows three spectral peaks (
<figureCitation id="4DD12A63C013553AB867F826B811F864" box="[1042,1113,1944,1970]" captionStart="FIGURE 5" captionStartId="12.[151,258,1915,1939]" captionTargetBox="[154,1436,198,1899]" captionTargetId="figure@12.[151,1436,194,1903]" captionTargetPageId="12" captionText="FIGURE 5. Calls of Engystomops from western Ecuador. A – E: E. puyango (QCAZ 26968) from Bosque Protector Puyango, Provincia El Oro, Ecuador; F – J: E. pustulatus (QCAZ 26852) from El Empalme, Provincia Guayas, Ecuador. A and F are oscilograms; B and G spectrograms; C and H power spectra of complete call, D and I power spectra of the first component, E and J power spectra of the third component." httpUri="https://zenodo.org/record/197715/files/figure.png" pageId="10" pageNumber="35">Fig. 5</figureCitation>
C). Dominant frequency of the call was on the second spectral peak in calls from nine out of ten males (mean = 2022.3 Hz, SD = 30.8).
</paragraph>
<paragraph id="D55536E6C012553BBCE2FF26BDBCFE91" blockId="11.[151,1436,152,327]" pageId="11" pageNumber="36">Dominant frequency was 835.5 Hz in calls from the only male on which the dominant frequency was on the first spectral peak. The third spectral peak has high energy only in calls with the third component (see below). The dominant frequency of the first component was on the second spectral peak in calls of nine out of ten males (mean = 2017.6, SD = 40.5). Dominant frequency was 994 Hz in the only male on which the dominant was on the first spectral peak.</paragraph>
<caption id="8195666EC012553BBCE2FEC7B80DFE39" ID-Table-UUID="8195666EC012553BBCE2FEC7B80DFE39" httpUri="http://table.plazi.org/id/8195666EC012553BBCE2FEC7B80DFE39" pageId="11" pageNumber="36" targetBox="[151,1436,522,1167]" targetIsTable="true" targetPageId="11">
<paragraph id="D55536E6C012553BBCE2FEC7B80DFE39" blockId="11.[151,1436,375,495]" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBCE2FEC7BD4AFE59" bold="true" box="[151,258,375,399]" pageId="11" pageNumber="36">TABLE 5.</emphasis>
Descriptive statistics for morphometric measurements of male 
<taxonomicName id="12EA4D65C012553BBFDAFEC6B8E9FE59" box="[943,1185,376,399]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="11" pageNumber="36" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C012553BBFDAFEC6B8E9FE59" box="[943,1185,376,399]" italics="true" pageId="11" pageNumber="36">Engystomops puyango</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C012553BB8A2FEC6B92BFE59" box="[1239,1379,376,399]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="11" pageNumber="36" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C012553BB8A2FEC6B92BFE59" box="[1239,1379,376,399]" italics="true" pageId="11" pageNumber="36">E. pustulatus</emphasis>
</taxonomicName>
used for Principal Component Analysis. Mean ± SD is given with range below. Bold figures are combined for males of all populations. Abbreviations are: SVL = snout-vent length; TL = tibia length; FL = femur length; AL = arm length; HL = head length; HW = head width; EN = eye-nostril distance. All measurements are in mm.
</paragraph>
</caption>
<paragraph id="D55536E6C012553BBCEAFDB4B922FB59" pageId="11" pageNumber="36">
<table id="A7EAC446C012AACFBCE2FDB4B9D4FB59" box="[151,1436,522,1167]" gridcols="8" gridrows="10" pageId="11" pageNumber="36">
<tr id="6BDA34A4C012AACFBCE2FDB4B9D4FDC9" box="[151,1436,522,543]" gridrow="0" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FDB4BD66FDC9" box="[151,302,522,543]" gridcol="0" gridrow="0" pageId="11" pageNumber="36">Species</th>
<th id="280B5DD8C012AACFBD15FDB4BD91FDC9" box="[352,473,522,543]" gridcol="1" gridrow="0" pageId="11" pageNumber="36">SVL</th>
<th id="280B5DD8C012AACFBD89FDB4BE3EFDC9" box="[508,630,522,543]" gridcol="2" gridrow="0" pageId="11" pageNumber="36">TL</th>
<th id="280B5DD8C012AACFBEECFDB4BF5AFDC9" box="[665,786,522,543]" gridcol="3" gridrow="0" pageId="11" pageNumber="36">FL</th>
<th id="280B5DD8C012AACFBF40FDB4BFEAFDC9" box="[821,930,522,543]" gridcol="4" gridrow="0" pageId="11" pageNumber="36">AL</th>
<th id="280B5DD8C012AACFBFA4FDB4B876FDC9" box="[977,1086,522,543]" gridcol="5" gridrow="0" pageId="11" pageNumber="36">HL</th>
<th id="280B5DD8C012AACFB818FDB4B892FDC9" box="[1133,1242,522,543]" gridcol="6" gridrow="0" pageId="11" pageNumber="36">HW</th>
<th id="280B5DD8C012AACFB97FFDB4B9D4FDC9" box="[1290,1436,522,543]" gridcol="7" gridrow="0" pageId="11" pageNumber="36">EN</th>
</tr>
<tr id="6BDA34A4C012AACFBCE2FD8CB9D4FDB3" box="[151,1436,562,613]" gridrow="1" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FD8CBD66FDB3" box="[151,302,562,613]" gridcol="0" gridrow="1" pageId="11" pageNumber="36">
<taxonomicName id="12EA4D65C012553BBCEAFD8CBD54FD91" box="[159,284,562,583]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="11" pageNumber="36" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C012553BBCEAFD8CBD54FD91" box="[159,284,562,583]" italics="true" pageId="11" pageNumber="36">E. pustulatus</emphasis>
</taxonomicName>
(
<emphasis id="E79EEAF4C012553BBCD2FDEEBCFBFDB3" box="[167,179,592,613]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 51)
</th>
<td id="280B5DD8C012AACFBD15FD8CBD91FDB3" box="[352,473,562,613]" gridcol="1" gridrow="1" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBD15FD8CBD91FD91" bold="true" box="[352,473,562,583]" pageId="11" pageNumber="36">27.08 ± 1.36</emphasis>
</td>
<td id="280B5DD8C012AACFBD89FD8CBE3EFDB3" box="[508,630,562,613]" gridcol="2" gridrow="1" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBD89FD8CBE3CFD91" bold="true" box="[508,628,562,583]" pageId="11" pageNumber="36">11.42 ± 0.59</emphasis>
</td>
<td id="280B5DD8C012AACFBEECFD8CBF5AFDB3" box="[665,786,562,613]" gridcol="3" gridrow="1" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBEECFD8CBF58FD91" bold="true" box="[665,784,562,583]" pageId="11" pageNumber="36">11.50 ± 1.06</emphasis>
</td>
<td id="280B5DD8C012AACFBF40FD8CBFEAFDB3" box="[821,930,562,613]" gridcol="4" gridrow="1" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBF40FD8CBFEAFD91" bold="true" box="[821,930,562,583]" pageId="11" pageNumber="36">6.26 ± 0.36</emphasis>
</td>
<td id="280B5DD8C012AACFBFA4FD8CB876FDB3" box="[977,1086,562,613]" gridcol="5" gridrow="1" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBFA4FD8CB876FD91" bold="true" box="[977,1086,562,583]" pageId="11" pageNumber="36">8.73 ± 0.42</emphasis>
</td>
<td id="280B5DD8C012AACFB818FD8CB892FDB3" box="[1133,1242,562,613]" gridcol="6" gridrow="1" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BB818FD8CB891FD91" bold="true" box="[1133,1241,562,583]" pageId="11" pageNumber="36">9.11 ± 0.80</emphasis>
</td>
<td id="280B5DD8C012AACFB97FFD8CB9D4FDB3" box="[1290,1436,562,613]" gridcol="7" gridrow="1" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BB97FFD8CB93FFD91" bold="true" box="[1290,1399,562,583]" pageId="11" pageNumber="36">2.75 ± 0.23</emphasis>
</td>
</tr>
<tr id="6BDA34A4C012AACFBCE2FDC9B9D4FD7C" box="[151,1436,631,682]" gridrow="2" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FDC9BD66FD7C" box="[151,302,631,682]" gridcol="0" gridrow="2" pageId="11" pageNumber="36">
Arenillas (
<emphasis id="E79EEAF4C012553BBCD2FD2BBCFBFD7C" box="[167,179,661,682]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 10)
</th>
<td id="280B5DD8C012AACFBD15FDC9BD91FD7C" box="[352,473,631,682]" gridcol="1" gridrow="2" pageId="11" pageNumber="36">
28.33 
<emphasis id="E79EEAF4C012553BBDE9FDC9BDE1FD5A" bold="true" box="[412,425,631,652]" pageId="11" pageNumber="36">±</emphasis>
0.80 26.84–29.34
</td>
<td id="280B5DD8C012AACFBD89FDC9BE3EFD7C" box="[508,630,631,682]" gridcol="2" gridrow="2" pageId="11" pageNumber="36">
11.78 
<emphasis id="E79EEAF4C012553BBE42FDC9BE0CFD5A" bold="true" box="[567,580,631,652]" pageId="11" pageNumber="36">±</emphasis>
0.39 11.24–12.37
</td>
<td id="280B5DD8C012AACFBEECFDC9BF5AFD7C" box="[665,786,631,682]" gridcol="3" gridrow="2" pageId="11" pageNumber="36">
11.35 
<emphasis id="E79EEAF4C012553BBEA1FDC9BEA9FD5A" bold="true" box="[724,737,631,652]" pageId="11" pageNumber="36">±</emphasis>
0.41 10.85–12.28
</td>
<td id="280B5DD8C012AACFBF40FDC9BFEAFD7C" box="[821,930,631,682]" gridcol="4" gridrow="2" pageId="11" pageNumber="36">
6.50 
<emphasis id="E79EEAF4C012553BBF10FDC9BF3AFD5A" bold="true" box="[869,882,631,652]" pageId="11" pageNumber="36">±</emphasis>
0.24 6.22–6.99
</td>
<td id="280B5DD8C012AACFBFA4FDC9B876FD7C" box="[977,1086,631,682]" gridcol="5" gridrow="2" pageId="11" pageNumber="36">
8.99 
<emphasis id="E79EEAF4C012553BB874FDC9B846FD5A" bold="true" box="[1025,1038,631,652]" pageId="11" pageNumber="36">±</emphasis>
0.19 8.76–9.37
</td>
<td id="280B5DD8C012AACFB818FDC9B892FD7C" box="[1133,1242,631,682]" gridcol="6" gridrow="2" pageId="11" pageNumber="36">
8.72 
<emphasis id="E79EEAF4C012553BB8E8FDC9B8E2FD5A" bold="true" box="[1181,1194,631,652]" pageId="11" pageNumber="36">±</emphasis>
0.22 8.34–9.21
</td>
<td id="280B5DD8C012AACFB97FFDC9B9D4FD7C" box="[1290,1436,631,682]" gridcol="7" gridrow="2" pageId="11" pageNumber="36">
2.68 
<emphasis id="E79EEAF4C012553BB94FFDC9B90FFD5A" bold="true" box="[1338,1351,631,652]" pageId="11" pageNumber="36">±</emphasis>
0.13 2.37–2.85
</td>
</tr>
<tr id="6BDA34A4C012AACFBCE2FD03B9D4FD39" box="[151,1436,701,751]" gridrow="3" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FD03BD66FD39" box="[151,302,701,751]" gridcol="0" gridrow="3" pageId="11" pageNumber="36">
Huaquillas (
<emphasis id="E79EEAF4C012553BBCD2FD64BCFBFD39" box="[167,179,730,751]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 10)
</th>
<td id="280B5DD8C012AACFBD15FD03BD91FD39" box="[352,473,701,751]" gridcol="1" gridrow="3" pageId="11" pageNumber="36">
25.53 
<emphasis id="E79EEAF4C012553BBDE9FD03BDE1FD04" bold="true" box="[412,425,701,722]" pageId="11" pageNumber="36">±</emphasis>
1.13 23.90–28.14
</td>
<td id="280B5DD8C012AACFBD89FD03BE3EFD39" box="[508,630,701,751]" gridcol="2" gridrow="3" pageId="11" pageNumber="36">
10.64 
<emphasis id="E79EEAF4C012553BBE4DFD03BE0DFD04" bold="true" box="[568,581,701,722]" pageId="11" pageNumber="36">±</emphasis>
0.48 9.98–11.44
</td>
<td id="280B5DD8C012AACFBEECFD03BF5AFD39" box="[665,786,701,751]" gridcol="3" gridrow="3" pageId="11" pageNumber="36">
9.82 
<emphasis id="E79EEAF4C012553BBEBCFD03BE9EFD04" bold="true" box="[713,726,701,722]" pageId="11" pageNumber="36">±</emphasis>
0.50 9.02–10.73
</td>
<td id="280B5DD8C012AACFBF40FD03BFEAFD39" box="[821,930,701,751]" gridcol="4" gridrow="3" pageId="11" pageNumber="36">
5.98 
<emphasis id="E79EEAF4C012553BBF10FD03BF3AFD04" bold="true" box="[869,882,701,722]" pageId="11" pageNumber="36">±</emphasis>
0.33 5.25–6.46
</td>
<td id="280B5DD8C012AACFBFA4FD03B876FD39" box="[977,1086,701,751]" gridcol="5" gridrow="3" pageId="11" pageNumber="36">
8.54 
<emphasis id="E79EEAF4C012553BB874FD03B846FD04" bold="true" box="[1025,1038,701,722]" pageId="11" pageNumber="36">±</emphasis>
0.37 8.08–9.33
</td>
<td id="280B5DD8C012AACFB818FD03B892FD39" box="[1133,1242,701,751]" gridcol="6" gridrow="3" pageId="11" pageNumber="36">
7.92 
<emphasis id="E79EEAF4C012553BB8E8FD03B8E2FD04" bold="true" box="[1181,1194,701,722]" pageId="11" pageNumber="36">±</emphasis>
0.23 7.56–8.25
</td>
<td id="280B5DD8C012AACFB97FFD03B9D4FD39" box="[1290,1436,701,751]" gridcol="7" gridrow="3" pageId="11" pageNumber="36">
2.58 
<emphasis id="E79EEAF4C012553BB94FFD03B90FFD04" bold="true" box="[1338,1351,701,722]" pageId="11" pageNumber="36">±</emphasis>
0.07 2.49–2.72
</td>
</tr>
<tr id="6BDA34A4C012AACFBCE2FCBCB9D4FCE3" box="[151,1436,770,821]" gridrow="4" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FCBCBD66FCE3" box="[151,302,770,821]" gridcol="0" gridrow="4" pageId="11" pageNumber="36">
Patricia Pilar (
<emphasis id="E79EEAF4C012553BBCD2FC9EBCFBFCE3" box="[167,179,800,821]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 17)
</th>
<td id="280B5DD8C012AACFBD15FCBCBD91FCE3" box="[352,473,770,821]" gridcol="1" gridrow="4" pageId="11" pageNumber="36">
27.43 
<emphasis id="E79EEAF4C012553BBDE9FCBCBDE1FCC1" bold="true" box="[412,425,770,791]" pageId="11" pageNumber="36">±</emphasis>
1.24 25.17–29.88
</td>
<td id="280B5DD8C012AACFBD89FCBCBE3EFCE3" box="[508,630,770,821]" gridcol="2" gridrow="4" pageId="11" pageNumber="36">
11.59 
<emphasis id="E79EEAF4C012553BBE42FCBCBE0CFCC1" bold="true" box="[567,580,770,791]" pageId="11" pageNumber="36">±</emphasis>
0.53 10.65–12.72
</td>
<td id="280B5DD8C012AACFBEECFCBCBF5AFCE3" box="[665,786,770,821]" gridcol="3" gridrow="4" pageId="11" pageNumber="36">
12.09 
<emphasis id="E79EEAF4C012553BBEA1FCBCBEA9FCC1" bold="true" box="[724,737,770,791]" pageId="11" pageNumber="36">±</emphasis>
0.71 10.45–13.01
</td>
<td id="280B5DD8C012AACFBF40FCBCBFEAFCE3" box="[821,930,770,821]" gridcol="4" gridrow="4" pageId="11" pageNumber="36">
6.24 
<emphasis id="E79EEAF4C012553BBF10FCBCBF3AFCC1" bold="true" box="[869,882,770,791]" pageId="11" pageNumber="36">±</emphasis>
0.41 5.69–6.84
</td>
<td id="280B5DD8C012AACFBFA4FCBCB876FCE3" box="[977,1086,770,821]" gridcol="5" gridrow="4" pageId="11" pageNumber="36">
8.73 
<emphasis id="E79EEAF4C012553BB874FCBCB846FCC1" bold="true" box="[1025,1038,770,791]" pageId="11" pageNumber="36">±</emphasis>
0.54 7.68–9.6
</td>
<td id="280B5DD8C012AACFB818FCBCB892FCE3" box="[1133,1242,770,821]" gridcol="6" gridrow="4" pageId="11" pageNumber="36">
9.60 
<emphasis id="E79EEAF4C012553BB8E8FCBCB8E2FCC1" bold="true" box="[1181,1194,770,791]" pageId="11" pageNumber="36">±</emphasis>
0.47 8.63–10.28
</td>
<td id="280B5DD8C012AACFB97FFCBCB9D4FCE3" box="[1290,1436,770,821]" gridcol="7" gridrow="4" pageId="11" pageNumber="36">
2.84 
<emphasis id="E79EEAF4C012553BB94FFCBCB90FFCC1" bold="true" box="[1338,1351,770,791]" pageId="11" pageNumber="36">±</emphasis>
0.27 2.36–3.42
</td>
</tr>
<tr id="6BDA34A4C012AACFBCE2FCF9B9D4FCAC" box="[151,1436,839,890]" gridrow="5" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FCF9BD66FCAC" box="[151,302,839,890]" gridcol="0" gridrow="5" pageId="11" pageNumber="36">
Puerto Rico (
<emphasis id="E79EEAF4C012553BBCD2FCDBBCFBFCAC" box="[167,179,869,890]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 14)
</th>
<td id="280B5DD8C012AACFBD15FCF9BD91FCAC" box="[352,473,839,890]" gridcol="1" gridrow="5" pageId="11" pageNumber="36">26.86 ± 0.77 25.85–27.96</td>
<td id="280B5DD8C012AACFBD89FCF9BE3EFCAC" box="[508,630,839,890]" gridcol="2" gridrow="5" pageId="11" pageNumber="36">11.50 ± 0.33 10.94–12.14</td>
<td id="280B5DD8C012AACFBEECFCF9BF5AFCAC" box="[665,786,839,890]" gridcol="3" gridrow="5" pageId="11" pageNumber="36">12.08 ± 0.64 10.44–13.03</td>
<td id="280B5DD8C012AACFBF40FCF9BFEAFCAC" box="[821,930,839,890]" gridcol="4" gridrow="5" pageId="11" pageNumber="36">6.32 ± 0.26 5.76–6.71</td>
<td id="280B5DD8C012AACFBFA4FCF9B876FCAC" box="[977,1086,839,890]" gridcol="5" gridrow="5" pageId="11" pageNumber="36">8.70 ± 0.32 8.15–9.29</td>
<td id="280B5DD8C012AACFB818FCF9B892FCAC" box="[1133,1242,839,890]" gridcol="6" gridrow="5" pageId="11" pageNumber="36">9.65 ± 0.53 8.76–10.52</td>
<td id="280B5DD8C012AACFB97FFCF9B9D4FCAC" box="[1290,1436,839,890]" gridcol="7" gridrow="5" pageId="11" pageNumber="36">2.82 ± 0.23 2.31–3.17</td>
</tr>
<tr id="6BDA34A4C012AACFBCE2FC33B9D4FC69" box="[151,1436,909,959]" gridrow="6" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FC33BD66FC69" box="[151,302,909,959]" gridcol="0" gridrow="6" pageId="11" pageNumber="36">
<taxonomicName id="12EA4D65C012553BBCEAFC33BD44FC74" box="[159,268,909,930]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="11" pageNumber="36" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C012553BBCEAFC33BD44FC74" box="[159,268,909,930]" italics="true" pageId="11" pageNumber="36">E. puyango</emphasis>
</taxonomicName>
(
<emphasis id="E79EEAF4C012553BBCD2FC14BCFBFC69" box="[167,179,938,959]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 30)
</th>
<td id="280B5DD8C012AACFBD15FC33BD91FC69" box="[352,473,909,959]" gridcol="1" gridrow="6" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBD15FC33BD91FC74" bold="true" box="[352,473,909,930]" pageId="11" pageNumber="36">27.31 ± 1.16</emphasis>
</td>
<td id="280B5DD8C012AACFBD89FC33BE3EFC69" box="[508,630,909,959]" gridcol="2" gridrow="6" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBD89FC33BE3CFC74" bold="true" box="[508,628,909,930]" pageId="11" pageNumber="36">11.88 ± 0.58</emphasis>
</td>
<td id="280B5DD8C012AACFBEECFC33BF5AFC69" box="[665,786,909,959]" gridcol="3" gridrow="6" pageId="11" pageNumber="36">
11.10 
<emphasis id="E79EEAF4C012553BBEA1FC33BEA9FC74" bold="true" box="[724,737,909,930]" pageId="11" pageNumber="36">±</emphasis>
0.64
</td>
<td id="280B5DD8C012AACFBF40FC33BFEAFC69" box="[821,930,909,959]" gridcol="4" gridrow="6" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBF40FC33BFEAFC74" bold="true" box="[821,930,909,930]" pageId="11" pageNumber="36">6.36 ± 0.31</emphasis>
</td>
<td id="280B5DD8C012AACFBFA4FC33B876FC69" box="[977,1086,909,959]" gridcol="5" gridrow="6" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BBFA4FC33B876FC74" bold="true" box="[977,1086,909,930]" pageId="11" pageNumber="36">8.49 ± 0.44</emphasis>
</td>
<td id="280B5DD8C012AACFB818FC33B892FC69" box="[1133,1242,909,959]" gridcol="6" gridrow="6" pageId="11" pageNumber="36">
8.81 
<emphasis id="E79EEAF4C012553BB8E8FC33B8E2FC74" bold="true" box="[1181,1194,909,930]" pageId="11" pageNumber="36">±</emphasis>
0.61
</td>
<td id="280B5DD8C012AACFB97FFC33B9D4FC69" box="[1290,1436,909,959]" gridcol="7" gridrow="6" pageId="11" pageNumber="36">
<emphasis id="E79EEAF4C012553BB97FFC33B93FFC74" bold="true" box="[1290,1399,909,930]" pageId="11" pageNumber="36">2.65 ± 0.16</emphasis>
</td>
</tr>
<tr id="6BDA34A4C012AACFBCE2FC6CB9D4FBD3" box="[151,1436,978,1029]" gridrow="7" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FC6CBD66FBD3" box="[151,302,978,1029]" gridcol="0" gridrow="7" pageId="11" pageNumber="36">
Catamayo (
<emphasis id="E79EEAF4C012553BBCD2FC4EBCFBFBD3" box="[167,179,1008,1029]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 3)
</th>
<td id="280B5DD8C012AACFBD15FC6CBD91FBD3" box="[352,473,978,1029]" gridcol="1" gridrow="7" pageId="11" pageNumber="36">
29.31 
<emphasis id="E79EEAF4C012553BBDE9FC6CBDE1FC31" bold="true" box="[412,425,978,999]" pageId="11" pageNumber="36">±</emphasis>
0.51 28.76–29.79
</td>
<td id="280B5DD8C012AACFBD89FC6CBE3EFBD3" box="[508,630,978,1029]" gridcol="2" gridrow="7" pageId="11" pageNumber="36">
12.31 
<emphasis id="E79EEAF4C012553BBE4DFC6CBE0DFC31" bold="true" box="[568,581,978,999]" pageId="11" pageNumber="36">±</emphasis>
0.61 11.86–13.01
</td>
<td id="280B5DD8C012AACFBEECFC6CBF5AFBD3" box="[665,786,978,1029]" gridcol="3" gridrow="7" pageId="11" pageNumber="36">
11.84 
<emphasis id="E79EEAF4C012553BBEA1FC6CBEA9FC31" bold="true" box="[724,737,978,999]" pageId="11" pageNumber="36">±</emphasis>
0.82 11.36–12.80
</td>
<td id="280B5DD8C012AACFBF40FC6CBFEAFBD3" box="[821,930,978,1029]" gridcol="4" gridrow="7" pageId="11" pageNumber="36">
6.80 
<emphasis id="E79EEAF4C012553BBF10FC6CBF3AFC31" bold="true" box="[869,882,978,999]" pageId="11" pageNumber="36">±</emphasis>
0.32 6.58–7.17
</td>
<td id="280B5DD8C012AACFBFA4FC6CB876FBD3" box="[977,1086,978,1029]" gridcol="5" gridrow="7" pageId="11" pageNumber="36">
9.11 
<emphasis id="E79EEAF4C012553BB875FC6CB845FC31" bold="true" box="[1024,1037,978,999]" pageId="11" pageNumber="36">±</emphasis>
0.12 9.02–9.26
</td>
<td id="280B5DD8C012AACFB818FC6CB892FBD3" box="[1133,1242,978,1029]" gridcol="6" gridrow="7" pageId="11" pageNumber="36">
8.50 
<emphasis id="E79EEAF4C012553BB8E8FC6CB8E2FC31" bold="true" box="[1181,1194,978,999]" pageId="11" pageNumber="36">±</emphasis>
0.08 8.42–8.59
</td>
<td id="280B5DD8C012AACFB97FFC6CB9D4FBD3" box="[1290,1436,978,1029]" gridcol="7" gridrow="7" pageId="11" pageNumber="36">
2.65 
<emphasis id="E79EEAF4C012553BB94FFC6CB90FFC31" bold="true" box="[1338,1351,978,999]" pageId="11" pageNumber="36">±</emphasis>
0.05 2.61–2.71
</td>
</tr>
<tr id="6BDA34A4C012AACFBCE2FBA9B9D4FB9C" box="[151,1436,1047,1098]" gridrow="8" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FBA9BD66FB9C" box="[151,302,1047,1098]" gridcol="0" gridrow="8" pageId="11" pageNumber="36">
Mangaurquillo (
<emphasis id="E79EEAF4C012553BBCD2FB8BBCFBFB9C" box="[167,179,1077,1098]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 4)
</th>
<td id="280B5DD8C012AACFBD15FBA9BD91FB9C" box="[352,473,1047,1098]" gridcol="1" gridrow="8" pageId="11" pageNumber="36">
26.17 
<emphasis id="E79EEAF4C012553BBDE9FBA9BDE1FBFA" bold="true" box="[412,425,1047,1068]" pageId="11" pageNumber="36">±</emphasis>
1.75 23.78–27.67
</td>
<td id="280B5DD8C012AACFBD89FBA9BE3EFB9C" box="[508,630,1047,1098]" gridcol="2" gridrow="8" pageId="11" pageNumber="36">
11.47 
<emphasis id="E79EEAF4C012553BBE42FBA9BE0CFBFA" bold="true" box="[567,580,1047,1068]" pageId="11" pageNumber="36">±</emphasis>
0.19 11.21–11.68
</td>
<td id="280B5DD8C012AACFBEECFBA9BF5AFB9C" box="[665,786,1047,1098]" gridcol="3" gridrow="8" pageId="11" pageNumber="36">
10.28 
<emphasis id="E79EEAF4C012553BBEA1FBA9BEA9FBFA" bold="true" box="[724,737,1047,1068]" pageId="11" pageNumber="36">±</emphasis>
0.45 9.79–10.83
</td>
<td id="280B5DD8C012AACFBF40FBA9BFEAFB9C" box="[821,930,1047,1098]" gridcol="4" gridrow="8" pageId="11" pageNumber="36">
6.04 
<emphasis id="E79EEAF4C012553BBF10FBA9BF3AFBFA" bold="true" box="[869,882,1047,1068]" pageId="11" pageNumber="36">±</emphasis>
0.23 5.81–6.36
</td>
<td id="280B5DD8C012AACFBFA4FBA9B876FB9C" box="[977,1086,1047,1098]" gridcol="5" gridrow="8" pageId="11" pageNumber="36">
8.35 
<emphasis id="E79EEAF4C012553BB874FBA9B846FBFA" bold="true" box="[1025,1038,1047,1068]" pageId="11" pageNumber="36">±</emphasis>
0.52 7.74–8.97
</td>
<td id="280B5DD8C012AACFB818FBA9B892FB9C" box="[1133,1242,1047,1098]" gridcol="6" gridrow="8" pageId="11" pageNumber="36">
9.36 
<emphasis id="E79EEAF4C012553BB8E8FBA9B8E2FBFA" bold="true" box="[1181,1194,1047,1068]" pageId="11" pageNumber="36">±</emphasis>
0.37 9.15–9.92
</td>
<td id="280B5DD8C012AACFB97FFBA9B9D4FB9C" box="[1290,1436,1047,1098]" gridcol="7" gridrow="8" pageId="11" pageNumber="36">
2.54 
<emphasis id="E79EEAF4C012553BB94FFBA9B90FFBFA" bold="true" box="[1338,1351,1047,1068]" pageId="11" pageNumber="36">±</emphasis>
0.21 2.33–2.81
</td>
</tr>
<tr id="6BDA34A4C012AACFBCE2FBE3B9D4FB59" box="[151,1436,1117,1167]" gridrow="9" pageId="11" pageNumber="36">
<th id="280B5DD8C012AACFBCE2FBE3BD66FB59" box="[151,302,1117,1167]" gridcol="0" gridrow="9" pageId="11" pageNumber="36">
<taxonomicName id="12EA4D65C012553BBCEAFBE3BCBBFBA4" box="[159,243,1117,1138]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="11" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
(
<emphasis id="E79EEAF4C012553BBCD2FBC4BCFBFB59" box="[167,179,1146,1167]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 23)
</th>
<td id="280B5DD8C012AACFBD15FBE3BD91FB59" box="[352,473,1117,1167]" gridcol="1" gridrow="9" pageId="11" pageNumber="36">
27.43 
<emphasis id="E79EEAF4C012553BBDE9FBE3BDE1FBA4" bold="true" box="[412,425,1117,1138]" pageId="11" pageNumber="36">±</emphasis>
0.83 25.98–29.29
</td>
<td id="280B5DD8C012AACFBD89FBE3BE3EFB59" box="[508,630,1117,1167]" gridcol="2" gridrow="9" pageId="11" pageNumber="36">
12.03 
<emphasis id="E79EEAF4C012553BBE4DFBE3BE0DFBA4" bold="true" box="[568,581,1117,1138]" pageId="11" pageNumber="36">±</emphasis>
0.47 11.33–12.77
</td>
<td id="280B5DD8C012AACFBEECFBE3BF5AFB59" box="[665,786,1117,1167]" gridcol="3" gridrow="9" pageId="11" pageNumber="36">
11.29 
<emphasis id="E79EEAF4C012553BBEA1FBE3BEA9FBA4" bold="true" box="[724,737,1117,1138]" pageId="11" pageNumber="36">±</emphasis>
0.46 10.38–12.19
</td>
<td id="280B5DD8C012AACFBF40FBE3BFEAFB59" box="[821,930,1117,1167]" gridcol="4" gridrow="9" pageId="11" pageNumber="36">
6.42 
<emphasis id="E79EEAF4C012553BBF10FBE3BF3AFBA4" bold="true" box="[869,882,1117,1138]" pageId="11" pageNumber="36">±</emphasis>
0.25 5.90–6.84
</td>
<td id="280B5DD8C012AACFBFA4FBE3B876FB59" box="[977,1086,1117,1167]" gridcol="5" gridrow="9" pageId="11" pageNumber="36">
8.50 
<emphasis id="E79EEAF4C012553BB874FBE3B846FBA4" bold="true" box="[1025,1038,1117,1138]" pageId="11" pageNumber="36">±</emphasis>
0.36 7.91–9.45
</td>
<td id="280B5DD8C012AACFB818FBE3B892FB59" box="[1133,1242,1117,1167]" gridcol="6" gridrow="9" pageId="11" pageNumber="36">
8.80 
<emphasis id="E79EEAF4C012553BB8E8FBE3B8E2FBA4" bold="true" box="[1181,1194,1117,1138]" pageId="11" pageNumber="36">±</emphasis>
0.57 7.54–9.8
</td>
<td id="280B5DD8C012AACFB97FFBE3B9D4FB59" box="[1290,1436,1117,1167]" gridcol="7" gridrow="9" pageId="11" pageNumber="36">
2.68 
<emphasis id="E79EEAF4C012553BB94FFBE3B90FFBA4" bold="true" box="[1338,1351,1117,1138]" pageId="11" pageNumber="36">±</emphasis>
0.15 2.36–2.92
</td>
</tr>
</table>
</paragraph>
<paragraph id="D55536E6C012553BBCB0FB6BBFBAFA7C" blockId="11.[151,1437,1237,2010]" pageId="11" pageNumber="36">
The second component follows immediately and is a whine-like note. Each harmonic is a nearly pure tone with descending frequency (
<figureCitation id="4DD12A63C012553BBD91FB44BE64FAC2" box="[484,556,1274,1300]" captionStart="FIGURE 5" captionStartId="12.[151,258,1915,1939]" captionTargetBox="[154,1436,198,1899]" captionTargetId="figure@12.[151,1436,194,1903]" captionTargetPageId="12" captionText="FIGURE 5. Calls of Engystomops from western Ecuador. A – E: E. puyango (QCAZ 26968) from Bosque Protector Puyango, Provincia El Oro, Ecuador; F – J: E. pustulatus (QCAZ 26852) from El Empalme, Provincia Guayas, Ecuador. A and F are oscilograms; B and G spectrograms; C and H power spectra of complete call, D and I power spectra of the first component, E and J power spectra of the third component." httpUri="https://zenodo.org/record/197715/files/figure.png" pageId="11" pageNumber="36">Fig. 5</figureCitation>
). The average decrease in frequency of the first harmonic is 448 Hz (SD = 45.6, 
<emphasis id="E79EEAF4C012553BBC95FA9EBCA7FAEF" box="[224,239,1312,1337]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 10; measured to the end of the call). The average frequency at the beginning of the second component is 937.7 Hz (SD = 41.0, 
<emphasis id="E79EEAF4C012553BBE33FAFBBE1DFA88" box="[582,597,1349,1374]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 10) for the first harmonic and 1924 Hz (SD = 67.15, 
<emphasis id="E79EEAF4C012553BB89BFAFBB8B5FA88" box="[1262,1277,1349,1374]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 10) for the second. The first spectral peak had more energy than the second in calls of 
<specimenCount id="C3ECFD6FC012553BB842FAD4B8DDFA52" box="[1079,1173,1386,1412]" pageId="11" pageNumber="36" type="male">8 males</specimenCount>
out of 
<specimenCount id="C3ECFD6FC012553BB89AFAD4B929FA52" box="[1263,1377,1386,1412]" pageId="11" pageNumber="36" type="male">10 males</specimenCount>
. The number of visible harmonics in the spectrogram varies between 6 and 12.
</paragraph>
<paragraph id="D55536E6C012553BBCB0FA0BBF06F92C" blockId="11.[151,1437,1237,2010]" pageId="11" pageNumber="36">
A facultative third component can be added after the whine and is characterized by a gradual increase in amplitude and a switch in dominant frequency from the first to the fifth or sixth harmonic (
<figureCitation id="4DD12A63C012553BB8ACFA64B96AFA22" box="[1241,1314,1498,1524]" captionStart="FIGURE 5" captionStartId="12.[151,258,1915,1939]" captionTargetBox="[154,1436,198,1899]" captionTargetId="figure@12.[151,1436,194,1903]" captionTargetPageId="12" captionText="FIGURE 5. Calls of Engystomops from western Ecuador. A – E: E. puyango (QCAZ 26968) from Bosque Protector Puyango, Provincia El Oro, Ecuador; F – J: E. pustulatus (QCAZ 26852) from El Empalme, Provincia Guayas, Ecuador. A and F are oscilograms; B and G spectrograms; C and H power spectra of complete call, D and I power spectra of the first component, E and J power spectra of the third component." httpUri="https://zenodo.org/record/197715/files/figure.png" pageId="11" pageNumber="36">Fig. 5</figureCitation>
). Average fundamental frequency of the third component is 509 Hz (SD = 24.9, 
<emphasis id="E79EEAF4C012553BBF9BF9BEBFB5F9CF" box="[1006,1021,1536,1561]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 10; Table 4); average dominant frequency is 2851 Hz (SD = 135.5; 
<emphasis id="E79EEAF4C012553BBE24F99BBE28F9E8" box="[593,608,1573,1598]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 10). Infrequently, the third component exhibits a doubling in the number of harmonics (as shown in 
<figureCitation id="4DD12A63C012553BBE47F9F4BE3FF9B2" box="[562,631,1610,1636]" captionStart="FIGURE 5" captionStartId="12.[151,258,1915,1939]" captionTargetBox="[154,1436,198,1899]" captionTargetId="figure@12.[151,1436,194,1903]" captionTargetPageId="12" captionText="FIGURE 5. Calls of Engystomops from western Ecuador. A – E: E. puyango (QCAZ 26968) from Bosque Protector Puyango, Provincia El Oro, Ecuador; F – J: E. pustulatus (QCAZ 26852) from El Empalme, Provincia Guayas, Ecuador. A and F are oscilograms; B and G spectrograms; C and H power spectra of complete call, D and I power spectra of the first component, E and J power spectra of the third component." httpUri="https://zenodo.org/record/197715/files/figure.png" pageId="11" pageNumber="36">Fig. 5</figureCitation>
B). The majority of males may be capable of doubling the harmonics because in captivity we recorded repeatedly calls from eight males and all of them produced at least once a call with doubling. This 
<typeStatus id="0A518844C012553BBDBCF92BBDB6F979" box="[457,510,1685,1711]" pageId="11" pageNumber="36">type</typeStatus>
of call, however, was produced infrequently: out of 12 recordings of third components from seven captive males, only two presented doubling in the number of harmonics; out of ten males recorded at Bosque Protector 
<taxonomicName id="12EA4D65C012553BBE4AF95EBEE1F92C" box="[575,681,1760,1786]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="11" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
, only one did.
</paragraph>
<paragraph id="D55536E6C012553BBCB0F8BBBE70F80C" blockId="11.[151,1437,1237,2010]" pageId="11" pageNumber="36">
Observations at Bosque Protector 
<taxonomicName id="12EA4D65C012553BBE17F8BBBE82F8C9" box="[610,714,1797,1823]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="11" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
suggest that the third component is produced when another individual approaches the calling male. Our experiment to test the influence of the proximity of other individuals in the production of the third component shown a significant positive effect. The average proportion of calls with third component for accompanied males was 0.682 (SD = 0.1943, 
<emphasis id="E79EEAF4C012553BB8A9F8CBB8A3F858" box="[1244,1259,1909,1934]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 6) while the proportion for lonely males was 0.085 (SD = 0.140, 
<emphasis id="E79EEAF4C012553BBF74F825BF58F862" box="[769,784,1947,1972]" italics="true" pageId="11" pageNumber="36">n</emphasis>
= 6). Differences between proportions were significant (Wilcoxon’s 
<emphasis id="E79EEAF4C012553BBD5BF87EBD71F80F" box="[302,313,1984,2009]" italics="true" pageId="11" pageNumber="36">z</emphasis>
= -2.207; 
<emphasis id="E79EEAF4C012553BBDC1F87EBD8EF80F" box="[436,454,1984,2009]" italics="true" pageId="11" pageNumber="36">P</emphasis>
= 0.027).
</paragraph>
<caption id="8195666EC015553CBCE2F8C5BF70F824" httpUri="https://zenodo.org/record/197715/files/figure.png" pageId="12" pageNumber="37" targetBox="[154,1436,198,1899]" targetPageId="12">
<paragraph id="D55536E6C015553CBCE2F8C5BF70F824" blockId="12.[151,1437,1915,2034]" pageId="12" pageNumber="37">
<emphasis id="E79EEAF4C015553CBCE2F8C5BD56F845" bold="true" box="[151,286,1915,1939]" pageId="12" pageNumber="37">FIGURE 5.</emphasis>
Calls of 
<taxonomicName id="12EA4D65C015553CBDF0F8C5BE50F844" box="[389,536,1915,1938]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="12" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C015553CBDF0F8C5BE50F844" box="[389,536,1915,1938]" italics="true" pageId="12" pageNumber="37">Engystomops</emphasis>
</taxonomicName>
from western Ecuador. A–E: 
<taxonomicName id="12EA4D65C015553CBF12F8C5BFADF844" box="[871,997,1915,1938]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="12" pageNumber="37" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C015553CBF12F8C5BFADF844" box="[871,997,1915,1938]" italics="true" pageId="12" pageNumber="37">E. puyango</emphasis>
</taxonomicName>
(QCAZ 26968) from Bosque Protector 
<taxonomicName id="12EA4D65C015553CBCE2F825BCBFF864" box="[151,247,1947,1970]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="12" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
, Provincia El Oro, Ecuador; F–J: 
<taxonomicName id="12EA4D65C015553CBE11F825BEA7F864" box="[612,751,1947,1970]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="12" pageNumber="37" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C015553CBE11F825BEA7F864" box="[612,751,1947,1970]" italics="true" pageId="12" pageNumber="37">E. pustulatus</emphasis>
</taxonomicName>
(QCAZ 26852) from El Empalme, Provincia Guayas, Ecuador. A and F are oscilograms; B and G spectrograms; C and H power spectra of complete call, D and I power spectra of the first component, E and J power spectra of the third component.
</paragraph>
</caption>
<caption id="8195666EC014553DBCE2F9BDBF57F98C" httpUri="https://zenodo.org/record/197716/files/figure.png" pageId="13" pageNumber="38" targetBox="[321,1261,185,1519]" targetPageId="13">
<paragraph id="D55536E6C014553DBCE2F9BDBF57F98C" blockId="13.[151,1437,1539,1626]" pageId="13" pageNumber="38">
<emphasis id="E79EEAF4C014553DBCE2F9BDBD54F9CD" bold="true" box="[151,284,1539,1563]" pageId="13" pageNumber="38">FIGURE 6.</emphasis>
Dorsal and ventral views of adult 
<taxonomicName id="12EA4D65C014553DBEE1F9BDBFC0F9CC" box="[660,904,1539,1562]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C014553DBEE1F9BDBFC0F9CC" box="[660,904,1539,1562]" italics="true" pageId="13" pageNumber="38">Engystomops puyango</emphasis>
</taxonomicName>
showing variation in dorsal and ventral patterns. From left to right, above: QCAZ 26976, 26982–83, 26986; below: QCAZ 27009, 27013, 37276 (all males). Provincia Loja and El Oro, Ecuador (See Appendix I for locality data).
</paragraph>
</caption>
<paragraph id="D55536E6C014553DBCB0F936B818F8EE" blockId="13.[151,1437,1672,2034]" pageId="13" pageNumber="38">
Two components with eigenvalues&gt; 1.0 were extracted from the PCA of calls from 
<specimenCount id="C3ECFD6FC014553DB892F936B91EF974" box="[1255,1366,1672,1698]" pageId="13" pageNumber="38" type="male">49 males</specimenCount>
of 
<taxonomicName id="12EA4D65C014553DB9F7F937BCB5F911" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C014553DB9F7F937BCB5F911" italics="true" pageId="13" pageNumber="38">E. puyango</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C014553DBD4DF910BD9CF911" box="[312,468,1710,1735]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C014553DBD4DF910BD9CF911" box="[312,468,1710,1735]" italics="true" pageId="13" pageNumber="38">E. pustulatus</emphasis>
</taxonomicName>
. The two PCs accounted for 76.6% of the total variation. Principal Component I loads mostly on frequency variables while PC II on call duration (
<tableCitation id="9868035DC014553DBFEBF96DBFB2F93B" box="[926,1018,1747,1773]" captionStart="TABLE 2" captionStartId="2.[151,231,1233,1254]" captionTargetBox="[151,1437,1360,1828]" captionTargetPageId="2" captionText="TABLE 2. Character loading and percentage of explained variance for Principal Components (PC) I – II for eight acoustic variables from the advertisement call of Engystomops puyango and E. pustulatus. Bold figures indicate highest loadings. Variable Character Loading" httpUri="http://table.plazi.org/id/8195666EC01B5532BCE2FB6FB806FAEA" pageId="13" pageNumber="38" tableUuid="8195666EC01B5532BCE2FB6FB806FAEA">Table 2</tableCitation>
). The acoustic space of 
<taxonomicName id="12EA4D65C014553DB963F96DB9D4F93A" box="[1302,1436,1747,1772]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C014553DB963F96DB9D4F93A" box="[1302,1436,1747,1772]" italics="true" pageId="13" pageNumber="38">E. puyango</emphasis>
</taxonomicName>
is distinctive from that of 
<taxonomicName id="12EA4D65C014553DBDB3F947BE17F8C4" box="[454,607,1785,1810]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C014553DBDB3F947BE17F8C4" box="[454,607,1785,1810]" italics="true" pageId="13" pageNumber="38">E. pustulatus</emphasis>
</taxonomicName>
as shown by significant differences between both species in scores for PC I (
<emphasis id="E79EEAF4C014553DBCA8F8A0BCADF8E1" box="[221,229,1822,1847]" italics="true" pageId="13" pageNumber="38">t</emphasis>
= 5.34, 
<emphasis id="E79EEAF4C014553DBD33F8A0BD15F8E1" box="[326,349,1822,1847]" italics="true" pageId="13" pageNumber="38">df</emphasis>
= 32, 
<emphasis id="E79EEAF4C014553DBDDDF8A0BDF2F8E1" box="[424,442,1822,1847]" italics="true" pageId="13" pageNumber="38">P</emphasis>
&lt;0.001) and PC II (
<emphasis id="E79EEAF4C014553DBED8F8A0BEFDF8E1" box="[685,693,1822,1847]" italics="true" pageId="13" pageNumber="38">t</emphasis>
= 6.34, 
<emphasis id="E79EEAF4C014553DBF62F8A0BF65F8E1" box="[791,813,1822,1847]" italics="true" pageId="13" pageNumber="38">df</emphasis>
= 13, 
<emphasis id="E79EEAF4C014553DBF0CF8A0BFC3F8E1" box="[889,907,1822,1847]" italics="true" pageId="13" pageNumber="38">P</emphasis>
&lt;0.001; 
<figureCitation id="4DD12A63C014553DBF8EF8A0B80AF8EE" box="[1019,1090,1822,1848]" captionStart="FIGURE 4" captionStartId="9.[151,255,983,1007]" captionTargetBox="[373,1217,189,958]" captionTargetId="figure@9.[369,1218,180,962]" captionTargetPageId="9" captionText="FIGURE 4. Axes I and II from Principal Components Analysis based on eight acoustic variables from the advertisement calls of Engystomops puyango (10 males) and E. pustulatus (39). See Table 2 for character loadings on each component." httpUri="https://zenodo.org/record/197714/files/figure.png" pageId="13" pageNumber="38">Fig. 4</figureCitation>
).
</paragraph>
<paragraph id="D55536E6C014553DBCB0F8FDB8EFF87E" blockId="13.[151,1437,1672,2034]" pageId="13" pageNumber="38">
In the DFA classification procedure, calls from all 
<specimenCount id="C3ECFD6FC014553DBF69F8FDBFCEF88B" box="[796,902,1859,1885]" pageId="13" pageNumber="38" type="male">10 males</specimenCount>
of 
<taxonomicName id="12EA4D65C014553DBFD8F8FDB87AF88A" box="[941,1074,1859,1884]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C014553DBFD8F8FDB87AF88A" box="[941,1074,1859,1884]" italics="true" pageId="13" pageNumber="38">E. puyango</emphasis>
</taxonomicName>
were correctly classified as 
<taxonomicName id="12EA4D65C014553DB9F6F8FDBCB4F854" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C014553DB9F6F8FDBCB4F854" italics="true" pageId="13" pageNumber="38">E. puyango</emphasis>
</taxonomicName>
and calls from all 
<specimenCount id="C3ECFD6FC014553DBDA1F8D6BE75F854" box="[468,573,1896,1922]" pageId="13" pageNumber="38" type="male">29 males</specimenCount>
of 
<taxonomicName id="12EA4D65C014553DBE17F8D7BEB3F854" box="[610,763,1897,1922]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C014553DBE17F8D7BEB3F854" box="[610,763,1897,1922]" italics="true" pageId="13" pageNumber="38">E. pustulatus</emphasis>
</taxonomicName>
were classified as 
<taxonomicName id="12EA4D65C014553DBFA3F8D7B826F854" box="[982,1134,1897,1922]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C014553DBFA3F8D7B826F854" box="[982,1134,1897,1922]" italics="true" pageId="13" pageNumber="38">E. pustulatus</emphasis>
</taxonomicName>
. The absence of incorrect assignments confirms the high acoustic distinctiveness between the calls of both species.
</paragraph>
<paragraph id="D55536E6C014553EBCB0F80DB864FE61" blockId="13.[151,1437,1672,2034]" lastBlockId="14.[151,1436,152,626]" lastPageId="14" lastPageNumber="39" pageId="13" pageNumber="38">
<emphasis id="E79EEAF4C014553DBCB0F80DBE61F81B" bold="true" box="[197,553,1971,1997]" pageId="13" pageNumber="38">Morphometric comparisons.</emphasis>
Three components with eigenvalues&gt; 1.0 were extracted from the PCA of 
<specimenCount id="C3ECFD6FC014553DBCE2F866BD7DF824" box="[151,309,2008,2034]" pageId="13" pageNumber="38" type="generic">30 specimens</specimenCount>
of 
<taxonomicName id="12EA4D65C014553DBD2EF866BD97F827" box="[347,479,2008,2033]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C014553DBD2EF866BD97F827" box="[347,479,2008,2033]" italics="true" pageId="13" pageNumber="38">E. puyango</emphasis>
</taxonomicName>
, 51 of 
<taxonomicName id="12EA4D65C014553DBE44F866BE81F827" box="[561,713,2008,2033]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="13" pageNumber="38" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C014553DBE44F866BE81F827" box="[561,713,2008,2033]" italics="true" pageId="13" pageNumber="38">E. pustulatus</emphasis>
</taxonomicName>
, and 11 of 
<emphasis id="E79EEAF4C014553DBF3EF866BF15F827" box="[843,861,2008,2033]" italics="true" pageId="13" pageNumber="38">E</emphasis>
. sp. B. The three PCs accounted for 76.1% of the total variation. The highest loadings were head width and femur length for PC I, arm length for PC II, and head length for PC III (
<tableCitation id="9868035DC017553EBDB6FF03BE6EFF01" box="[451,550,189,215]" captionStart="TABLE 6" captionStartId="15.[151,235,153,174]" captionTargetBox="[151,1437,277,599]" captionTargetPageId="15" captionText="TABLE 6. Character loading and percentage of explained variance for Principal Components (PC) I – II for six morphometric variables. Bold figures indicate highest loadings." httpUri="http://table.plazi.org/id/8195666EC016553FBCE2FF27BF76FF19" pageId="14" pageNumber="39" tableUuid="8195666EC016553FBCE2FF27BF76FF19">Table 6</tableCitation>
). The morphometric space of 
<taxonomicName id="12EA4D65C017553EBFD2FF03B87FFF00" box="[935,1079,189,214]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C017553EBFD2FF03B87FFF00" box="[935,1079,189,214]" italics="true" pageId="14" pageNumber="39">E. puyango</emphasis>
</taxonomicName>
overlaps with 
<taxonomicName id="12EA4D65C017553EB883FF03B9D4FF00" box="[1270,1436,189,214]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C017553EB883FF03B9D4FF00" box="[1270,1436,189,214]" italics="true" pageId="14" pageNumber="39">E. pustulatus</emphasis>
</taxonomicName>
populations from Arenillas and Huaquillas (
<figureCitation id="4DD12A63C017553EBEEBFF5CBEACFF2A" box="[670,740,226,252]" captionStart="FIGURE 8" captionStartId="15.[151,255,1682,1706]" captionTargetBox="[254,1323,685,1631]" captionTargetId="figure@15.[245,1342,677,1661]" captionTargetPageId="15" captionText="FIGURE 8. Axes I and II from Principal Components Analysis based on six size-corrected morphological variables for Engystomops puyango (30 specimens), E. pustulatus (51), and E. sp. B (11). See Table 6 for character loadings on each component." httpUri="https://zenodo.org/record/197718/files/figure.png" pageId="14" pageNumber="39">Fig. 8</figureCitation>
) but only slightly with 
<taxonomicName id="12EA4D65C017553EBF8EFF5CB8DDFF2D" box="[1019,1173,226,251]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C017553EBF8EFF5CB8DDFF2D" box="[1019,1173,226,251]" italics="true" pageId="14" pageNumber="39">E. pustulatus</emphasis>
</taxonomicName>
from 
<collectingCountry id="ADFD7676C017553EB8ABFF5CB921FF2A" box="[1246,1385,226,252]" name="Puerto Rico" pageId="14" pageNumber="39">Puerto Rico</collectingCountry>
and Patricia Pilar. In contrast, 
<emphasis id="E79EEAF4C017553EBDB8FEB6BDAEFEF7" box="[461,486,264,289]" italics="true" pageId="14" pageNumber="39">E.</emphasis>
sp. B overlaps widely in morphometric space with 
<taxonomicName id="12EA4D65C017553EB838FEB6B89CFEF7" box="[1101,1236,264,289]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C017553EB838FEB6B89CFEF7" box="[1101,1236,264,289]" italics="true" pageId="14" pageNumber="39">E. puyango</emphasis>
</taxonomicName>
as shown by the lack of significant differences along both PCs (PC I: 
<emphasis id="E79EEAF4C017553EBF70FE93BF45FE90" box="[773,781,301,326]" italics="true" pageId="14" pageNumber="39">t</emphasis>
= 0.44, df = 23, 
<emphasis id="E79EEAF4C017553EBFA6FE93BFADFE90" box="[979,997,301,326]" italics="true" pageId="14" pageNumber="39">P</emphasis>
= 0.661; PC II: 
<emphasis id="E79EEAF4C017553EB8D1FE93B8E4FE90" box="[1188,1196,301,326]" italics="true" pageId="14" pageNumber="39">t</emphasis>
= 0.55, df = 15, 
<emphasis id="E79EEAF4C017553EB907FE93B9CCFE90" box="[1394,1412,301,326]" italics="true" pageId="14" pageNumber="39">P</emphasis>
= 0.587). Comparisons between 
<taxonomicName id="12EA4D65C017553EBE77FEECBEC2FEBD" box="[514,650,338,363]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C017553EBE77FEECBEC2FEBD" box="[514,650,338,363]" italics="true" pageId="14" pageNumber="39">E. puyango</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C017553EBEB0FEECBF29FEBD" box="[709,865,338,363]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C017553EBEB0FEECBF29FEBD" box="[709,865,338,363]" italics="true" pageId="14" pageNumber="39">E. pustulatus</emphasis>
</taxonomicName>
of the variables with the highest loadings show significant differences in residual head width (
<emphasis id="E79EEAF4C017553EBECDFEC6BE88FE47" box="[696,704,376,401]" italics="true" pageId="14" pageNumber="39">t</emphasis>
= 2.20, df = 65, 
<emphasis id="E79EEAF4C017553EBFF3FEC6BFD0FE47" box="[902,920,376,401]" italics="true" pageId="14" pageNumber="39">P</emphasis>
= 0.03) and residual femur length (
<emphasis id="E79EEAF4C017553EB94FFEC6B90AFE47" box="[1338,1346,376,401]" italics="true" pageId="14" pageNumber="39">t</emphasis>
= 2.45, df = 78, 
<emphasis id="E79EEAF4C017553EBC8FFE23BD44FE60" box="[250,268,413,438]" italics="true" pageId="14" pageNumber="39">P</emphasis>
= 0.016) but not in residual arm length (
<emphasis id="E79EEAF4C017553EBE9CFE23BEB9FE60" box="[745,753,413,438]" italics="true" pageId="14" pageNumber="39">t</emphasis>
= 1.38, df = 65, 
<emphasis id="E79EEAF4C017553EBFC3FE23BF80FE60" box="[950,968,413,438]" italics="true" pageId="14" pageNumber="39">P</emphasis>
= 0.17).
</paragraph>
<paragraph id="D55536E6C017553EBCB0FE7CBE07FDA4" blockId="14.[151,1436,152,626]" pageId="14" pageNumber="39">
In the DFA classification procedure, 16 out of 
<specimenCount id="C3ECFD6FC017553EBE9EFE7CBFC3FE0A" box="[747,907,450,476]" pageId="14" pageNumber="39" type="generic">30 specimens</specimenCount>
of 
<taxonomicName id="12EA4D65C017553EBFC6FE7CB870FE0D" box="[947,1080,450,475]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C017553EBFC6FE7CB870FE0D" box="[947,1080,450,475]" italics="true" pageId="14" pageNumber="39">E. puyango</emphasis>
</taxonomicName>
were classified correctly. The misclassified specimens were assigned to 
<taxonomicName id="12EA4D65C017553EBEF6FE56BF54FDD7" box="[643,796,488,513]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C017553EBEF6FE56BF54FDD7" box="[643,796,488,513]" italics="true" pageId="14" pageNumber="39">E. pustulatus</emphasis>
</taxonomicName>
(four specimens) and 
<emphasis id="E79EEAF4C017553EB856FE56B874FDD7" box="[1059,1084,488,513]" italics="true" pageId="14" pageNumber="39">E.</emphasis>
sp. B (
<specimenCount id="C3ECFD6FC017553EB8E5FE56B97CFDD4" box="[1168,1332,488,514]" pageId="14" pageNumber="39" type="generic">10 specimens</specimenCount>
). Six out of 11 
<emphasis id="E79EEAF4C017553EBCA9FDB3BCBEFDF0" box="[220,246,525,550]" italics="true" pageId="14" pageNumber="39">E.</emphasis>
sp. B were correctly classified; all misclassified specimens were assigned 
<taxonomicName id="12EA4D65C017553EB802FDB3B8B7FDF0" box="[1143,1279,525,550]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C017553EB802FDB3B8B7FDF0" box="[1143,1279,525,550]" italics="true" pageId="14" pageNumber="39">E. puyango</emphasis>
</taxonomicName>
. Overall, the multivariate analyses indicate some morphometric differentiation between 
<taxonomicName id="12EA4D65C017553EB864FD8CB8DFFD9D" box="[1041,1175,562,587]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C017553EB864FD8CB8DFFD9D" box="[1041,1175,562,587]" italics="true" pageId="14" pageNumber="39">E. puyango</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C017553EB8A6FD8CB926FD9D" box="[1235,1390,562,587]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C017553EB8A6FD8CB926FD9D" box="[1235,1390,562,587]" italics="true" pageId="14" pageNumber="39">E. pustulatus</emphasis>
</taxonomicName>
but not between 
<taxonomicName id="12EA4D65C017553EBD59FDE6BDFAFDA7" box="[300,434,600,625]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C017553EBD59FDE6BDFAFDA7" box="[300,434,600,625]" italics="true" pageId="14" pageNumber="39">E. puyango</emphasis>
</taxonomicName>
and 
<emphasis id="E79EEAF4C017553EBD9EFDE6BE4CFDA7" box="[491,516,600,625]" italics="true" pageId="14" pageNumber="39">E.</emphasis>
sp. B.
</paragraph>
<caption id="8195666EC017553EBCE2F802BE24F822" httpUri="https://zenodo.org/record/197717/files/figure.png" pageId="14" pageNumber="39" targetBox="[204,1382,674,1959]" targetPageId="14">
<paragraph id="D55536E6C017553EBCE2F802BE24F822" blockId="14.[151,1436,1980,2036]" pageId="14" pageNumber="39">
<emphasis id="E79EEAF4C017553EBCE2F802BD52F802" bold="true" box="[151,282,1980,2004]" pageId="14" pageNumber="39">FIGURE 7.</emphasis>
Ventral views of the right hand and foot of 
<taxonomicName id="12EA4D65C017553EBE80F803BFAFF802" box="[757,999,1981,2004]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="39" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C017553EBE80F803BFAFF802" box="[757,999,1981,2004]" italics="true" pageId="14" pageNumber="39">Engystomops puyango</emphasis>
</taxonomicName>
, QCAZ 26975 (adult male from Bosque Protector 
<taxonomicName id="12EA4D65C017553EBC8AF863BD17F822" box="[255,351,2013,2036]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="14" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
, Ecuador; SVL = 28.16).
</paragraph>
</caption>
<caption id="8195666EC016553FBCE2FF27BF76FF19" ID-Table-UUID="8195666EC016553FBCE2FF27BF76FF19" httpUri="http://table.plazi.org/id/8195666EC016553FBCE2FF27BF76FF19" pageId="15" pageNumber="40" targetBox="[151,1437,277,599]" targetIsTable="true" targetPageId="15">
<paragraph id="D55536E6C016553FBCE2FF27BF76FF19" blockId="15.[151,1437,151,257]" pageId="15" pageNumber="40">
<emphasis id="E79EEAF4C016553FBCE2FF27BD42FF79" bold="true" box="[151,266,151,175]" pageId="15" pageNumber="40">TABLE 6.</emphasis>
Character loading and percentage of explained variance for Principal Components (PC) I–II for six morphometric variables. Bold figures indicate highest loadings.
</paragraph>
</caption>
<paragraph id="D55536E6C016553FBCEAFF54B8C8FED7" blockId="15.[151,1437,151,257]" box="[159,1152,234,257]" pageId="15" pageNumber="40">Variable Size-free morphology</paragraph>
</subSubSection>
<paragraph id="D55536E6C016553FBF44FEABB8B1FD81" pageId="15" pageNumber="40">
<table id="A7EAC446C016AACFBCE2FEABB9D5FD81" box="[151,1437,277,599]" gridcols="3" gridrows="4" pageId="15" pageNumber="40">
<tr id="6BDA34A4C016AACFBCE2FEABB9D5FEFA" box="[151,1437,277,300]" gridrow="0" pageId="15" pageNumber="40" rowspan-0="1">
<th id="280B5DD8C016AACFBF5EFEABBF2FFEFA" box="[811,871,277,300]" gridcol="1" gridrow="0" pageId="15" pageNumber="40">PC I</th>
<th id="280B5DD8C016AACFB8C3FEABB9D5FEFA" box="[1206,1437,277,300]" gridcol="2" gridrow="0" pageId="15" pageNumber="40">PC II</th>
</tr>
<tr id="6BDA34A4C016AACFBCE2FEFEB9D5FE7A" box="[151,1437,320,428]" gridrow="1" pageId="15" pageNumber="40">
<th id="280B5DD8C016AACFBCE2FEFEBD3BFE7A" box="[151,371,320,428]" gridcol="0" gridrow="1" pageId="15" pageNumber="40">Tibia length Femur length Arm length</th>
<td id="280B5DD8C016AACFBF5EFEFEBF2FFE7A" box="[811,871,320,428]" gridcol="1" gridrow="1" pageId="15" pageNumber="40">
0.413 
<emphasis id="E79EEAF4C016553FBF5EFED4BF2FFE54" bold="true" box="[811,871,362,386]" pageId="15" pageNumber="40">0.526</emphasis>
0.293
</td>
<td id="280B5DD8C016AACFB8C3FEFEB9D5FE7A" box="[1206,1437,320,428]" gridcol="2" gridrow="1" pageId="15" pageNumber="40">
0.409 –0.128 
<emphasis id="E79EEAF4C016553FB8C8FE2AB8B1FE7A" bold="true" box="[1213,1273,404,428]" pageId="15" pageNumber="40">0.664</emphasis>
</td>
</tr>
<tr id="6BDA34A4C016AACFBCE2FE7EB9D5FDFA" box="[151,1437,448,556]" gridrow="2" pageId="15" pageNumber="40">
<th id="280B5DD8C016AACFBCE2FE7EBD3BFDFA" box="[151,371,448,556]" gridcol="0" gridrow="2" pageId="15" pageNumber="40">Head length Head width Eye-nostril distance</th>
<td id="280B5DD8C016AACFBF5EFE7EBF2FFDFA" box="[811,871,448,556]" gridcol="1" gridrow="2" pageId="15" pageNumber="40">
0.249 
<emphasis id="E79EEAF4C016553FBF5EFE54BF2FFDD4" bold="true" box="[811,871,490,514]" pageId="15" pageNumber="40">0.543</emphasis>
0.328
</td>
<td id="280B5DD8C016AACFB8C3FE7EB9D5FDFA" box="[1206,1437,448,556]" gridcol="2" gridrow="2" pageId="15" pageNumber="40">
0.073 –0.242 
<emphasis id="E79EEAF4C016553FB8C3FDAAB948FDFA" bold="true" box="[1206,1280,532,556]" pageId="15" pageNumber="40">–0.557</emphasis>
</td>
</tr>
<tr id="6BDA34A4C016AACFBCE2FDFEB9D5FD81" box="[151,1437,576,599]" gridrow="3" pageId="15" pageNumber="40">
<th id="280B5DD8C016AACFBCE2FDFEBD3BFD81" box="[151,371,576,599]" gridcol="0" gridrow="3" pageId="15" pageNumber="40">Eigenvalue</th>
<td id="280B5DD8C016AACFBF5EFDFEBF2FFD81" box="[811,871,576,599]" gridcol="1" gridrow="3" pageId="15" pageNumber="40">2.233</td>
<td id="280B5DD8C016AACFB8C3FDFEB9D5FD81" box="[1206,1437,576,599]" gridcol="2" gridrow="3" pageId="15" pageNumber="40">1.315</td>
</tr>
</table>
</paragraph>
<caption id="8195666EC016553FBCE2F92CBD5DF93F" httpUri="https://zenodo.org/record/197718/files/figure.png" pageId="15" pageNumber="40" targetBox="[254,1323,685,1631]" targetPageId="15">
<paragraph id="D55536E6C016553FBCE2F92CBD5DF93F" blockId="15.[151,1436,1682,1769]" pageId="15" pageNumber="40">
<emphasis id="E79EEAF4C016553FBCE2F92CBD51F97C" bold="true" box="[151,281,1682,1706]" pageId="15" pageNumber="40">FIGURE 8.</emphasis>
Axes I and II from Principal Components Analysis based on six size-corrected morphological variables for 
<taxonomicName id="12EA4D65C016553FBCE2F90CBDC0F91F" box="[151,392,1714,1737]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="15" pageNumber="40" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C016553FBCE2F90CBDC0F91F" box="[151,392,1714,1737]" italics="true" pageId="15" pageNumber="40">Engystomops puyango</emphasis>
</taxonomicName>
(30 specimens), 
<taxonomicName id="12EA4D65C016553FBE34F90CBE85F91F" box="[577,717,1714,1737]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="15" pageNumber="40" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C016553FBE34F90CBE85F91F" box="[577,717,1714,1737]" italics="true" pageId="15" pageNumber="40">E. pustulatus</emphasis>
</taxonomicName>
(51), and 
<emphasis id="E79EEAF4C016553FBF49F90CBF04F91F" box="[828,844,1714,1737]" italics="true" pageId="15" pageNumber="40">E</emphasis>
. sp. B (11). See Table 6 for character loadings on each component.
</paragraph>
</caption>
<subSubSection id="9DF0656DC0165521BCB0F8A9BD24FF01" lastPageId="17" lastPageNumber="42" pageId="15" pageNumber="40" type="distribution">
<paragraph id="D55536E6C016553FBCB0F8A9BF15F877" blockId="15.[151,1437,1815,2028]" pageId="15" pageNumber="40">
<emphasis id="E79EEAF4C016553FBCB0F8A9BE48F8E7" bold="true" box="[197,512,1815,1841]" pageId="15" pageNumber="40">Distribution and ecology.</emphasis>
<taxonomicName id="12EA4D65C016553FBE7DF8A6BF59F8E7" box="[520,785,1816,1841]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="15" pageNumber="40" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C016553FBE7DF8A6BF59F8E7" box="[520,785,1816,1841]" italics="true" pageId="15" pageNumber="40">Engystomops puyango</emphasis>
</taxonomicName>
has been recorded in southwestern 
<collectingCountry id="ADFD7676C016553FB8CDF8A9B952F8E7" box="[1208,1306,1815,1841]" name="Ecuador" pageId="15" pageNumber="40">Ecuador</collectingCountry>
(Provincia Loja and Provincia El Oro) between 
<quantity id="12129B03C016553FBE26F883BEA2F881" box="[595,746,1853,1879]" metricMagnitude="2" metricUnit="m" metricValue="8.055" metricValueMax="12.91" metricValueMin="3.2" pageId="15" pageNumber="40" unit="m" value="805.5" valueMax="1291.0" valueMin="320.0">320–1291 m</quantity>
above sea level (
<figureCitation id="4DD12A63C016553FBFCEF883B84EF881" box="[955,1030,1853,1879]" captionStart="FIGURE 9" captionStartId="16.[151,257,1725,1749]" captionTargetBox="[328,1258,387,1691]" captionTargetId="figure@16.[318,1269,381,1705]" captionTargetPageId="16" captionText="FIGURE 9. Known records of Engystomops puyango (circles) and E. pustulatus (solid triangles for northern range, hollow triangles for the southern range). Locality data are based on specimens deposited in, California Academy of Sciences, Museo de Zoología de la Pontificia Universidad Católica del Ecuador, and Museum of Comparative Zoology Harvard University (Appendix I)." httpUri="https://zenodo.org/record/197719/files/figure.png" pageId="15" pageNumber="40">Fig. 9</figureCitation>
). Maximum straight-line distance between localities is 
<quantity id="12129B03C016553FBDE9F8DCBDB6F8AA" box="[412,510,1890,1916]" metricMagnitude="5" metricUnit="m" metricValue="1.2" pageId="15" pageNumber="40" unit="km" value="120.0">120 km</quantity>
. The occurrence of 
<taxonomicName id="12EA4D65C016553FBE80F8DCBFCAF8AD" box="[757,898,1890,1915]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="15" pageNumber="40" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C016553FBE80F8DCBFCAF8AD" box="[757,898,1890,1915]" italics="true" pageId="15" pageNumber="40">E. puyango</emphasis>
</taxonomicName>
in 
<collectingCountry id="ADFD7676C016553FBFD9F8DCBFACF8AA" box="[940,996,1890,1916]" name="Peru" pageId="15" pageNumber="40">Peru</collectingCountry>
is highly probable because Bosque Protector 
<taxonomicName id="12EA4D65C016553FBD7CF839BD27F877" box="[265,367,1927,1953]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="15" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
and Zapotillo are in the border with 
<collectingCountry id="ADFD7676C016553FBF6AF839BF11F877" box="[799,857,1927,1953]" name="Peru" pageId="15" pageNumber="40">Peru</collectingCountry>
.
</paragraph>
<paragraph id="D55536E6C0165520BCB0F813B858FE91" blockId="15.[151,1437,1815,2028]" lastBlockId="16.[151,1436,152,327]" lastPageId="16" lastPageNumber="41" pageId="15" pageNumber="40">
The geographic range of 
<taxonomicName id="12EA4D65C016553FBD9FF813BE27F810" box="[490,623,1965,1990]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="15" pageNumber="40" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C016553FBD9FF813BE27F810" box="[490,623,1965,1990]" italics="true" pageId="15" pageNumber="40">E. puyango</emphasis>
</taxonomicName>
is characterized by scant and highly seasonal precipitation. The rainy season lasts between February and April (
<bibRefCitation id="B17B4B17C016553FBEE8F86CBFA8F83A" author="Lynch" box="[669,992,2002,2028]" pageId="15" pageNumber="40" refString="Lynch, J. D. &amp; Duellman, W. E. (1997) Frogs of the genus Eleutherodactylus in Western Ecuador. Special Publication The University of Kansas Natural History Museum, 23, 1 - 236." type="journal article" year="1997">Lynch &amp; Duellman, 1997</bibRefCitation>
). Localities are in Evergreen Lower Montane Forest of the Western Andes (Catamayo) and Foothill Semideciduous Costa Forest (
<taxonomicName id="12EA4D65C0095520B953FF26B9DFFF64" box="[1318,1431,152,178]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="16" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
, Mangaurquillo, and Zapotillo; vegetation 
<typeStatus id="0A518844C0095520BEF1FF03BE8AFF01" box="[644,706,189,215]" pageId="16" pageNumber="41">types</typeStatus>
are as defined by Sierra, 
<emphasis id="E79EEAF4C0095520BF85FF03B863FF00" box="[1008,1067,189,214]" italics="true" pageId="16" pageNumber="41">et al.</emphasis>
, 1999 and Cerón, 
<emphasis id="E79EEAF4C0095520B973FF03B909FF00" box="[1286,1345,189,214]" italics="true" pageId="16" pageNumber="41">et al.</emphasis>
, 1999). The Evergreen Lower Montane Forest is a transitional form between humid forest and the dry forests that predominates in the Andes of southern 
<collectingCountry id="ADFD7676C0095520BE16FEB6BE8FFEF4" box="[611,711,264,290]" name="Ecuador" pageId="16" pageNumber="41">Ecuador</collectingCountry>
. The Foothill Semideciduous Costa Forest has scattered trees, less than 
<quantity id="12129B03C0095520BD71FE93BD77FE91" box="[260,319,301,327]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="16" pageNumber="41" unit="m" value="20.0">20 m</quantity>
tall, and a dense understory dominated by herbaceous plants.
</paragraph>
<caption id="8195666EC0095520BCE2F903BDB5F8E3" httpUri="https://zenodo.org/record/197719/files/figure.png" pageId="16" pageNumber="41" targetBox="[328,1258,387,1691]" targetPageId="16">
<paragraph id="D55536E6C0095520BCE2F903BDB5F8E3" blockId="16.[151,1436,1725,1845]" pageId="16" pageNumber="41">
<emphasis id="E79EEAF4C0095520BCE2F903BD55F903" bold="true" box="[151,285,1725,1749]" pageId="16" pageNumber="41">FIGURE 9.</emphasis>
Known records of 
<taxonomicName id="12EA4D65C0095520BD81F900BEA4F903" box="[500,748,1726,1749]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="16" pageNumber="41" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0095520BD81F900BEA4F903" box="[500,748,1726,1749]" italics="true" pageId="16" pageNumber="41">Engystomops puyango</emphasis>
</taxonomicName>
(circles) and 
<taxonomicName id="12EA4D65C0095520BFF0F900B85EF903" box="[901,1046,1726,1749]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="16" pageNumber="41" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C0095520BFF0F900B85EF903" box="[901,1046,1726,1749]" italics="true" pageId="16" pageNumber="41">E. pustulatus</emphasis>
</taxonomicName>
(solid triangles for northern range, hollow triangles for the southern range). Locality data are based on specimens deposited in, California Academy of Sciences, Museo de Zoología de la Pontificia Universidad Católica del Ecuador, and Museum of Comparative Zoology Harvard University (Appendix I).
</paragraph>
</caption>
<paragraph id="D55536E6C0095521BCB0F8DDBD24FF01" blockId="16.[151,1437,1891,2029]" lastBlockId="17.[151,1437,152,738]" lastPageId="17" lastPageNumber="42" pageId="16" pageNumber="41">
All individuals were found in open to sparsely forested areas where the original natural vegetation has been partly or completely removed by humans. At Bosque Protector 
<taxonomicName id="12EA4D65C0095520BF8FF837B822F875" box="[1018,1130,1929,1955]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="16" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
, choruses were found in 
<date id="A1541026C0095520BCE2F810BD0DF81E" box="[151,325,1966,1992]" pageId="16" pageNumber="41" value="2004-02">February 2004</date>
and 
<date id="A1541026C0095520BDF4F810BE7BF81E" box="[385,563,1966,1992]" pageId="16" pageNumber="41" value="2005-02">February 2005</date>
. Males were calling from water in ponds and ditches. Amplexus and egg deposition take place at the same sites where choruses call. 
<taxonomicName id="12EA4D65C0095520BF11F86DB839F83A" box="[868,1137,2003,2028]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="16" pageNumber="41" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0095520BF11F86DB839F83A" box="[868,1137,2003,2028]" italics="true" pageId="16" pageNumber="41">Engystomops puyango</emphasis>
</taxonomicName>
constructs floating foam nests during amplexus. While the female discharges the egg masses, the male beats them with his legs to produce the foam.
</paragraph>
</subSubSection>
<subSubSection id="9DF0656DC0085522BCB0FF5CBF59FE44" lastPageId="18" lastPageNumber="43" pageId="17" pageNumber="42" type="reference_group">
<paragraph id="D55536E6C0085521BCB0FF5CBDD1FD34" blockId="17.[151,1437,152,738]" pageId="17" pageNumber="42">
Males 
<taxonomicName id="12EA4D65C0085521BD6DFF5CBDEDFF2D" box="[280,421,226,251]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0085521BD6DFF5CBDEDFF2D" box="[280,421,226,251]" italics="true" pageId="17" pageNumber="42">E. puyango</emphasis>
</taxonomicName>
and the smaller 
<taxonomicName id="12EA4D65C0085521BE01FF5CBE92FF2D" box="[628,730,226,252]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C0085521BE01FF5CBECEFF2D" box="[628,646,226,251]" italics="true" pageId="17" pageNumber="42">E</emphasis>
. 
<emphasis id="E79EEAF4C0085521BEE2FF5CBE92FF2D" box="[663,730,226,251]" italics="true" pageId="17" pageNumber="42">randi</emphasis>
</taxonomicName>
were frequently calling at the same ponds, sometimes at distances less than 
<quantity id="12129B03C0085521BD0BFEB6BD84FEF4" box="[382,460,264,290]" metricMagnitude="-1" metricUnit="m" metricValue="2.0" pageId="17" pageNumber="42" unit="cm" value="20.0">20 cm</quantity>
. At a dense chorus of both species at Bosque Protector 
<taxonomicName id="12EA4D65C0085521B819FEB6B890FEF4" box="[1132,1240,264,290]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="49" phylum="Chordata" rank="species" species="puyango">Puyango</taxonomicName>
, on 
<date id="A1541026C0085521B97FFEB6BC9DFE91" pageId="17" pageNumber="42" value="2005-02-19">February 19 2005</date>
, we found a male (QCAZ 28801) with an unusual advertisement call. Average call duration (
<geoCoordinate id="B0DE5021C0085521B967FE93B921FE91" box="[1298,1385,301,327]" direction="south" orientation="latitude" pageId="17" pageNumber="42" precision="55" value="-0.222">0.222 s</geoCoordinate>
, SD = 0.006, 
<emphasis id="E79EEAF4C0085521BD74FEECBD58FEBD" box="[257,272,338,363]" italics="true" pageId="17" pageNumber="42">n</emphasis>
= 10) was below the range for 
<taxonomicName id="12EA4D65C0085521BEF1FEECBF43FEBD" box="[644,779,338,363]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0085521BEF1FEECBF43FEBD" box="[644,779,338,363]" italics="true" pageId="17" pageNumber="42">E. puyango</emphasis>
</taxonomicName>
(Table 4) and close to the average of 
<taxonomicName id="12EA4D65C0085521B8B9FEECB966FEBD" box="[1228,1326,338,363]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C0085521B8B9FEECB966FEBD" box="[1228,1326,338,363]" italics="true" pageId="17" pageNumber="42">E. randi</emphasis>
</taxonomicName>
(
<geoCoordinate id="B0DE5021C0085521B948FEECB9DEFEBA" box="[1341,1430,338,364]" direction="south" orientation="latitude" pageId="17" pageNumber="42" precision="55" value="-0.206">0.206 s</geoCoordinate>
, SD = 0.013, based on recordings from 
<specimenCount id="C3ECFD6FC0085521BE13FEC6BE8AFE44" box="[614,706,376,402]" pageId="17" pageNumber="42" type="male">7 males</specimenCount>
from the same locality). Call interval was closer to that of 
<taxonomicName id="12EA4D65C0085521B9F6FEC6BC9FFE60" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C0085521B9F6FEC6BC9FFE60" italics="true" pageId="17" pageNumber="42">E. randi</emphasis>
</taxonomicName>
(QCAZ 28801 mean = 0.203, SD = 0.011, 
<emphasis id="E79EEAF4C0085521BEA8FE23BEA4FE60" box="[733,748,413,438]" italics="true" pageId="17" pageNumber="42">n</emphasis>
= 10 calls; 
<taxonomicName id="12EA4D65C0085521BF0CFE23BF93FE60" box="[889,987,413,438]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C0085521BF0CFE23BF93FE60" box="[889,987,413,438]" italics="true" pageId="17" pageNumber="42">E. randi</emphasis>
</taxonomicName>
= 0.324, SD = 0.091, 
<emphasis id="E79EEAF4C0085521B892FE23B8BEFE60" box="[1255,1270,413,438]" italics="true" pageId="17" pageNumber="42">n</emphasis>
= 
<specimenCount id="C3ECFD6FC0085521B963FE23B93EFE61" box="[1302,1398,413,439]" pageId="17" pageNumber="42" type="male">7 males</specimenCount>
; 
<taxonomicName id="12EA4D65C0085521B9F6FE23BCB6FE0D" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0085521B9F6FE23BCB6FE0D" italics="true" pageId="17" pageNumber="42">E. puyango</emphasis>
</taxonomicName>
= 1.083, SD = 0.146, 
<emphasis id="E79EEAF4C0085521BE7EFE7CBE52FE0D" box="[523,538,450,475]" italics="true" pageId="17" pageNumber="42">n</emphasis>
= 
<specimenCount id="C3ECFD6FC0085521BE4EFE7CBEE4FE0A" box="[571,684,450,476]" pageId="17" pageNumber="42" type="male">10 males</specimenCount>
). However, similarly to 
<taxonomicName id="12EA4D65C0085521BFB8FE7CB81DFE0D" box="[973,1109,450,475]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0085521BFB8FE7CB81DFE0D" box="[973,1109,450,475]" italics="true" pageId="17" pageNumber="42">E. puyango</emphasis>
</taxonomicName>
, the first component of the call lacked the well-defined pulses present in calls of 
<taxonomicName id="12EA4D65C0085521BF7BFE56B877FDD4" authority="Ron et al. 2004" authorityName="Ron et al." authorityYear="2004" box="[782,1087,488,514]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C0085521BF7BFE56BF27FDD7" box="[782,879,488,513]" italics="true" pageId="17" pageNumber="42">E. randi</emphasis>
(
<bibRefCitation id="B17B4B17C0085521BF0BFE56B87FFDD4" author="Ron" box="[894,1079,488,514]" pageId="17" pageNumber="42" refString="Ron, S. R., Cannatella, D. C. &amp; Coloma, L. A. (2004) Two new species of Physalaemus (Anura: Leptodactylidae) from western Ecuador. Herpetologica, 60, 261 - 275." type="journal article" year="2004">
Ron 
<emphasis id="E79EEAF4C0085521BFCDFE56BFA4FDD7" box="[952,1004,488,513]" italics="true" pageId="17" pageNumber="42">et al</emphasis>
. 2004
</bibRefCitation>
)
</taxonomicName>
and its SVL (
<quantity id="12129B03C0085521B891FE56B917FDD4" box="[1252,1375,488,514]" metricMagnitude="-2" metricUnit="m" metricValue="2.831" pageId="17" pageNumber="42" unit="mm" value="28.31">28.31 mm</quantity>
) was outside the range of adult males of 
<taxonomicName id="12EA4D65C0085521BE32FDB3BEE3FDF0" box="[583,683,525,550]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C0085521BE32FDB3BEE3FDF0" box="[583,683,525,550]" italics="true" pageId="17" pageNumber="42">E. randi</emphasis>
</taxonomicName>
(20.09–24.00 mm, 
<emphasis id="E79EEAF4C0085521BFE9FDB3BFE3FDF0" box="[924,939,525,550]" italics="true" pageId="17" pageNumber="42">n</emphasis>
= 24). Because the characteristics of the advertisement call of QCAZ 28801 are intermediate between 
<taxonomicName id="12EA4D65C0085521BF19FD8CBFBAFD9D" box="[876,1010,562,587]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0085521BF19FD8CBFBAFD9D" box="[876,1010,562,587]" italics="true" pageId="17" pageNumber="42">E. puyango</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C0085521B85EFD8CB8C3FD9D" box="[1067,1163,562,587]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C0085521B85EFD8CB8C3FD9D" box="[1067,1163,562,587]" italics="true" pageId="17" pageNumber="42">E. randi</emphasis>
</taxonomicName>
, we hypothesize that it is a hybrid individual. A sequence of 750 bases of mtDNA, gene 16S, is identical to that of 
<taxonomicName id="12EA4D65C0085521B897FDE6B90CFDA7" box="[1250,1348,600,625]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C0085521B897FDE6B90CFDA7" box="[1250,1348,600,625]" italics="true" pageId="17" pageNumber="42">E. randi</emphasis>
</taxonomicName>
QCAZ 23768 from the same population. Thus, the hybrid could be the product of the cross of a female 
<taxonomicName id="12EA4D65C0085521B957FDC3B9CDFD40" box="[1314,1413,637,662]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="randi">
<emphasis id="E79EEAF4C0085521B957FDC3B9CDFD40" box="[1314,1413,637,662]" italics="true" pageId="17" pageNumber="42">E. randi</emphasis>
</taxonomicName>
x male 
<taxonomicName id="12EA4D65C0085521BCA9FD1CBD23FD6D" box="[220,363,674,699]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0085521BCA9FD1CBD23FD6D" box="[220,363,674,699]" italics="true" pageId="17" pageNumber="42">E. puyango</emphasis>
</taxonomicName>
. Other species calling in syntopy with 
<taxonomicName id="12EA4D65C0085521BF2FFD1CBFA1FD6D" box="[858,1001,674,699]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0085521BF2FFD1CBFA1FD6D" box="[858,1001,674,699]" italics="true" pageId="17" pageNumber="42">E. puyango</emphasis>
</taxonomicName>
were 
<taxonomicName id="12EA4D65C0085521B842FD1CB92FFD6D" ID-CoL="4VJYQ" box="[1079,1383,674,699]" class="Amphibia" family="Hylidae" genus="Scinax" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="quinquefasciatus">
<emphasis id="E79EEAF4C0085521B842FD1CB92FFD6D" box="[1079,1383,674,699]" italics="true" pageId="17" pageNumber="42">Scinax quinquefasciatus</emphasis>
</taxonomicName>
and 
<taxonomicName id="12EA4D65C0085521BCE2FD76BDDAFD37" box="[151,402,712,737]" class="Amphibia" family="Hylidae" genus="Phrynohyas" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="venulosa">
<emphasis id="E79EEAF4C0085521BCE2FD76BDDAFD37" box="[151,402,712,737]" italics="true" pageId="17" pageNumber="42">Phrynohyas venulosa</emphasis>
</taxonomicName>
.
</paragraph>
<caption id="8195666EC0085521BCE2FA9EBF9CFA81" httpUri="https://zenodo.org/record/197720/files/figure.png" pageId="17" pageNumber="42" targetBox="[156,1432,802,1272]" targetPageId="17">
<paragraph id="D55536E6C0085521BCE2FA9EBF9CFA81" blockId="17.[151,1436,1312,1367]" pageId="17" pageNumber="42">
<emphasis id="E79EEAF4C0085521BCE2FA9EBD62FAEE" bold="true" box="[151,298,1312,1336]" pageId="17" pageNumber="42">FIGURE 10.</emphasis>
Karyotype of 
<taxonomicName id="12EA4D65C0085521BDBFFA9EBE88FAE1" box="[458,704,1312,1335]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C0085521BDBFFA9EBE88FAE1" box="[458,704,1312,1335]" italics="true" pageId="17" pageNumber="42">Engystomops puyango</emphasis>
</taxonomicName>
. (A) Giemsa stained, (B) Ag-NOR stained, (C) C-banded. Arrows indicate the secondary constriction (A), Ag-NOR (B), C-banded in pair 9 (C).
</paragraph>
</caption>
<paragraph id="D55536E6C0085521BCB0FA3BBFF4F823" blockId="17.[151,1437,1413,2037]" pageId="17" pageNumber="42">
<emphasis id="E79EEAF4C0085521BCB0FA3BBD3BFA49" bold="true" box="[197,371,1413,1439]" pageId="17" pageNumber="42">Cytogenetics.</emphasis>
All eight analyzed individuals had a chromosome number of 2 
<emphasis id="E79EEAF4C0085521B80AFA38B8C6FA49" box="[1151,1166,1414,1439]" italics="true" pageId="17" pageNumber="42">n</emphasis>
= 20. This is the first known departure from the 2 
<emphasis id="E79EEAF4C0085521BE74FA15BE58FA12" box="[513,528,1451,1476]" italics="true" pageId="17" pageNumber="42">n</emphasis>
= 22 karyotype characteristic of 
<taxonomicName id="12EA4D65C0085521BFB6FA15B82AFA13" box="[963,1122,1451,1477]" class="Amphibia" family="Leiuperidae" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="family">Leiuperidae</taxonomicName>
. All species analyzed of 
<emphasis id="E79EEAF4C0085521BCE2FA6FBD56FA3C" box="[151,286,1489,1514]" italics="true" pageId="17" pageNumber="42">Eladorhina</emphasis>
, 
<taxonomicName id="12EA4D65C0085521BD58FA6FBDFAFA3C" box="[301,434,1489,1514]" class="Amphibia" family="Leiuperidae" genus="Eupemphix" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C0085521BD58FA6FBDFAFA3C" box="[301,434,1489,1514]" italics="true" pageId="17" pageNumber="42">Eupemphix</emphasis>
</taxonomicName>
, 
<taxonomicName id="12EA4D65C0085521BDB6FA6FBE1BFA3C" box="[451,595,1489,1514]" class="Amphibia" family="Leiuperidae" genus="Pleurodema" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C0085521BDB6FA6FBE1BFA3C" box="[451,595,1489,1514]" italics="true" pageId="17" pageNumber="42">Pleurodema</emphasis>
</taxonomicName>
, 
<taxonomicName id="12EA4D65C0085521BE16FA6FBF72FA3C" box="[611,826,1489,1514]" class="Amphibia" family="Leiuperidae" genus="Pseudopaludicola" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C0085521BE16FA6FBF72FA3C" box="[611,826,1489,1514]" italics="true" pageId="17" pageNumber="42">Pseudopaludicola</emphasis>
</taxonomicName>
, 
<taxonomicName id="12EA4D65C0085521BF3CFA6FBFAEFA3C" box="[841,998,1489,1514]" class="Amphibia" family="Leiuperidae" genus="Physalaemus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C0085521BF3CFA6FBFAEFA3C" box="[841,998,1489,1514]" italics="true" pageId="17" pageNumber="42">Physalaemus</emphasis>
</taxonomicName>
, and 
<taxonomicName id="12EA4D65C0085521B85DFA6FB88FFA3C" box="[1064,1223,1489,1514]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="17" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="E79EEAF4C0085521B85DFA6FB88FFA3C" box="[1064,1223,1489,1514]" italics="true" pageId="17" pageNumber="42">Engystomops</emphasis>
</taxonomicName>
have a karyotype 2 
<emphasis id="E79EEAF4C0085521BCD0FA48BCFCF9D9" box="[165,180,1526,1551]" italics="true" pageId="17" pageNumber="42">n</emphasis>
= 22 (e.g., 
<bibRefCitation id="B17B4B17C0085521BD4CFA48BE66F9C6" author="Ananias" box="[313,558,1526,1552]" pageId="17" pageNumber="42" refString="Ananias, F., Bombeiro, A. L., Silva, C. D., Silva, P., Haddad, C. F. B. &amp; Kasahara, S. (2007) Cytogenetics of Eupemphix nattereri Steindachner, 1863 (Anura: Leiuperidae) and karyotypic similarity with species of related genera: taxonomic implications. Acta Zoologica Sinica, 53, 285 - 293." type="journal article" year="2007">
Ananias, 
<emphasis id="E79EEAF4C0085521BDDDFA48BDAAF9D9" box="[424,482,1526,1551]" italics="true" pageId="17" pageNumber="42">et al.</emphasis>
, 2007
</bibRefCitation>
; 
<bibRefCitation id="B17B4B17C0085521BE4FFA48BEB2F9C6" author="Duellman" box="[570,762,1526,1552]" pageId="17" pageNumber="42" refString="Duellman, W. E. (1967) Additional studies of chromosomes of anuran amphibians. Systematic Zoology, 16, 38 - 43." type="journal article" year="1967">Duellman, 1967</bibRefCitation>
; 
<bibRefCitation id="B17B4B17C0085521BF73FA48B843F9C6" author="Lourenco" box="[774,1035,1526,1552]" pageId="17" pageNumber="42" refString="Lourenco, L. B., Cardoso, A. J. &amp; Recco-Pimentel, S. M. (2000) Cytogenetics of Edalorhina perezi (Anura, Leptodactylidae). Cytologia, 65, 359 - 363." type="journal article" year="2000">
Lourenço, 
<emphasis id="E79EEAF4C0085521BFF1FA48BFF7F9D9" box="[900,959,1526,1551]" italics="true" pageId="17" pageNumber="42">et al.</emphasis>
, 2000
</bibRefCitation>
; 
<bibRefCitation id="B17B4B17C0085521B862FA48B954F9C6" author="Lourenco" box="[1047,1308,1526,1552]" pageId="17" pageNumber="42" refString="Lourenco, L. B., Nascimento, A. A., Andrade, G. V., Rossa-Feres, D. C. &amp; Recco-Pimentel, S. M. (2006) Chromosomal analysis of the leptodactylids Pleurodema diplolistris and Physalaemus nattereri. Amphibia-Reptilia, 27, 481 - 489." type="journal article" year="2006">
Lourenço, 
<emphasis id="E79EEAF4C0085521B8E0FA48B898F9D9" box="[1173,1232,1526,1551]" italics="true" pageId="17" pageNumber="42">et al.</emphasis>
, 2006
</bibRefCitation>
; 
<bibRefCitation id="B17B4B17C0085521B95DFA48BD6AF9E3" author="Quindere" pageId="17" pageNumber="42" refString="Quindere, Y. R., Lourenco, L. B., Andrade, G. V., Tomatis, C., Baldo, D. &amp; Recco-Pimentel, S. M. (2009) Polytypic and polymorphic cytogenetic variations in the widespread anuran Physalaemus cuvieri (Anura, Leiuperidae) with emphasis on nucleolar organizing regions. Biological Research, 42, 79 - 92." type="journal article" year="2009">
Quindere, 
<emphasis id="E79EEAF4C0085521BCE2F9A5BC9BF9E2" box="[151,211,1563,1588]" italics="true" pageId="17" pageNumber="42">et al.</emphasis>
, 2009
</bibRefCitation>
; Tomatis, 
<emphasis id="E79EEAF4C0085521BDE9F9A5BD91F9E2" box="[412,473,1563,1588]" italics="true" pageId="17" pageNumber="42">et al.</emphasis>
, 2009). Chromosome pairs 1, 5, 6, 7 and 10 were metacentric while pairs 2, 3, 4 and 8 were submetacentric (
<tableCitation id="9868035DC0085521BD89F9FEBE17F98C" box="[508,607,1600,1626]" captionStart="TABLE 3" captionStartId="6.[151,231,153,174]" captionTargetBox="[151,1436,423,863]" captionTargetPageId="6" captionText="TABLE 3. Morphometry of karyotypes of Engystomops puyango and E. pustulatus. Means and standard deviations are from 25 metaphases from three males (QCAZ 47296, 46932, 47295) and five females (QCAZ 35690 – 91, 46934, 47297, SC 26360) of E. puyango from Bosque Protector Puyango, Ecuador; and four of E. pustulatus (from Cerro Blanco, Ecuador; QCAZ 47031 – 33, QCAZ 46931). Abbreviations are: m = metacentric; sm = submetacentric." httpUri="http://table.plazi.org/id/8195666EC01F5536BCE2FF27B8D9FEC0" pageId="17" pageNumber="42" tableUuid="8195666EC01F5536BCE2FF27B8D9FEC0">Table 3</tableCitation>
, 
<figureCitation id="4DD12A63C0085521BE1AF9FEBE85F98C" box="[623,717,1600,1626]" captionStart="FIGURE 10" captionStartId="17.[151,256,1312,1336]" captionTargetBox="[156,1432,802,1272]" captionTargetId="figure@17.[151,1436,791,1290]" captionTargetPageId="17" captionText="FIGURE 10. Karyotype of Engystomops puyango. (A) Giemsa stained, (B) Ag-NOR stained, (C) C-banded. Arrows indicate the secondary constriction (A), Ag-NOR (B), C-banded in pair 9 (C)." httpUri="https://zenodo.org/record/197720/files/figure.png" pageId="17" pageNumber="42">Fig. 10</figureCitation>
). Pair 9 was heteromorphic with one chromosome being submetacentric and the other metacentric and slightly smaller. An extended secondary constriction was detected with Giemsa staining in the long arm of two chromosomes 9 (
<figureCitation id="4DD12A63C0085521BFA0F935B862F973" box="[981,1066,1675,1701]" captionStart="FIGURE 10" captionStartId="17.[151,256,1312,1336]" captionTargetBox="[156,1432,802,1272]" captionTargetId="figure@17.[151,1436,791,1290]" captionTargetPageId="17" captionText="FIGURE 10. Karyotype of Engystomops puyango. (A) Giemsa stained, (B) Ag-NOR stained, (C) C-banded. Arrows indicate the secondary constriction (A), Ag-NOR (B), C-banded in pair 9 (C)." httpUri="https://zenodo.org/record/197720/files/figure.png" pageId="17" pageNumber="42">Fig. 10</figureCitation>
A); using the Ag-NOR method, it was determined that the constriction corresponds to the nucleolar organizing region (NOR) (
<figureCitation id="4DD12A63C0085521B898F90EB90DF91C" box="[1261,1349,1712,1738]" captionStart="FIGURE 10" captionStartId="17.[151,256,1312,1336]" captionTargetBox="[156,1432,802,1272]" captionTargetId="figure@17.[151,1436,791,1290]" captionTargetPageId="17" captionText="FIGURE 10. Karyotype of Engystomops puyango. (A) Giemsa stained, (B) Ag-NOR stained, (C) C-banded. Arrows indicate the secondary constriction (A), Ag-NOR (B), C-banded in pair 9 (C)." httpUri="https://zenodo.org/record/197720/files/figure.png" pageId="17" pageNumber="42">Fig. 10</figureCitation>
B). The difference in size and morphology of pair 9 is a result of the size of NOR: in the submetacentric chromosome 9 the NOR is a large block that doubles the size of the NOR of the metacentric chromosome 9. This heteromorphism in the size of the NOR may be due to varying amounts of its transcription units associated with events of duplication and/or deletion of NOR constituents; NORs are made of highly repetitive ribosomal genes that allow such genetic events. It is also possible that the NOR heteromorphism could result from other mechanisms such as unequal sister chromatid exchange or non-peer linkage in meiosis (
<bibRefCitation id="B17B4B17C0085521B953F82EBC92F806" author="Kasahara" pageId="17" pageNumber="42" refString="Kasahara, S. (2009) Introducao a pesquisa em citogenetica de Vertebrados. Sociedade Brasileira de Genetica, Ribeirao Preto." type="book" year="2009">Kasahara, 2009</bibRefCitation>
). It could also be the result of differential activity of rDNA transcription, which could explain the existence of inter-cellular polymorphism present in some specimens.
</paragraph>
<paragraph id="D55536E6C00B5522BCB0FF26BFDCFEF4" blockId="18.[151,1437,152,402]" pageId="18" pageNumber="43">
C-banding revealed small blocks of heterochromatin in the centromere of all chromosomes. C-positive bands were observed in the telomeres of the long and short arms of pairs 1 and 5, in the long arm of pairs 3, 6 and 8. Faint intercalated C-bands were observed in the long arms of pairs 1 and 8 (
<figureCitation id="4DD12A63C00B5522B8F5FF5CB891FF2A" box="[1152,1241,226,252]" captionStart="FIGURE 10" captionStartId="17.[151,256,1312,1336]" captionTargetBox="[156,1432,802,1272]" captionTargetId="figure@17.[151,1436,791,1290]" captionTargetPageId="17" captionText="FIGURE 10. Karyotype of Engystomops puyango. (A) Giemsa stained, (B) Ag-NOR stained, (C) C-banded. Arrows indicate the secondary constriction (A), Ag-NOR (B), C-banded in pair 9 (C)." httpUri="https://zenodo.org/record/197720/files/figure.png" pageId="18" pageNumber="43">Fig. 10</figureCitation>
C). Pair 9 had C bands in the internal and external borders of the NOR (
<figureCitation id="4DD12A63C00B5522BF69FEB6BF3AFEF4" box="[796,882,264,290]" captionStart="FIGURE 10" captionStartId="17.[151,256,1312,1336]" captionTargetBox="[156,1432,802,1272]" captionTargetId="figure@17.[151,1436,791,1290]" captionTargetPageId="17" captionText="FIGURE 10. Karyotype of Engystomops puyango. (A) Giemsa stained, (B) Ag-NOR stained, (C) C-banded. Arrows indicate the secondary constriction (A), Ag-NOR (B), C-banded in pair 9 (C)." httpUri="https://zenodo.org/record/197720/files/figure.png" pageId="18" pageNumber="43">Fig. 10</figureCitation>
C).
</paragraph>
<paragraph id="D55536E6C00B5522BCB0FE93BF59FE44" blockId="18.[151,1437,152,402]" pageId="18" pageNumber="43">
The karyotype of 
<taxonomicName id="12EA4D65C00B5522BDE2FE93BEE8FE90" box="[407,672,301,326]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="18" pageNumber="43" phylum="Chordata" rank="species" species="puyango">
<emphasis id="E79EEAF4C00B5522BDE2FE93BEE8FE90" box="[407,672,301,326]" italics="true" pageId="18" pageNumber="43">Engystomops puyango</emphasis>
</taxonomicName>
shares with 
<taxonomicName id="12EA4D65C00B5522BF40FE93BF86FE90" box="[821,974,301,326]" class="Amphibia" family="Leiuperidae" genus="Engystomops" kingdom="Animalia" order="Anura" pageId="18" pageNumber="43" phylum="Chordata" rank="species" species="pustulatus">
<emphasis id="E79EEAF4C00B5522BF40FE93BF86FE90" box="[821,974,301,326]" italics="true" pageId="18" pageNumber="43">E. pustulatus</emphasis>
</taxonomicName>
the same number of chromosomes and the position of the NOR in pair 9 (M. Rivera &amp; J. P. Falconí, unpublished). However, there are morphological differences in chromosome pairs 4, 5, and 6 (
<tableCitation id="9868035DC00B5522BED2FEC6BF48FE44" box="[679,768,376,402]" captionStart="TABLE 3" captionStartId="6.[151,231,153,174]" captionTargetBox="[151,1436,423,863]" captionTargetPageId="6" captionText="TABLE 3. Morphometry of karyotypes of Engystomops puyango and E. pustulatus. Means and standard deviations are from 25 metaphases from three males (QCAZ 47296, 46932, 47295) and five females (QCAZ 35690 – 91, 46934, 47297, SC 26360) of E. puyango from Bosque Protector Puyango, Ecuador; and four of E. pustulatus (from Cerro Blanco, Ecuador; QCAZ 47031 – 33, QCAZ 46931). Abbreviations are: m = metacentric; sm = submetacentric." httpUri="http://table.plazi.org/id/8195666EC01F5536BCE2FF27B8D9FEC0" pageId="18" pageNumber="43" tableUuid="8195666EC01F5536BCE2FF27B8D9FEC0">Table 3</tableCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>