treatments-xml/data/03/99/FA/0399FA75FFE12367FF4DFA12FF68DB47.xml
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<document id="F07DC9A102514EA1EDF59D77CAFDDCB6" ID-ISSN="1175­5334" IM.materialsCitations_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1629710095404" checkinUser="felipe" docAuthor="Páez-Vacas, Mónica I., Coloma, Luis A. &amp; Santos, Juan C." docDate="2010" docId="0399FA75FFE12367FF4DFA12FF68DB47" docLanguage="en" docName="zt02711p075.pdf" docOrigin="Zootaxa 2711" docStyle="DocumentStyle:D239614CE4198176A422035174489AB1.4:Zootaxa.2009-2012.monograph" docStyleId="D239614CE4198176A422035174489AB1" docStyleName="Zootaxa.2009-2012.monograph" docStyleVersion="4" docTitle="Hyloxalus sauli" docType="treatment" docVersion="3" lastPageNumber="47" masterDocId="FFA0820DFFCD2349FFDBFFECFF84D935" masterDocTitle="Systematics of the Hyloxalus bocagei complex (Anura: Dendrobatidae), description of two new cryptic species, and recognition of H. maculosus 2711" masterLastPageNumber="75" masterPageNumber="1" pageNumber="45" updateTime="1699099446680" updateUser="plazi" zenodo-license-document="CLOSED">
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<mods:title id="6F9466304EA37F1D3E2B19F19F38201B">Systematics of the Hyloxalus bocagei complex (Anura: Dendrobatidae), description of two new cryptic species, and recognition of H. maculosus 2711</mods:title>
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<mods:namePart id="4648AA60DF00096A5473D8650B135410">Páez-Vacas, Mónica I.</mods:namePart>
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<mods:namePart id="7E3FB1C404AE3F452B1E0C6B30799BCA">Coloma, Luis A.</mods:namePart>
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<mods:namePart id="440A6D3B62E1F5000E7D13E8323AF1B2">Santos, Juan C.</mods:namePart>
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<mods:title id="B71DC2C3B8A65256B1B1F88D0AFC35EA">Zootaxa</mods:title>
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<mods:number id="7CA61D9AC504EB8C386A894B1079994B">2711</mods:number>
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<subSubSection id="C32A18E8FFE12365FF4DFA12FDA5DF2D" box="[150,545,1534,1560]" pageId="44" pageNumber="45" type="nomenclature">
<paragraph id="8B8F4B63FFE12365FF4DFA12FDA5DF2D" blockId="44.[150,545,1534,1560]" box="[150,545,1534,1560]" pageId="44" pageNumber="45">
<heading id="D0C7FC0FFFE12365FF4DFA12FDA5DF2D" bold="true" box="[150,545,1534,1560]" fontSize="11" level="1" pageId="44" pageNumber="45" reason="1">
<taxonomicName id="4C3030E0FFE12365FF4DFA12FDA5DF2D" ID-CoL="3NGYB" authority="(Edwards 1974)" baseAuthorityName="Edwards" baseAuthorityYear="1974" box="[150,545,1534,1560]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="sauli">
<emphasis id="B9449771FFE12365FF4DFA12FDA5DF2D" bold="true" box="[150,545,1534,1560]" pageId="44" pageNumber="45">
<emphasis id="B9449771FFE12365FF4DFA12FED0DF2D" bold="true" box="[150,340,1534,1560]" italics="true" pageId="44" pageNumber="45">Hyloxalus sauli</emphasis>
(Edwards 1974)
</emphasis>
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="C32A18E8FFE12365FF4CF9A5FDC7DF95" pageId="44" pageNumber="45" type="reference_group">
<paragraph id="8B8F4B63FFE12365FF4CF9A5FCA1DFB5" blockId="44.[151,1436,1609,1696]" pageId="44" pageNumber="45">
<treatmentCitationGroup id="AB206C4DFFE12365FF4CF9A5FCA1DFB5" pageId="44" pageNumber="45">
<emphasis id="B9449771FFE12365FF4CF9A5FEE8DF55" box="[151,364,1609,1632]" italics="true" pageId="44" pageNumber="45">
<taxonomicName id="4C3030E0FFE12365FF4CF9A5FED6DF55" ID-CoL="X8GR" authorityName="Edwards" authorityYear="1974" box="[151,338,1609,1632]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="sauli">Colostethus sauli</taxonomicName>
</emphasis>
<materialsCitation id="3B58413EFFE12365FEB7F9A5FCA1DFB5" accessionNumber="KU 122217" collectingDate="1968-07-12" collectionCode="KU" collectorName="William G. Saul" country="Ecuador" elevation="340" location="Napo" pageId="44" pageNumber="45" specimenCount="1" stateProvince="Napo" typeStatus="holotype">
<treatmentCitation id="0A916D72FFE12365FEB7F9A5FDB3DF55" author="Edwards, S. R." box="[364,567,1609,1632]" page="10" pageId="44" pageNumber="45" year="1974">
<bibRefCitation id="EFA13692FFE12365FEB7F9A5FDB3DF55" author="Edwards, S. R." box="[364,567,1609,1632]" pageId="44" pageNumber="45" pagination="1 - 14" refId="ref45111" refString="Edwards, S. R. (1974 b) Taxonomic notes on South American dendrobatid frogs of the genus Colostethus. University of Kansas, Natural History Museum, Occasional Papers, 30, 1 - 14." type="journal article" year="1974">Edwards 1974b:10</bibRefCitation>
</treatmentCitation>
.—
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: “
<collectionCode id="ED21D3A6FFE12365FD07F9A5FC80DF55" box="[732,772,1609,1632]" country="USA" name="Biodiversity Institute, University of Kansas" pageId="44" pageNumber="45" type="University or college">KU</collectionCode>
122217, an adult male from Santa Cecilia,
<quantity id="4CC8E686FFE12365FADBF9A5FACFDF55" box="[1280,1355,1609,1632]" metricMagnitude="2" metricUnit="m" metricValue="3.4" pageId="44" pageNumber="45" unit="m" value="340.0">
<elevation id="001DAC50FFE12365FADBF9A5FACFDF55" box="[1280,1355,1609,1632]" metricMagnitude="2" metricUnit="m" metricValue="3.4" pageId="44" pageNumber="45" unit="m" value="340.0">340 m</elevation>
</quantity>
,
<collectingRegion id="49F48581FFE12365FA80F9A5FA13DF55" box="[1371,1431,1609,1632]" country="Ecuador" name="Napo" pageId="44" pageNumber="45">Napo</collectingRegion>
,
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; collected by
<collectorName id="26C52EB5FFE12365FE6DF985FDDBDFB5" box="[438,607,1641,1664]" pageId="44" pageNumber="45">William G. Saul</collectorName>
on
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<collectingDate id="EFCA944BFFE12365FD53F985FC92DFB5" box="[648,790,1641,1664]" pageId="44" pageNumber="45" value="1968-07-12">12 July 1968</collectingDate>
</date>
”.
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</paragraph>
<paragraph id="8B8F4B63FFE12365FF4CF965FDC7DF95" blockId="44.[151,1436,1609,1696]" box="[151,579,1673,1696]" pageId="44" pageNumber="45">
<treatmentCitationGroup id="AB206C4DFFE12365FF4CF965FDC7DF95" box="[151,579,1673,1696]" pageId="44" pageNumber="45">
<taxonomicName id="4C3030E0FFE12365FF4CF965FEBADF95" ID-CoL="3NGYB" baseAuthorityName="Edwards" baseAuthorityYear="1974" box="[151,318,1673,1696]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="sauli">
<emphasis id="B9449771FFE12365FF4CF965FEBADF95" box="[151,318,1673,1696]" italics="true" pageId="44" pageNumber="45">Hyloxalus sauli</emphasis>
</taxonomicName>
<treatmentCitation id="0A916D72FFE12365FE83F965FDBBDF95" author="Grant, T. &amp; Frost, D. R. &amp; Caldwell, J. P. &amp; Gagliardo, R. &amp; Haddad, C. F. B. &amp; Kok, P. J. R. &amp; Means, D. B. &amp; Noonan, B. P. &amp; Schargel, W. E. &amp; Wheeler, W. C." box="[344,575,1673,1696]" page="168" pageId="44" pageNumber="45" year="2006">
<bibRefCitation id="EFA13692FFE12365FE83F965FDBBDF95" author="Grant, T. &amp; Frost, D. R. &amp; Caldwell, J. P. &amp; Gagliardo, R. &amp; Haddad, C. F. B. &amp; Kok, P. J. R. &amp; Means, D. B. &amp; Noonan, B. P. &amp; Schargel, W. E. &amp; Wheeler, W. C." box="[344,575,1673,1696]" pageId="44" pageNumber="45" pagination="1 - 262" refId="ref45590" refString="Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F. B., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. &amp; Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262." type="journal article" year="2006">
Grant
<emphasis id="B9449771FFE12365FE47F965FE55DF95" box="[412,465,1673,1696]" italics="true" pageId="44" pageNumber="45">et al.</emphasis>
2006:168
</bibRefCitation>
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.
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</paragraph>
</subSubSection>
<subSubSection id="C32A18E8FFE12364FF4CF93CFD62DB12" lastPageId="45" lastPageNumber="46" pageId="44" pageNumber="45" type="diagnosis">
<paragraph id="8B8F4B63FFE12364FF4CF93CFD11D817" blockId="44.[151,1436,1744,2035]" lastBlockId="45.[151,1436,152,1972]" lastPageId="45" lastPageNumber="46" pageId="44" pageNumber="45">
<emphasis id="B9449771FFE12365FF4CF93CFE9CDFDF" bold="true" box="[151,280,1744,1770]" pageId="44" pageNumber="45">Diagnosis.</emphasis>
A species with (1) mean SVL in adult males
<quantity id="4CC8E686FFE12365FCE6F93DFC2CDFDE" box="[829,936,1745,1771]" metricMagnitude="-2" metricUnit="m" metricValue="2.27" pageId="44" pageNumber="45" unit="mm" value="22.7">22.7 mm</quantity>
(19.724.9, SD = 1,
<emphasis id="B9449771FFE12365FB7DF93DFB31DFDF" box="[1190,1205,1745,1770]" italics="true" pageId="44" pageNumber="45">n</emphasis>
= 30), and in adult females
<quantity id="4CC8E686FFE12365FEDBF91AFEEADE25" box="[256,366,1782,1808]" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="44" pageNumber="45" unit="mm" value="25.4">25.4 mm</quantity>
(22.228.7, SD = 1.4,
<emphasis id="B9449771FFE12365FD54F91AFD1ADE3A" box="[655,670,1782,1807]" italics="true" pageId="44" pageNumber="45">n</emphasis>
= 29); (2) disc of Finger IV expanded 1.5 times the width of adjacent phalange; (3) Finger II shorter than Finger III; (4) fringes present on fingers; (5) disc of Toe IV expanded 1.5 times the width of adjacent phalange; (6) fringes present on toes; (7) outer metatarsal fold present; (8) basal toe webbing, formula
<emphasis id="B9449771FFE12365FDA2F884FD0ADEB7" bold="true" box="[633,654,1896,1922]" pageId="44" pageNumber="45">I(</emphasis>
2)—(23)
<emphasis id="B9449771FFE12365FCE2F884FCDFDEB7" bold="true" box="[825,859,1896,1922]" pageId="44" pageNumber="45">II(</emphasis>
2)—(23½)
<emphasis id="B9449771FFE12365FBC7F884FBBADEB7" bold="true" box="[1052,1086,1896,1922]" pageId="44" pageNumber="45">III</emphasis>
(2
<superScript id="7C45E62BFFE12365FB83F88AFBE5DE41" attach="none" box="[1112,1121,1894,1908]" fontSize="6" pageId="44" pageNumber="45">+</superScript>
3)—(3½4
<superScript id="7C45E62BFFE12365FB28F88AFB78DE41" attach="none" box="[1267,1276,1894,1908]" fontSize="6" pageId="44" pageNumber="45">+</superScript>
)
<emphasis id="B9449771FFE12365FADCF884FAACDEB7" bold="true" box="[1287,1320,1896,1922]" pageId="44" pageNumber="45">IV</emphasis>
(44½)— (23)
<emphasis id="B9449771FFE12365FF0CF862FF6EDE9D" bold="true" box="[215,234,1934,1960]" pageId="44" pageNumber="45">V</emphasis>
; (9) oblique lateral stripe complete, ventrolateral stripe complete, diffuse or interrupted; (10) dorsum with three large blotches; (11) gular-chest region uniformly brown or gray in adult males, cream-colored or slightly brown with white spots in adult females; (12) venter uniformly brown or gray in adult males, cream slightly translucent or slightly flecked with brown in adult females; (13) sexual dimorphism in ventral coloration present; (14) tadpoles light brown with scattered brown blotches on tail and a large longitudinal brown mark on dorsum of tail musculature; (15) tadpole spiracular opening oriented dorsoposteriorly; (16) call is a long trill composed of single notes.
</paragraph>
<paragraph id="8B8F4B63FFE02364FF1DFEC1FD62DB12" blockId="45.[151,1436,152,1972]" pageId="45" pageNumber="46">
<taxonomicName id="4C3030E0FFE02364FF1DFEC1FEF9D873" baseAuthorityName="Edwards" baseAuthorityYear="1974" box="[198,381,301,326]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="sauli">
<emphasis id="B9449771FFE02364FF1DFEC1FEF9D873" box="[198,381,301,326]" italics="true" pageId="45" pageNumber="46">Hyloxalus sauli</emphasis>
</taxonomicName>
is similar to those species with basal toe webbing, oblique lateral stripe complete, and that lack dorsolateral stripe and pectoral marks (
<taxonomicName id="4C3030E0FFE02364FD7EFEBEFCE1D859" authority="Coloma" authorityName="Coloma" box="[677,869,338,364]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="awa">
<emphasis id="B9449771FFE02364FD7EFEBEFD78D85E" box="[677,764,338,363]" italics="true" pageId="45" pageNumber="46">H. awa</emphasis>
Coloma
</taxonomicName>
,
<taxonomicName id="4C3030E0FFE02364FCA9FEBEFB06D859" authority="Duellman" authorityName="Duellman" box="[882,1154,338,364]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="insulatus">
<emphasis id="B9449771FFE02364FCA9FEBEFB86D85E" box="[882,1026,338,363]" italics="true" pageId="45" pageNumber="46">H. insulatus</emphasis>
Duellman
</taxonomicName>
,
<taxonomicName id="4C3030E0FFE02364FB54FEBEFAF6D859" authority="Pefaur" authorityName="Pefaur" box="[1167,1394,338,364]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="littoralis">
<emphasis id="B9449771FFE02364FB54FEBEFA9FD85E" box="[1167,1307,338,363]" italics="true" pageId="45" pageNumber="46">H. littoralis</emphasis>
Péfaur
</taxonomicName>
,
<taxonomicName id="4C3030E0FFE02364FAA4FEBEFED5D8A7" authority="Coloma" authorityName="Coloma" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="toachi">
<emphasis id="B9449771FFE02364FAA4FEBEFF60D8A4" italics="true" pageId="45" pageNumber="46">H. toachi</emphasis>
Coloma
</taxonomicName>
,
<taxonomicName id="4C3030E0FFE02364FE85FE94FDDFD8A7" authority="Hellmich" authorityName="Hellmich" authorityYear="1940" box="[350,603,376,402]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="vergeli">
<emphasis id="B9449771FFE02364FE85FE94FE5DD8A4" box="[350,473,376,401]" italics="true" pageId="45" pageNumber="46">H. vergeli</emphasis>
Hellmich
</taxonomicName>
).
<taxonomicName id="4C3030E0FFE02364FDABFE94FCACD8A4" box="[624,808,376,401]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="awa">
<emphasis id="B9449771FFE02364FDABFE94FCACD8A4" box="[624,808,376,401]" italics="true" pageId="45" pageNumber="46">Hyloxalus awa</emphasis>
</taxonomicName>
has white spots over dark brown throat and chest.
<taxonomicName id="4C3030E0FFE02364FF4CFE71FE09D883" box="[151,397,413,438]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="insulatus">
<emphasis id="B9449771FFE02364FF4CFE71FE09D883" box="[151,397,413,438]" italics="true" pageId="45" pageNumber="46">Hyloxalus insulatus</emphasis>
</taxonomicName>
has Finger II longer than Finger III and lacks outer metatarsal fold and metatarsal tubercle.
<taxonomicName id="4C3030E0FFE02364FED8FE2EFE62D8EE" box="[259,486,450,475]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="littoralis">
<emphasis id="B9449771FFE02364FED8FE2EFE62D8EE" box="[259,486,450,475]" italics="true" pageId="45" pageNumber="46">Hyloxalus littoralis</emphasis>
</taxonomicName>
exhibits brown spots on gular-chest region, forming a subtle collar-like mark and Fingers II and III are equal in size.
<taxonomicName id="4C3030E0FFE02364FDEAFE04FD7CDB34" box="[561,760,488,513]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="toachi">
<emphasis id="B9449771FFE02364FDEAFE04FD7CDB34" box="[561,760,488,513]" italics="true" pageId="45" pageNumber="46">Hyloxalus toachi</emphasis>
</taxonomicName>
has no fringes on fingers and testes are brown.
<taxonomicName id="4C3030E0FFE02364FAFEFE04FF63DB13" authorityName="Hellmich" authorityYear="1940" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="vergeli">
<emphasis id="B9449771FFE02364FAFEFE04FF63DB13" italics="true" pageId="45" pageNumber="46">Hyloxalus vergeli</emphasis>
</taxonomicName>
is larger in size (SVL greater than
<quantity id="4CC8E686FFE02364FD59FDE1FD5EDB12" box="[642,730,525,551]" metricMagnitude="-2" metricUnit="m" metricValue="2.8" pageId="45" pageNumber="46" unit="mm" value="28.0">28 mm</quantity>
).
</paragraph>
</subSubSection>
<subSubSection id="C32A18E8FFE02367FF1EFDDEFC31D8A7" lastPageId="46" lastPageNumber="47" pageId="45" pageNumber="46" type="description">
<paragraph id="8B8F4B63FFE02364FF1EFDDEFBF4DB89" blockId="45.[151,1436,152,1972]" pageId="45" pageNumber="46">
<emphasis id="B9449771FFE02364FF1EFDDEFEE5DB79" bold="true" box="[197,353,562,588]" pageId="45" pageNumber="46">Description.</emphasis>
<bibRefCitation id="EFA13692FFE02364FEB1FDDEFDBEDB79" author="Edwards, S. R." box="[362,570,562,588]" pageId="45" pageNumber="46" pagination="1 - 14" refId="ref45111" refString="Edwards, S. R. (1974 b) Taxonomic notes on South American dendrobatid frogs of the genus Colostethus. University of Kansas, Natural History Museum, Occasional Papers, 30, 1 - 14." type="journal article" year="1974">Edwards (1974b)</bibRefCitation>
provided a morphological description of the species under the name of
<taxonomicName id="4C3030E0FFE02364FF4CFDB4FEE5DB44" authorityName="Edwards" authorityYear="1974" box="[151,353,600,625]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="sauli">
<emphasis id="B9449771FFE02364FF4CFDB4FEE5DB44" box="[151,353,600,625]" italics="true" pageId="45" pageNumber="46">Colostethus sauli</emphasis>
</taxonomicName>
. Variation in measurements is given in
<tableCitation id="C6B27ED8FFE02364FCF4FDB4FC02DB47" box="[815,902,600,626]" captionStart="TABLE 4" captionStartId="12.[151,241,399,423]" captionText="TABLE 4. Measurements (in mm) of adults of Hyloxalus bocagei, H. maculosus and H. sauli. Mean ± SD, and range are given. Abbreviations are defined in the text." pageId="45" pageNumber="46">Table 4</tableCitation>
(
<emphasis id="B9449771FFE02364FC4DFDB4FC21DB44" box="[918,933,600,625]" italics="true" pageId="45" pageNumber="46">n</emphasis>
= 59:
<specimenCount id="9D3680EAFFE02364FC2BFDB4FBD8DB47" box="[1008,1116,600,626]" pageId="45" pageNumber="46" type="male">30 males</specimenCount>
,
<specimenCount id="9D3680EAFFE02364FBBCFDB4FB69DB47" box="[1127,1261,600,626]" pageId="45" pageNumber="46" type="female">29 females</specimenCount>
). Head slightly longer than wide; snout round in dorsal view, angular in lateral view; loreal region flat; nostrils slightly protuberant; supratympanic bulge diffuse, covering upper edge of tympanum or not.
</paragraph>
<paragraph id="8B8F4B63FFE02364FF1DFD24FACDDADF" blockId="45.[151,1436,152,1972]" pageId="45" pageNumber="46">
Finger II shorter than Finger III, fingers unwebbed; fringes present but discrete; terminal discs 1.5 times width of adjacent phalange; subarticular tubercles round, that of Finger II slightly oval; outer metacarpal tubercle large, round; inner metacarpal tubercle at base of thumb small, elliptical and subtly protuberant. Outer metatarsal fold present; inner sigmoid tarsal fold present on distal third of half of tarsus; outer metatarsal tubercle round; inner metatarsal tubercle smaller than outer, elliptical; protuberance between both metatarsal tubercles in 60% of examined specimens; toe webbing formula
<emphasis id="B9449771FFE02364FBFFFC6EFBBEDAA9" bold="true" box="[1060,1082,898,924]" pageId="45" pageNumber="46">I(</emphasis>
2)—(23)
<emphasis id="B9449771FFE02364FB3CFC6EFA8CDAA9" bold="true" box="[1255,1288,898,924]" pageId="45" pageNumber="46">II(</emphasis>
2)—(2 3½)
<emphasis id="B9449771FFE02364FF1DFC46FF6CDAF1" bold="true" box="[198,232,938,964]" pageId="45" pageNumber="46">III</emphasis>
(2
<superScript id="7C45E62BFFE02364FEDAFC44FE8EDA83" attach="none" box="[257,266,936,950]" fontSize="6" pageId="45" pageNumber="46">+</superScript>
3)—(3½4
<superScript id="7C45E62BFFE02364FE47FC44FE21DA83" attach="none" box="[412,421,936,950]" fontSize="6" pageId="45" pageNumber="46">+</superScript>
)
<emphasis id="B9449771FFE02364FE74FC46FE54DAF1" bold="true" box="[431,464,938,964]" pageId="45" pageNumber="46">IV</emphasis>
(44½)—(23)
<emphasis id="B9449771FFE02364FD5FFC46FD13DAF1" bold="true" box="[644,663,938,964]" pageId="45" pageNumber="46">V</emphasis>
; lateral fringes present but discrete; terminal discs expanded 1.5 times width of adjacent phalange; subarticular tubercles small, round; supernumerary tubercles absent.
</paragraph>
<paragraph id="8B8F4B63FFE02364FF1DFC19FD0EDD6F" blockId="45.[151,1436,152,1972]" pageId="45" pageNumber="46">Skin of dorsum, limbs and venter smooth; flanks rough; tubercle posterior to mandible junction; anal sheath conspicuous. Tongue more than twice as long as wide, thinner and broader posteriorly, posterior half not attached to mouths floor. Testis white.</paragraph>
<paragraph id="8B8F4B63FFE02364FF1EFB89FED6DCC1" blockId="45.[151,1436,152,1972]" pageId="45" pageNumber="46">
<emphasis id="B9449771FFE02364FF1EFB89FE49DD4A" bold="true" box="[197,461,1125,1151]" pageId="45" pageNumber="46">Color in preservative</emphasis>
(~70% ethanol). Variation in color patterns is depicted on
<figureCitation id="130B57E6FFE02364FBA0FB89FB5BDD4A" box="[1147,1247,1125,1151]" captionStart="FIGURE 5" captionStartId="17.[151,259,1774,1798]" captionTargetBox="[301,1270,194,1744]" captionTargetId="figure-155@17.[301,1270,194,1744]" captionTargetPageId="17" captionText="FIGURE 5. Dorsal view of adults showing intra and inter-population variation in dorsal coloration pattern: (A) Hyloxalus bocagei, left to right: QCAZ 32388 (Río Leo), 20824 (La Virgen), 37620 (Hostería El Reventador), 37260 (La Virgen); (B) H. maculosus, left to right: QCAZ 37156 (Puyo-Shell road), 42074 (Santa Rosa), 19975 (Puerto Misahuallí), 37870 (Puyo-Shell road); (C) H. sauli, left to right: QCAZ 33382, 39893 (PUCEs Yasuní Research Station), 6233 (Jatunsacha), 33384 (PUCEs Yasuní Research Station); (D) H. faciopunctulatus, left to right: MCZ-A 93782, 94746, 94749, 94761 (Puerto Nariño); (E) H. italoi, left to right: QCAZ 33200, 37202, 33194, 33199 (Reserva Hola Vida); (F) H. yasuni, left to right: QCAZ 25065, 32467, 25059, 22557 (PUCEs Yasuní Research Station). Labels were removed from specimens of H. faciopunctulatus using Adobe Photoshop. Scale bar = 10 mm. Photos by LAC." pageId="45" pageNumber="46">Figs. 5C</figureCitation>
<figureCitation id="130B57E6FFE02364FB34FB89FA97DD4A" box="[1263,1299,1125,1151]" captionStart="FIGURE 7" captionStartId="19.[151,255,1780,1804]" captionTargetBox="[298,1282,180,1753]" captionTargetId="figure-156@19.[298,1282,180,1753]" captionTargetPageId="19" captionText="FIGURE 7. Ventral view of adult females showing intra and inter-population variation in ventral patterns: (A) Hyloxalus bocagei, left to right: QCAZ 32390, 32393 (Río Leo), 37620 (Hostería El Reventador), 37260 (La Virgen); (B) H. maculosus, left to right: QCAZ 19975 (Puerto Misahuallí), 37870 (Shell-Puyo road), 42078, 42082 (Santa Rosa); (C) H. sauli, left to right: QCAZ 1435 (Locación Petrolera Garza 1), 35462 (PUCEs Yasuní Research Station), 6233 (Jatunsacha), 33384 (PUCEs Yasuní Research Station); (D) H. faciopunctulatus, left to right: MCZ-A 94759, 94761, 94750, 94752 (Puerto Nariño); (E) H. italoi, left to right: QCAZ 37171, 33194, 33199, 37213 (Reserva Hola Vida); (F) H. yasuni, left to right: QCAZ 22588, 25059, 33379, 22557 (PUCEs Yasuní Research Station). Labels were removed from specimens of H. faciopunctulatus using Adobe Photoshop. Scale bar = 10 mm. Photos by LAC." pageId="45" pageNumber="46">7C</figureCitation>
. Dorsum of head and body brown, slightly reddish; a dark brown interorbital blotch, a V-shaped mark at scapular level and a blotch at posterior portion of dorsum; dark brown lateral stripe from snout to groin. Pale oblique lateral stripe conspicuous from posterior corner of eye to groin, sometimes diffuse at anterior portion; pale ventrolateral stripe complete, diffuse or interrupted. Forelimbs brown with darker transverse bars on forearm, longitudinal dark brown stripe at anterior surface of arm, cream spot at arm-body junction; dorsal surfaces of hind limbs brown with darker transverse bars (911 at leg and tarsus), cream spot posterior to groin; ventral surface cream in females, brown in males. Gular-chest region uniformly dark brown in males, cream or light brown in females; venter uniformly gray or brown in males, cream slightly flecked with brown in females; axillas translucent, brown band at each axilla in some male specimens. Palmar and plantar surfaces brown; toe webbing cream.
</paragraph>
<paragraph id="8B8F4B63FFE02364FF1EF9ECFEAEDF51" blockId="45.[151,1436,152,1972]" pageId="45" pageNumber="46">
<emphasis id="B9449771FFE02364FF1EF9ECFEF4DF2F" bold="true" box="[197,368,1536,1562]" pageId="45" pageNumber="46">Color in life.</emphasis>
<bibRefCitation id="EFA13692FFE02364FEA0F9ECFDD0DF2F" author="Edwards, S. R." box="[379,596,1536,1562]" pageId="45" pageNumber="46" pagination="1 - 14" refId="ref45111" refString="Edwards, S. R. (1974 b) Taxonomic notes on South American dendrobatid frogs of the genus Colostethus. University of Kansas, Natural History Museum, Occasional Papers, 30, 1 - 14." type="journal article" year="1974">Edwards (1974b)</bibRefCitation>
described color in life. An adult male and a female from PUCEs (Pontificia Universidad Católica del
<collectingCountry id="F3270BF3FFE02364FD85F9C9FD41DF0A" box="[606,709,1573,1599]" name="Ecuador" pageId="45" pageNumber="46">Ecuador</collectingCountry>
) Yasuní Research Station are depicted in
<figureCitation id="130B57E6FFE02364FB10F9C9FAB7DF0A" box="[1227,1331,1573,1599]" captionStart="FIGURE 3" captionStartId="15.[151,257,1878,1902]" captionTargetBox="[353,1257,194,1846]" captionTargetId="figure-95@15.[321,1257,194,1846]" captionTargetPageId="15" captionText="FIGURE 3. Dorsal (left) and ventral (right) views of males of (A) Hyloxalus bocagei from Hostería El Reventador (QCAZ 42095, SVL = 23.7 mm); (B) H. maculosus from Santa Rosa (QCAZ 42085, SVL = 24 mm); (C) H. sauli from PUCEs Yasuní Research Station (QCAZ 45516, SVL = 22.6 mm); (D) H. italoi from Reserva Hola Vida (Holotype, QCAZ 33197, SVL = 24.5 mm); (E) H. yasuni from PUCEs Yasuní Research Station (QCAZ 45517, SVL = 24.5 mm). Photos by LAC." pageId="45" pageNumber="46">Figs. 3C</figureCitation>
and
<figureCitation id="130B57E6FFE02364FAAAF9C9FA12DF0A" box="[1393,1430,1573,1599]" captionStart="FIGURE 4" captionStartId="16.[151,256,1867,1891]" captionTargetBox="[358,1262,194,1842]" captionTargetId="figure-95@16.[326,1262,194,1842]" captionTargetPageId="16" captionText="FIGURE 4. Dorsal (left) and ventral (right) views of females of (A) Hyloxalus bocagei from Hostería El Reventador (QCAZ 42099, SVL = 28 mm); (B) H. maculosus from Santa Rosa (QCAZ 42081, SVL =28.7 mm); (C) H. sauli from PUCEs Yasuní Research Station (QCAZ 45513, SVL = 27.4 mm); (D) H. italoi from Reserva Hola Vida (QCAZ 33191, SVL = 25.8 mm); (E) H. yasuni from PUCEs Yasuní Research Station (QCAZ 45488, SVL = 26.8 mm). Photos by LAC." pageId="45" pageNumber="46">4C</figureCitation>
, respectively.
</paragraph>
<paragraph id="8B8F4B63FFE02364FF1EF99CFD2FDFCF" blockId="45.[151,1436,152,1972]" pageId="45" pageNumber="46">
<emphasis id="B9449771FFE02364FF1EF99CFEC1DFBF" bold="true" box="[197,325,1648,1674]" pageId="45" pageNumber="46">Variation.</emphasis>
Variation in measurements is given in
<tableCitation id="C6B27ED8FFE02364FCCAF99CFCEFDFBF" box="[785,875,1648,1674]" captionStart="TABLE 4" captionStartId="12.[151,241,399,423]" captionText="TABLE 4. Measurements (in mm) of adults of Hyloxalus bocagei, H. maculosus and H. sauli. Mean ± SD, and range are given. Abbreviations are defined in the text." pageId="45" pageNumber="46">Table 4</tableCitation>
. Variation in color patterns is depicted on
<figureCitation id="130B57E6FFE02364FAB9F99CFF3EDF9A" captionStart="FIGURE 5" captionStartId="17.[151,259,1774,1798]" captionTargetBox="[301,1270,194,1744]" captionTargetId="figure-155@17.[301,1270,194,1744]" captionTargetPageId="17" captionText="FIGURE 5. Dorsal view of adults showing intra and inter-population variation in dorsal coloration pattern: (A) Hyloxalus bocagei, left to right: QCAZ 32388 (Río Leo), 20824 (La Virgen), 37620 (Hostería El Reventador), 37260 (La Virgen); (B) H. maculosus, left to right: QCAZ 37156 (Puyo-Shell road), 42074 (Santa Rosa), 19975 (Puerto Misahuallí), 37870 (Puyo-Shell road); (C) H. sauli, left to right: QCAZ 33382, 39893 (PUCEs Yasuní Research Station), 6233 (Jatunsacha), 33384 (PUCEs Yasuní Research Station); (D) H. faciopunctulatus, left to right: MCZ-A 93782, 94746, 94749, 94761 (Puerto Nariño); (E) H. italoi, left to right: QCAZ 33200, 37202, 33194, 33199 (Reserva Hola Vida); (F) H. yasuni, left to right: QCAZ 25065, 32467, 25059, 22557 (PUCEs Yasuní Research Station). Labels were removed from specimens of H. faciopunctulatus using Adobe Photoshop. Scale bar = 10 mm. Photos by LAC." pageId="45" pageNumber="46">Figs. 5C</figureCitation>
<figureCitation id="130B57E6FFE02364FF11F979FF6ADF9A" box="[202,238,1685,1711]" captionStart="FIGURE 7" captionStartId="19.[151,255,1780,1804]" captionTargetBox="[298,1282,180,1753]" captionTargetId="figure-156@19.[298,1282,180,1753]" captionTargetPageId="19" captionText="FIGURE 7. Ventral view of adult females showing intra and inter-population variation in ventral patterns: (A) Hyloxalus bocagei, left to right: QCAZ 32390, 32393 (Río Leo), 37620 (Hostería El Reventador), 37260 (La Virgen); (B) H. maculosus, left to right: QCAZ 19975 (Puerto Misahuallí), 37870 (Shell-Puyo road), 42078, 42082 (Santa Rosa); (C) H. sauli, left to right: QCAZ 1435 (Locación Petrolera Garza 1), 35462 (PUCEs Yasuní Research Station), 6233 (Jatunsacha), 33384 (PUCEs Yasuní Research Station); (D) H. faciopunctulatus, left to right: MCZ-A 94759, 94761, 94750, 94752 (Puerto Nariño); (E) H. italoi, left to right: QCAZ 37171, 33194, 33199, 37213 (Reserva Hola Vida); (F) H. yasuni, left to right: QCAZ 22588, 25059, 33379, 22557 (PUCEs Yasuní Research Station). Labels were removed from specimens of H. faciopunctulatus using Adobe Photoshop. Scale bar = 10 mm. Photos by LAC." pageId="45" pageNumber="46">7C</figureCitation>
. Specimens from Bloque Shiripuno (
<emphasis id="B9449771FFE02364FD79F97AFD35DF9A" box="[674,689,1686,1711]" italics="true" pageId="45" pageNumber="46">n</emphasis>
= 2, QCAZ 8984 and 9329) and Locación Petrolera Garza 1 (
<emphasis id="B9449771FFE02364FA56F97AFA18DF9A" box="[1421,1436,1686,1711]" italics="true" pageId="45" pageNumber="46">n</emphasis>
= 1, QCAZ 1435) lack ventrolateral stripe. In specimens from Puerto
<collectingRegion id="49F48581FFE02364FC1AF956FB9DDFE1" box="[961,1049,1722,1748]" country="Ecuador" name="Bolivar" pageId="45" pageNumber="46">Bolívar</collectingRegion>
(
<emphasis id="B9449771FFE02364FBF1F957FBBDDFE1" box="[1066,1081,1723,1748]" italics="true" pageId="45" pageNumber="46">n</emphasis>
= 3, QCAZ 2826228263 and 28339), the ventrolateral stripe is very subtle.
</paragraph>
<paragraph id="8B8F4B63FFE02364FF1EF8E9FAD1DE81" blockId="45.[151,1436,152,1972]" pageId="45" pageNumber="46">Twenty juveniles (QCAZ 484, 486487, 5318, 6879, 8082, 94049405, 22626, 2410124102, 24237 24238, 2505025051, 25056, 25060, 25066, 2826228263, 32476, 32478; USNM 282108282112) with SVL of 10.321.4 (mean = 15.2) exhibit same coloration pattern as adults. Nonetheless, they are slightly paler and, thus, dark dorsal blotches are more contrasting. Ventrally cream-colored, some specimens translucent. Two recently metamorphosed specimens (QCAZ 5313 and 25051) have SVL of 12.1 and 10.49 respectively.</paragraph>
<paragraph id="8B8F4B63FFE32367FF1EFF7BFC31D8A7" blockId="46.[151,1436,151,626]" pageId="46" pageNumber="47">
<emphasis id="B9449771FFE32367FF1EFF7BFEB6D984" bold="true" box="[197,306,151,177]" pageId="46" pageNumber="47">Tadpole.</emphasis>
<bibRefCitation id="EFA13692FFE32367FEE0FF74FD8DD987" author="Edwards, S. R." box="[315,521,152,178]" pageId="46" pageNumber="47" pagination="1 - 14" refId="ref45111" refString="Edwards, S. R. (1974 b) Taxonomic notes on South American dendrobatid frogs of the genus Colostethus. University of Kansas, Natural History Museum, Occasional Papers, 30, 1 - 14." type="journal article" year="1974">Edwards (1974b)</bibRefCitation>
described tadpoles in Stages 2736.
<bibRefCitation id="EFA13692FFE32367FC12FF74FB13D987" author="Duellman, W. E." box="[969,1175,152,178]" pageId="46" pageNumber="47" pagination="1 - 352" refId="ref44862" refString="Duellman, W. E. (1978) The biology of an Equatorial herpetofauna in Amazonian Ecuador. Miscellaneous Publications of the Museum of Natural History University of Kansas, 65, 1 - 352." type="journal article" year="1978">Duellman (1978)</bibRefCitation>
described tadpoles at Stage 36. Tadpoles of
<taxonomicName id="4C3030E0FFE32367FE7AFF51FDD8D9E3" baseAuthorityName="Edwards" baseAuthorityYear="1974" box="[417,604,189,214]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="sauli">
<emphasis id="B9449771FFE32367FE7AFF51FDD8D9E3" box="[417,604,189,214]" italics="true" pageId="46" pageNumber="47">Hyloxalus sauli</emphasis>
</taxonomicName>
are similar to
<taxonomicName id="4C3030E0FFE32367FCD1FF51FC09D9E3" baseAuthorityName="Jimenez de la Espada" baseAuthorityYear="1870" box="[778,909,189,214]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="bocagei">
<emphasis id="B9449771FFE32367FCD1FF51FC09D9E3" box="[778,909,189,214]" italics="true" pageId="46" pageNumber="47">H. bocagei</emphasis>
</taxonomicName>
tadpoles. Nevertheless, the latter are round, their snout is more elongated, tail musculature is slender, tip of the tail is round, dorsal fin is broader at its anterior region and they are darker; the upper edge of upper jaw is straight, whereas it is subtly winding in
<taxonomicName id="4C3030E0FFE32367FAA4FEE4FF56D873" baseAuthorityName="Edwards" baseAuthorityYear="1974" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="sauli">
<emphasis id="B9449771FFE32367FAA4FEE4FF56D873" italics="true" pageId="46" pageNumber="47">H. sauli</emphasis>
</taxonomicName>
. Tadpoles of
<taxonomicName id="4C3030E0FFE32367FEA3FEC1FE5DD873" baseAuthorityName="Edwards" baseAuthorityYear="1974" box="[376,473,301,326]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="sauli">
<emphasis id="B9449771FFE32367FEA3FEC1FE5DD873" box="[376,473,301,326]" italics="true" pageId="46" pageNumber="47">H. sauli</emphasis>
</taxonomicName>
are similar to
<taxonomicName id="4C3030E0FFE32367FD57FEC1FCB4D873" baseAuthorityName="Rivero" baseAuthorityYear="1991" box="[652,816,301,326]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="maculosus">
<emphasis id="B9449771FFE32367FD57FEC1FCB4D873" box="[652,816,301,326]" italics="true" pageId="46" pageNumber="47">H. maculosus</emphasis>
</taxonomicName>
tadpoles as well. However, tail in
<taxonomicName id="4C3030E0FFE32367FB02FEC1FABDD873" baseAuthorityName="Edwards" baseAuthorityYear="1974" box="[1241,1337,301,326]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="sauli">
<emphasis id="B9449771FFE32367FB02FEC1FABDD873" box="[1241,1337,301,326]" italics="true" pageId="46" pageNumber="47">H. sauli</emphasis>
</taxonomicName>
is more heavily pigmented, spiracular opening is directed dorsoposteriorly (laterally in
<taxonomicName id="4C3030E0FFE32367FB9DFEBEFB63D85E" baseAuthorityName="Rivero" baseAuthorityYear="1991" box="[1094,1255,338,363]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="maculosus">
<emphasis id="B9449771FFE32367FB9DFEBEFB63D85E" box="[1094,1255,338,363]" italics="true" pageId="46" pageNumber="47">H. maculosus</emphasis>
</taxonomicName>
) and the upper edge of the upper jaw is slightly winding (straight in
<taxonomicName id="4C3030E0FFE32367FCDFFE94FC27D8A4" baseAuthorityName="Rivero" baseAuthorityYear="1991" box="[772,931,376,401]" class="Amphibia" family="Dendrobatidae" genus="Hyloxalus" kingdom="Animalia" order="Anura" pageId="46" pageNumber="47" phylum="Chordata" rank="species" species="maculosus">
<emphasis id="B9449771FFE32367FCDFFE94FC27D8A4" box="[772,931,376,401]" italics="true" pageId="46" pageNumber="47">H. maculosus</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="C32A18E8FFE32367FF1EFE71FF68DB47" pageId="46" pageNumber="47" type="discussion">
<paragraph id="8B8F4B63FFE32367FF1EFE71FF68DB47" blockId="46.[151,1436,151,626]" pageId="46" pageNumber="47">
<emphasis id="B9449771FFE32367FF1EFE71FED7D882" bold="true" box="[197,339,413,439]" pageId="46" pageNumber="47">Comments.</emphasis>
Edwards (1974b) description is appropriate with the exceptions noticed by
<bibRefCitation id="EFA13692FFE32367FB3AFE71FA1CD882" author="Coloma, L. A." box="[1249,1432,413,439]" pageId="46" pageNumber="47" pagination="1 - 72" refId="ref44823" refString="Coloma, L. A. (1995) Ecuadorian frogs of the genus Colostethus (Anura: Dendrobatidae). Miscellaneous Publications of the Museum of Natural History University of Kansas, 87, 1 - 72." type="journal article" year="1995">Coloma (1995)</bibRefCitation>
. Additionally, Finger II is shorter than Finger III and females might exhibit brown spots on venter, whereas venter in males lacks spots.
<bibRefCitation id="EFA13692FFE32367FE06FE04FD23DB37" author="Edwards, S. R." box="[477,679,488,514]" pageId="46" pageNumber="47" pagination="1 - 14" refId="ref45111" refString="Edwards, S. R. (1974 b) Taxonomic notes on South American dendrobatid frogs of the genus Colostethus. University of Kansas, Natural History Museum, Occasional Papers, 30, 1 - 14." type="journal article" year="1974">Edwards (1974b)</bibRefCitation>
also remarks on the absence of ventrolateral stripe. Nonetheless, 88% of examined specimens exhibit ventrolateral stripe, complete, diffuse, or interrupted. Further information is included under Phylogeny and Chronogram and Ancestral Area Reconstruction sections (
<tableCitation id="C6B27ED8FFE32367FB38FDDEFACDDB79" box="[1251,1353,562,588]" captionStart="TABLE 2" captionStartId="8.[151,241,1036,1060]" captionText="TABLE 2. Pairwise patristic distances calculated from the 12S-16S sequences from ML tree (Fig. 1) branch lengths that describe the amount of phylogenetic change between individuals of the clade bocagei. Each patristic distance is the sum of the lengths of the branches that link two terminal nodes." pageId="46" pageNumber="47">Tables 2</tableCitation>
and
<tableCitation id="C6B27ED8FFE32367FA5EFDDEFA12DB79" box="[1413,1430,562,588]" captionStart="TABLE 3" captionStartId="11.[151,240,151,175]" captionText="TABLE 3. Pairwise patristic distances calculated from the Cytochrome b sequences from ML tree branch lengths that describe the amount of phylogenetic change between individuals of the clade bocagei. Numbers in the individual column and row correspond to those indicated in Table 2." pageId="46" pageNumber="47">3</tableCitation>
;
<figureCitation id="130B57E6FFE32367FF4CFDB4FF5EDB47" box="[151,218,600,626]" captionStart="FIGURE 1" captionStartId="10.[151,255,1516,1540]" captionTargetBox="[156,1436,223,1477]" captionTargetId="figure-107@10.[151,1436,197,1495]" captionTargetPageId="10" captionText="FIGURE 1. Maximum likelihood phylogeny of the bocagei clade. The population localities, museum numbers, nodal support are provided." pageId="46" pageNumber="47">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>