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<document id="B8FBD65D0D457AFE78625F812EE9373F" ID-CLB-Dataset="50816" ID-DOI="10.5281/zenodo.171517" ID-GBIF-Dataset="53382c85-70e8-4c8a-af71-0610d52c0ee9" ID-ISSN="1175­5326" ID-Zenodo-Dep="171517" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1459754193686" checkinUser="plazi" docAuthor="Barrio-Amorós, César L." docDate="2006" docId="E72287BAFFFDFF93BC126923FC0FFDD7" docLanguage="en" docName="zt01110p068.pdf" docOrigin="Zootaxa 1110" docStyle="DocumentStyle:FA7E419B012A62B0FC3AC15A186C3DAF.3:Zootaxa.2001-2006.journal_article" docStyleId="FA7E419B012A62B0FC3AC15A186C3DAF" docStyleName="Zootaxa.2001-2006.journal_article" docStyleVersion="3" docTitle="Colostethus breweri Barrio-Amorós, 2006, sp. nov." docType="treatment" docVersion="8" lastPageNumber="66" masterDocId="1B1BFFC2FFFCFF94BD1A6C32FFF4FFD6" masterDocTitle="A new dendrobatid frog (Anura: Dendrobatidae: Colostethus) from Aprada tepui, southern Venezuela" masterLastPageNumber="68" masterPageNumber="59" pageNumber="60" updateTime="1698216997379" updateUser="ExternalLinkService">
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<mods:title id="4CCB61F09831C9C842D198DCD504DC67">A new dendrobatid frog (Anura: Dendrobatidae: Colostethus) from Aprada tepui, southern Venezuela</mods:title>
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<mods:namePart id="D106172AD36E2D45C22048334165DDE7">Barrio-Amorós, César L.</mods:namePart>
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<mods:date id="069D6B9CCCA4A3A649029ADE76DF28F7">2006</mods:date>
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<treatment id="E72287BAFFFDFF93BC126923FC0FFDD7" ID-DOI="http://doi.org/10.5281/zenodo.6253850" ID-GBIF-Taxon="119338524" ID-Zenodo-Dep="6253850" LSID="urn:lsid:plazi:treatment:E72287BAFFFDFF93BC126923FC0FFDD7" httpUri="http://treatment.plazi.org/id/E72287BAFFFDFF93BC126923FC0FFDD7" lastPageId="7" lastPageNumber="66" pageId="1" pageNumber="60">
<subSubSection id="27916527FFFDFF95BC126923FE94FA98" pageId="1" pageNumber="60" type="nomenclature">
<paragraph id="6F3436ACFFFDFF95BC126923FDA9FAFD" blockId="1.[264,605,1297,1358]" box="[264,605,1297,1323]" pageId="1" pageNumber="60">
<heading id="347C81C0FFFDFF95BC126923FDA9FAFD" bold="true" box="[264,605,1297,1323]" fontSize="11" level="1" pageId="1" pageNumber="60" reason="1">
<emphasis id="5DFFEABEFFFDFF95BC126923FDA9FAFD" bold="true" box="[264,605,1297,1323]" pageId="1" pageNumber="60">
<taxonomicName id="A88B4D2FFFFDFF95BC126923FE0CFAFD" box="[264,504,1297,1323]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="1" pageNumber="60" phylum="Chordata" rank="species" species="breweri" status="sp. nov.">
<emphasis id="5DFFEABEFFFDFF95BC126923FE0CFAFD" bold="true" box="[264,504,1297,1323]" italics="true" pageId="1" pageNumber="60">Colostethus breweri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="46CC57C5FFFDFF95BF1A6923FDA9FAFD" box="[512,605,1297,1323]" pageId="1" pageNumber="60" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="6F3436ACFFFDFF95BC126906FE94FA98" blockId="1.[264,605,1297,1358]" box="[264,352,1332,1358]" pageId="1" pageNumber="60">
(
<figureCitation id="F7B02A29FFFDFF95BC0B6906FEA0FA98" box="[273,340,1332,1358]" captionStart="FIGURE 1" captionStartId="3.[264,368,1139,1163]" captionTargetBox="[273,1314,329,1112]" captionTargetId="figure@3.[273,1315,329,1113]" captionTargetPageId="3" captionText="FIGURE 1. Female holotype of Colostethus breweri in life. Photo by Charles Brewer­Carías." httpUri="https://zenodo.org/record/171518/files/figure.png" pageId="1" pageNumber="60">Fig. 1</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="27916527FFFDFF95BC126944FCFDF98F" pageId="1" pageNumber="60" type="materials_examined">
<paragraph id="6F3436ACFFFDFF95BC126944FCAAF9DF" blockId="1.[264,1325,1398,1905]" pageId="1" pageNumber="60">
<typeStatus id="B030880EFFFDFF95BC126944FE8EFA46" box="[264,378,1398,1424]" pageId="1" pageNumber="60" type="holotype">
<emphasis id="5DFFEABEFFFDFF95BC126944FE8EFA46" bold="true" box="[264,378,1398,1424]" pageId="1" pageNumber="60">Holotype</emphasis>
</typeStatus>
<collectionCode id="099AAE69FFFDFF95BC826945FE0AFA47" box="[408,510,1399,1425]" httpUri="http://grbio.org/cool/4kc6-e4ia" name="Coleccion de Mastozoologia, Museo de Historia Natural de La Salle" pageId="1" pageNumber="60">MHNLS</collectionCode>
17044. An adult female with circunvoluted oviducts from the entry of Cueva del Fantasma, northwestern slope of Aprada tepui,
<geoCoordinate id="0ABF506BFFFDFF95BEF969ADFBA5FA6F" box="[995,1105,1439,1465]" direction="north" orientation="latitude" pageId="1" pageNumber="60" precision="925" value="5.45">05º 27N</geoCoordinate>
,
<geoCoordinate id="0ABF506BFFFDFF95B94569ADFB26FA6F" box="[1119,1234,1439,1465]" direction="west" orientation="longitude" pageId="1" pageNumber="60" precision="925" value="-62.45">62º 27W</geoCoordinate>
,
<quantity id="A8739B49FFFDFF95B9C469ADFADFFA6F" box="[1246,1323,1439,1465]" metricMagnitude="2" metricUnit="m" metricValue="6.6" pageId="1" pageNumber="60" unit="m" value="660.0">660 m</quantity>
above sea level, Estado Bolívar,
<collectingCountry id="179C763CFFFDFF95BF9569F5FCFEFA37" box="[655,778,1479,1505]" name="Venezuela" pageId="1" pageNumber="60">Venezuela</collectingCountry>
, collected by Charles Brewer­Carías, Alberto Tovar and Fernando Tamayo, on
<date id="1B35106CFFFDFF95BF9769DDFCAEF9DF" box="[653,858,1519,1545]" pageId="1" pageNumber="60" value="2004-02-09">February 9, 2004</date>
.
</paragraph>
<paragraph id="6F3436ACFFFDFF95BC226A25FCFDF98F" blockId="1.[264,1325,1398,1905]" pageId="1" pageNumber="60">
Paratypes—MHNLS 17045, 17046, two adult males, and
<collectionCode id="099AAE69FFFDFF95BEF86A25FBBDF9E7" box="[994,1097,1559,1585]" httpUri="http://grbio.org/cool/4kc6-e4ia" name="Coleccion de Mastozoologia, Museo de Historia Natural de La Salle" pageId="1" pageNumber="60">MHNLS</collectionCode>
17047, an unsexed juvenile with the same data as the
<typeStatus id="B030880EFFFDFF95BF816A0DFCF2F98F" box="[667,774,1599,1625]" pageId="1" pageNumber="60" type="holotype">holotype</typeStatus>
.
</paragraph>
</subSubSection>
<subSubSection id="27916527FFFDFF96BC226A54FAF1FDF2" lastPageId="2" lastPageNumber="61" pageId="1" pageNumber="60" type="diagnosis">
<paragraph id="6F3436ACFFFDFF95BC226A54FEAEF8A7" blockId="1.[264,1325,1398,1905]" pageId="1" pageNumber="60">
<emphasis id="5DFFEABEFFFDFF95BC226A54FE44F956" bold="true" box="[312,432,1638,1664]" pageId="1" pageNumber="60">Diagnosis</emphasis>
—A relatively small species (male SVL up to
<quantity id="A8739B49FFFDFF95BEFA6A55FBC2F957" box="[992,1078,1639,1665]" metricMagnitude="-2" metricUnit="m" metricValue="2.1" pageId="1" pageNumber="60" unit="mm" value="21.0">21 mm</quantity>
, only known female
<quantity id="A8739B49FFFDFF95BC126ABDFE84F97F" box="[264,368,1679,1705]" metricMagnitude="-2" metricUnit="m" metricValue="2.38" pageId="1" pageNumber="60" unit="mm" value="23.8">23.8 mm</quantity>
SVL), Finger I approximately equal to Finger II; Finger III of males not swollen; venter in preservative dirty white in males, immaculate white in the only known female; in life venter of males dirty white, female yellow; toes moderately webbed; dorsolateral stripe absent; oblique lateral stripe short, white; ventrolateral stripe absent; median lingual process (MLP) present; cloacal tubercles absent; anal sheath absent; black armband absent.
</paragraph>
<paragraph id="6F3436ACFFFEFF96BC226D29FAF1FDF2" blockId="2.[264,1324,283,1899]" pageId="2" pageNumber="61">
<emphasis id="5DFFEABEFFFEFF96BC226D29FD3AFEE3" bold="true" box="[312,718,283,309]" pageId="2" pageNumber="61">Comparison with other species</emphasis>
<taxonomicName id="A88B4D2FFFFEFF96BFF16D2EFC2AFEE3" box="[747,990,284,309]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="breweri">
<emphasis id="5DFFEABEFFFEFF96BFF16D2EFC2AFEE3" box="[747,990,284,309]" italics="true" pageId="2" pageNumber="61">Colostethus breweri</emphasis>
</taxonomicName>
shares with the following species from Venezuelan Guayana the presence of a MLP:
<taxonomicName id="A88B4D2FFFFEFF96BEFA6D76FB84FE8B" box="[992,1136,324,349]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="parkerae">
<emphasis id="5DFFEABEFFFEFF96BEFA6D76FB84FE8B" box="[992,1136,324,349]" italics="true" pageId="2" pageNumber="61">C. parkerae</emphasis>
</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFEFF96B9996D76FB08FE8B" box="[1155,1276,324,349]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="shrevei">
<emphasis id="5DFFEABEFFFEFF96B9996D76FB08FE8B" box="[1155,1276,324,349]" italics="true" pageId="2" pageNumber="61">C. shrevei</emphasis>
</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFEFF96B80A6D76FE46FE53" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="tamacuarensis">
<emphasis id="5DFFEABEFFFEFF96B80A6D76FE46FE53" italics="true" pageId="2" pageNumber="61">C. tamacuarensis</emphasis>
</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFEFF96BCD26D5EFD98FE53" box="[456,620,364,389]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="tepuyensis">
<emphasis id="5DFFEABEFFFEFF96BCD26D5EFD98FE53" box="[456,620,364,389]" italics="true" pageId="2" pageNumber="61">C. tepuyensis</emphasis>
</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFEFF96BF986D5EFD0FFE53" box="[642,763,364,389]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="triunfo">
<emphasis id="5DFFEABEFFFEFF96BF986D5EFD0FFE53" box="[642,763,364,389]" italics="true" pageId="2" pageNumber="61">C. triunfo</emphasis>
</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFEFF96BE086D5EFC6EFE53" box="[786,922,364,389]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="wothuja">
<emphasis id="5DFFEABEFFFEFF96BE086D5EFC6EFE53" box="[786,922,364,389]" italics="true" pageId="2" pageNumber="61">C. wothuja</emphasis>
</taxonomicName>
(
<bibRefCitation id="0B1A4B5DFFFEFF96BEAA6D59FB67FE53" author="Grant" box="[944,1171,363,389]" pageId="2" pageNumber="66" refString="Grant, T., &amp; F. Castro. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of Cordillera Occidental of Colombia. Journal of Herpetology, 32 (3), 378 - 392." type="journal article" year="1998">
Grant
<emphasis id="5DFFEABEFFFEFF96B91E6D5EFBB2FE53" box="[1028,1094,364,389]" italics="true" pageId="2" pageNumber="61">et al.</emphasis>
1998
</bibRefCitation>
;
<bibRefCitation id="0B1A4B5DFFFEFF96B9BC6D59FE4FFE7B" author="Myers" pageId="2" pageNumber="67" refString="Myers, C. W. &amp; Donnelly, M. A. (1997) A Tepui herpetofauna on a granitic mountain (Tamacuari) in the borderland between Venezuela and Brazil: Report from the Phelps Tapirapeco Expedition. American Museum Novitates, 3213, 1 - 71." type="journal article" year="1997">Myers and Donnelly 1997</bibRefCitation>
;
<bibRefCitation id="0B1A4B5DFFFEFF96BCD36DA1FCFCFE7B" author="Barrio-Amoros" box="[457,776,403,429]" pageId="2" pageNumber="66" refString="Barrio-Amoros, C. L., Fuentes O., &amp; Rivas, G. (2004) Two new species of Colostethus (Anura: Dendrobatidae) from the Venezuelan Guayana. Salamandra, Rheinbach, 40 (3 / 4), 1 - 18." type="journal article" year="2004">
Barrio­Amorós
<emphasis id="5DFFEABEFFFEFF96BF9F6DA1FD35FE7A" box="[645,705,403,428]" italics="true" pageId="2" pageNumber="61">et al.</emphasis>
2004
</bibRefCitation>
),
<taxonomicName id="A88B4D2FFFFEFF96BE076DA1FC44FE7A" box="[797,944,403,428]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="praderioi">
<emphasis id="5DFFEABEFFFEFF96BE076DA1FC44FE7A" box="[797,944,403,428]" italics="true" pageId="2" pageNumber="61">C. praderioi</emphasis>
</taxonomicName>
and
<taxonomicName id="A88B4D2FFFFEFF96BEF16DA1FB9AFE7A" box="[1003,1134,403,428]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="roraima">
<emphasis id="5DFFEABEFFFEFF96BEF16DA1FB9AFE7A" box="[1003,1134,403,428]" italics="true" pageId="2" pageNumber="61">C. roraima</emphasis>
</taxonomicName>
(T. Grant, pers. comm.). Species from the Venezuelan Guayana lacking the MLP are
<taxonomicName id="A88B4D2FFFFEFF96B9796D89FADEFE02" box="[1123,1322,443,469]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="brunneus">
<emphasis id="5DFFEABEFFFEFF96B9796D89FB82FE02" box="[1123,1142,443,468]" italics="true" pageId="2" pageNumber="61">C</emphasis>
. aff.
<emphasis id="5DFFEABEFFFEFF96B9A76D89FADEFE02" box="[1213,1322,443,468]" italics="true" pageId="2" pageNumber="61">brunneus</emphasis>
</taxonomicName>
(probably a similar species to
<taxonomicName id="A88B4D2FFFFEFF96BF6A6DD1FCF5FE2A" box="[624,769,483,508]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="brunneus">
<emphasis id="5DFFEABEFFFEFF96BF6A6DD1FCF5FE2A" box="[624,769,483,508]" italics="true" pageId="2" pageNumber="61">C. brunneus</emphasis>
</taxonomicName>
: see
<bibRefCitation id="0B1A4B5DFFFEFF96BE5A6DD0FC13FE2A" author="Morales" box="[832,999,482,508]" pageId="2" pageNumber="67" refString="Morales, V. R. (2000) Sistematica y biogeografia del grupo trilineatus (Amphibia, Anura, Dendrobatidae, Colostethus) con descripcion de once nuevas especies. Publicaciones de la Asociacion de amigos de Donana, 13, 5 - 59." type="journal article" year="2000">Morales 2000</bibRefCitation>
),
<taxonomicName id="A88B4D2FFFFEFF96BEE76DD1FB07FE2A" box="[1021,1267,482,508]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="marchesianus">
<emphasis id="5DFFEABEFFFEFF96BEE76DD1FBECFE2A" box="[1021,1048,483,508]" italics="true" pageId="2" pageNumber="61">C.</emphasis>
aff.
<emphasis id="5DFFEABEFFFEFF96B94B6DD1FB07FE2A" box="[1105,1267,483,508]" italics="true" pageId="2" pageNumber="61">marchesianus</emphasis>
</taxonomicName>
(see
<bibRefCitation id="0B1A4B5DFFFEFF96BC126E38FE5EFDF2" author="Morales" box="[264,426,522,548]" pageId="2" pageNumber="67" refString="Morales, V. R. (1994) Taxonomia de algunos Colostethus (Anura: Dendrobatidae) de Sudamerica, con descripcion de dos especies nuevas. Revista Espanola Herpetologia, 8, 95 - 103." type="journal article" year="1994">Morales 1994</bibRefCitation>
and
<bibRefCitation id="0B1A4B5DFFFEFF96BCF96E38FD21FDF2" author="Caldwell" box="[483,725,522,548]" pageId="2" pageNumber="66" refString="Caldwell, J. P., Lima, A. P. &amp; Keller, C. (2002) Redescription of Colostethus marchesianus (Melin, 1941) from its type locality. Copeia, 2002, 157 - 165." type="journal article" year="2002">
Caldwell
<emphasis id="5DFFEABEFFFEFF96BF4E6E38FD7BFDF5" box="[596,655,522,547]" italics="true" pageId="2" pageNumber="61">et al.</emphasis>
2002
</bibRefCitation>
), and
<taxonomicName id="A88B4D2FFFFEFF96BE016E38FAF6FDF2" authority="Myers and Donnelly 2001" authorityName="Myers and Donnelly" authorityYear="2001" box="[795,1282,522,548]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="undulatus">
<emphasis id="5DFFEABEFFFEFF96BE016E38FC45FDF5" box="[795,945,522,547]" italics="true" pageId="2" pageNumber="61">C. undulatus</emphasis>
(
<bibRefCitation id="0B1A4B5DFFFEFF96BEA56E38FB0DFDF2" author="Myers" box="[959,1273,522,548]" pageId="2" pageNumber="67" refString="Myers, C. W. &amp; Donnelly, M. A. (2001) Herpetofauna of the Yutaje-Corocoro Massif, Venezuela: Second report from the Robert G. Goelet American Museum-Terramar Expedition to the Northwestern tepuis. Bulletin of the American Museum of Natural History, 261, 1 - 85." type="journal article" year="2001">Myers and Donnelly 2001</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="27916527FFFEFF91BC226E00FB6DFA30" lastPageId="5" lastPageNumber="64" pageId="2" pageNumber="61" type="description">
<paragraph id="6F3436ACFFFEFF96BC226E00FE6DFA60" blockId="2.[264,1324,283,1899]" pageId="2" pageNumber="61">
<taxonomicName id="A88B4D2FFFFEFF96BC226E00FDD2FD9D" box="[312,550,562,587]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="breweri">
<emphasis id="5DFFEABEFFFEFF96BC226E00FDD2FD9D" box="[312,550,562,587]" italics="true" pageId="2" pageNumber="61">Colostethus breweri</emphasis>
</taxonomicName>
can be also distinguished from those species having scanty toe webbing (
<emphasis id="5DFFEABEFFFEFF96BC656E68FCC4FDA5" box="[383,816,602,627]" italics="true" pageId="2" pageNumber="61">
<taxonomicName id="A88B4D2FFFFEFF96BC656E68FE05FDA5" box="[383,497,602,627]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="beebei">C. beebei</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFEFF96BCE46E68FD65FDA5" box="[510,657,602,627]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="brunneus">C. brunneus</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFEFF96BF846E68FCC4FDA5" box="[670,816,602,627]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="praderioi">C. praderioi</taxonomicName>
</emphasis>
and
<taxonomicName id="A88B4D2FFFFEFF96BE706E68FC18FDA5" box="[874,1004,602,627]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="roraima">
<emphasis id="5DFFEABEFFFEFF96BE706E68FC18FDA5" box="[874,1004,602,627]" italics="true" pageId="2" pageNumber="61">C. roraima</emphasis>
</taxonomicName>
) because of its moderately developed toe webbing. It is distinguishable from other species from the
<collectingCountry id="179C763CFFFEFF96B99B6EB0FB22FD4A" box="[1153,1238,642,668]" name="Guyana" pageId="2" pageNumber="61">Guiana</collectingCountry>
Shield that also present MLP and moderate toe webbing by the following character (those of
<taxonomicName id="A88B4D2FFFFEFF96B80A6E9BFE96FD3C" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="breweri">
<emphasis id="5DFFEABEFFFEFF96B80A6E9BFE96FD3C" italics="true" pageId="2" pageNumber="61">C. breweri</emphasis>
</taxonomicName>
in parentheses).
<taxonomicName id="A88B4D2FFFFEFF96BF296EE3FC92FD3C" box="[563,870,721,746]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="ayarzaguenai">
<emphasis id="5DFFEABEFFFEFF96BF296EE3FC92FD3C" box="[563,870,721,746]" italics="true" pageId="2" pageNumber="61">Colostethus ayarzaguenai</emphasis>
</taxonomicName>
has a dorsally­rounded snout (nearly truncate), Finger I shorter than II (equal); and an uniform pattern without dorsal spots (consistent pattern).
<taxonomicName id="A88B4D2FFFFEFF96BCE16F13FCE9FCEC" box="[507,797,801,826]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="degranvillei">
<emphasis id="5DFFEABEFFFEFF96BCE16F13FCE9FCEC" box="[507,797,801,826]" italics="true" pageId="2" pageNumber="61">Colostethus degranvillei</emphasis>
</taxonomicName>
has Finger I shorter than II (equal), oblique lateral stripe absent (present), a post tympanic white bar (absent) and ventral surfaces brown with white spots (whitish or yellow).
<taxonomicName id="A88B4D2FFFFEFF96BE0D6F42FBCFFC5F" box="[791,1083,880,905]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="guanayensis">
<emphasis id="5DFFEABEFFFEFF96BE0D6F42FBCFFC5F" box="[791,1083,880,905]" italics="true" pageId="2" pageNumber="61">Colostethus guanayensis</emphasis>
</taxonomicName>
has Finger I shorter than II (equal), dark ventral coloration (dirty white or yellow).
<taxonomicName id="A88B4D2FFFFEFF96BEFD6FAAFADFFC67" box="[999,1323,920,945]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="murisipanensis">
<emphasis id="5DFFEABEFFFEFF96BEFD6FAAFADFFC67" box="[999,1323,920,945]" italics="true" pageId="2" pageNumber="61">Colostethus murisipanensis</emphasis>
</taxonomicName>
has a Finger I shorter than II (equal), no oblique lateral stripe (present), dark ventral colouration (dirty white or yellow).
<taxonomicName id="A88B4D2FFFFEFF96BFCD6FDAFC27FBD7" box="[727,979,1000,1025]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="parimae">
<emphasis id="5DFFEABEFFFEFF96BFCD6FDAFC27FBD7" box="[727,979,1000,1025]" italics="true" pageId="2" pageNumber="61">Colostethus parimae</emphasis>
</taxonomicName>
has non consistent pattern (consistent), a dorsally rounded snout (truncate), Finger I shorter than II (equal).
<taxonomicName id="A88B4D2FFFFEFF96BC126805FDF1FB86" box="[264,517,1079,1104]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="parkerae">
<emphasis id="5DFFEABEFFFEFF96BC126805FDF1FB86" box="[264,517,1079,1104]" italics="true" pageId="2" pageNumber="61">Colostethus parkerae</emphasis>
</taxonomicName>
has a rounded snout in dorsal view (truncate), oblique lateral stripe absent (present).
<taxonomicName id="A88B4D2FFFFEFF96BCF8686DFD0FFBAE" box="[482,763,1119,1144]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="sanmartini">
<emphasis id="5DFFEABEFFFEFF96BCF8686DFD0FFBAE" box="[482,763,1119,1144]" italics="true" pageId="2" pageNumber="61">Colostethus sanmartini</emphasis>
</taxonomicName>
has tympanum large, 57% of eye diameter (indistinct), pale dorsolateral stripes (absent).
<taxonomicName id="A88B4D2FFFFEFF96BE3768B4FBE1FB49" box="[813,1045,1158,1183]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="shrevei">
<emphasis id="5DFFEABEFFFEFF96BE3768B4FBE1FB49" box="[813,1045,1158,1183]" italics="true" pageId="2" pageNumber="61">Colostethus shrevei</emphasis>
</taxonomicName>
is larger, up to
<quantity id="A8739B49FFFEFF96B9CC68B4FADEFB76" box="[1238,1322,1158,1184]" metricMagnitude="-2" metricUnit="m" metricValue="3.6" pageId="2" pageNumber="61" unit="mm" value="36.0">36 mm</quantity>
(up to 23.8), Finger I shorter than II (equal).
<taxonomicName id="A88B4D2FFFFEFF96BE0C689CFBA6FB11" box="[790,1106,1198,1223]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="tamacuarensis">
<emphasis id="5DFFEABEFFFEFF96BE0C689CFBA6FB11" box="[790,1106,1198,1223]" italics="true" pageId="2" pageNumber="61">Colostethus tamacuarensis</emphasis>
</taxonomicName>
has anal tubercles (absent), Finger III slightly swollen in males (not swollen).
<taxonomicName id="A88B4D2FFFFEFF96BECC68E4FB10FB39" box="[982,1252,1238,1263]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="tepuyensis">
<emphasis id="5DFFEABEFFFEFF96BECC68E4FB10FB39" box="[982,1252,1238,1263]" italics="true" pageId="2" pageNumber="61">Colostethus tepuyensis</emphasis>
</taxonomicName>
has a rounded snout in dorsal view (truncate), Finger I shorter than II (equal), oblique lateral stripe absent (present).
<taxonomicName id="A88B4D2FFFFEFF96BF346917FCE3FAE8" box="[558,791,1317,1342]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="triunfo">
<emphasis id="5DFFEABEFFFEFF96BF346917FCE3FAE8" box="[558,791,1317,1342]" italics="true" pageId="2" pageNumber="61">Colostethus triunfo</emphasis>
</taxonomicName>
has smooth dorsal skin (granular in life), metacarpal tubercle triangular (rounded), tarsal fold short, 1/3 of the tarsus (1/2).
<taxonomicName id="A88B4D2FFFFEFF96BC126947FDF6FA58" box="[264,514,1397,1422]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="61" phylum="Chordata" rank="species" species="wothuja">
<emphasis id="5DFFEABEFFFEFF96BC126947FDF6FA58" box="[264,514,1397,1422]" italics="true" pageId="2" pageNumber="61">Colostethus wothuja</emphasis>
</taxonomicName>
has a distinct tympanum (indistinct), and no particular pattern (consistent).
</paragraph>
<paragraph id="6F3436ACFFFEFF96BC2269F6FBD0F983" blockId="2.[264,1324,283,1899]" pageId="2" pageNumber="61">
<emphasis id="5DFFEABEFFFEFF96BC2269F6FDADFA08" bold="true" box="[312,601,1476,1502]" pageId="2" pageNumber="61">
Description of
<typeStatus id="B030880EFFFEFF96BCF469F6FDADFA08" box="[494,601,1476,1502]" pageId="2" pageNumber="61" type="holotype">holotype</typeStatus>
</emphasis>
—Dorsal and ventral skin smooth in preservative; shagreen in life with small but protuberant tubercles on the posterior part of the dorsum. Dorsal skin forming usually a well­defined rounded, posteriorly projecting flap well above vent, which opens at upper level of thighs; no anal tubercles; anal sheath absent.
</paragraph>
<paragraph id="6F3436ACFFFEFF96BC226A51FCA9F8BD" blockId="2.[264,1324,283,1899]" pageId="2" pageNumber="61">Head slightly longer than wide, head width (between angles of jaws) 33.6% of SVL. Snout protruding in profile, nearly truncate in dorsal and ventral views. Nares situated near the tip of the snout and directed slightly posterolaterally; nares visible from the front, barely or not visible dorsally, but well visible from below. Canthus rostralis rounded, rather indistinct; loreal region barely concave. Interorbital region slightly wider than upper eyelid. Snout slightly longer than eye diameter. Tympanum barely visible; positioned close behind eye and low, nearly touching angle of jaws.</paragraph>
<caption id="3BF46624FFFFFF97BC126841FB05FB5D" box="[264,1265,1139,1163]" httpUri="https://zenodo.org/record/171518/files/figure.png" pageId="3" pageNumber="62" targetBox="[273,1314,329,1112]" targetPageId="3">
<paragraph id="6F3436ACFFFFFF97BC126841FB05FB5D" blockId="3.[264,1265,1139,1163]" box="[264,1265,1139,1163]" pageId="3" pageNumber="62">
<emphasis id="5DFFEABEFFFFFF97BC126841FE7EFB5D" bold="true" box="[264,394,1139,1163]" pageId="3" pageNumber="62">FIGURE 1.</emphasis>
Female holotype of
<taxonomicName id="A88B4D2FFFFFFF97BF7D6846FCCAFB5D" box="[615,830,1140,1163]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="62" phylum="Chordata" rank="species" species="breweri">
<emphasis id="5DFFEABEFFFFFF97BF7D6846FCCAFB5D" box="[615,830,1140,1163]" italics="true" pageId="3" pageNumber="62">Colostethus breweri</emphasis>
</taxonomicName>
in life. Photo by Charles Brewer­Carías.
</paragraph>
</caption>
<paragraph id="6F3436ACFFFFFF97BC22688BFB93FA4D" blockId="3.[264,1325,1209,1875]" pageId="3" pageNumber="62">Hand length moderate, 27.7 % of SVL. Relative lengths of appressed fingers III&gt;IV&gt;I=II; Finger I equal in length to Finger II. Discs of all fingers moderately expanded; Finger III disc 1.5 times width of distal end of adjacent phalanx. Base of palm with large, nearly round, median palmar tubercle; oval inner thenar tubercle on base of Finger I; one subarticular tubercle on Fingers I and II, and two on Fingers III and IV, distal tubercles small, indistinct; all tubercles low, round. No fringes on fingers.</paragraph>
<paragraph id="6F3436ACFFFFFF97BC22699BFC25F885" blockId="3.[264,1325,1209,1875]" pageId="3" pageNumber="62">
<collectingCountry id="179C763CFFFFFF97BC22699BFE87FA15" box="[312,371,1449,1475]" name="India" pageId="3" pageNumber="62">Hind</collectingCountry>
limbs of moderate length; shank 49.5 % of SVL. Relative lengths of appressed toes IV&gt;III&gt;V&gt;II&gt;I; first toe reaching when appressed half subarticular tubercle of second toe. Toe discs moderately expanded; toe IV disc 1.4 times width of distal end of adjacent phalanx. Feet moderately webbed, the web distally continuous with a narrow fringe on all toes, including external edges of toes I and V. Webbing formula
<date id="1B35106CFFFFFF97B9186A7BFB99F9B5" box="[1026,1133,1609,1635]" pageId="3" pageNumber="62">I 12 II 1</date>
½3
<date id="1B35106CFFFFFF97B9B76A7BFB15F9B5" box="[1197,1249,1609,1635]" pageId="3" pageNumber="62">III 2</date>
½
<date id="1B35106CFFFFFF97BC126A43FEC8F95D" box="[264,316,1649,1675]" pageId="3" pageNumber="62">IV 3</date>
½1½ V. One to three non protuberant subarticular tubercles on toes as follows: one on toes I and II, two on toes III and V, and three on toe IV (distal tubercle ill defined). Sole with two metatarsal tubercles, similarly sized, a round outer metatarsal tubercle, and an elliptical inner metatarsal tubercle. A narrow tarsal fold or keel, straight, extending almost half the length of tarsus, distally continuous with the narrow fringe on free (preaxial) edge of toe I; fringe distinctly raised proximally, not tubercle­like.
</paragraph>
<paragraph id="6F3436ACFFF8FF90BC226D2EFB88FE88" blockId="4.[264,1323,284,350]" pageId="4" pageNumber="63">Maxillary teeth present. Tongue longer than wide, elliptical, posterior half free; MLP very small, longer than wide, positioned on the anterior third of the tongue.</paragraph>
<caption id="3BF46624FFF8FF90BC1269FFFCD1F9FC" httpUri="https://zenodo.org/record/171519/files/figure.png" pageId="4" pageNumber="63" targetBox="[409,1177,393,1473]" targetPageId="4">
<paragraph id="6F3436ACFFF8FF90BC1269FFFCD1F9FC" blockId="4.[264,1323,1485,1578]" pageId="4" pageNumber="63">
<emphasis id="5DFFEABEFFF8FF90BC1269FFFE7FFA33" bold="true" box="[264,395,1485,1509]" pageId="4" pageNumber="63">FIGURE 2.</emphasis>
View of the type locality (Cueva de El Fantasma, Aprada tepui) from inside. Note the size of the two helicopters at the entrance. This is the first geographic report and photograph of such immense “cave”. Photo by Charles Brewer­Carías.
</paragraph>
</caption>
<paragraph id="6F3436ACFFF8FF91BC226A6FFE94FE78" blockId="4.[264,1323,1629,1895]" lastBlockId="5.[264,1325,284,1876]" lastPageId="5" lastPageNumber="64" pageId="4" pageNumber="63">
<emphasis id="5DFFEABEFFF8FF90BC226A6FFE14F9A1" bold="true" box="[312,480,1629,1655]" pageId="4" pageNumber="63">Colour in life</emphasis>
—Dorsal ground colour pale brown with diffuse markings on the back, consisting of a dark brown interorbital bar, straight, usually with an apex at the posterior margin, a moderately large chevron between shoulders, three spots at midbody, forming an inverted V; and a single small and median posterior spot near the end of the body. Canthal stripe dark brown; supratympanic stripe black, and very distinct. In the
<typeStatus id="B030880EFFF8FF90B9556ACFFB41F8C1" box="[1103,1205,1789,1815]" pageId="4" pageNumber="63" type="holotype">holotype</typeStatus>
the upper lip bar is immaculate white, containing the indistinct tympanum. Upper lips also white in the other specimens. Supratympanic stripe is continuous above the arm, widening at mid body, forming a dark brown to black flank, where the white oblique lateral stripe is present. Flanks darker than dorsum but without definite borders. Upper flank dark brown. Throat, chest, and belly immaculate yellow; inferior surfaces of thighs dark orange. Iris bronze.
</paragraph>
<paragraph id="6F3436ACFFF9FF91BC226D8EFD9EFE28" blockId="5.[264,1325,284,1876]" pageId="5" pageNumber="64">Upper arms and forearms pale brown, the latter with dark crossbars. Thighs with illdefined dark brown crossbars.</paragraph>
<paragraph id="6F3436ACFFF9FF91BC226E3EFAE9FD98" blockId="5.[264,1325,284,1876]" pageId="5" pageNumber="64">After one year and a half in preservative, dorsal ground colour became dark brown; flanks dark brown; ventral surfaces white. The rest of the body remains without changes.</paragraph>
<paragraph id="6F3436ACFFF9FF91BC226E69FB20FD48" blockId="5.[264,1325,284,1876]" pageId="5" pageNumber="64">
<emphasis id="5DFFEABEFFF9FF91BC226E69FD70FDA3" bold="true" box="[312,644,603,629]" pageId="5" pageNumber="64">
Measurements of
<typeStatus id="B030880EFFF9FF91BF026E69FD70FDA3" box="[536,644,603,629]" pageId="5" pageNumber="64" type="holotype">holotype</typeStatus>
</emphasis>
(in mm)—SVL: 23.8; SL: 11.8; ThL: 12; HeL: 8.3; HW: 8; UEW: 2; IOD: 2.2; ED: 3; TD: 1; ETS: 3.2; FD: 1; 4TD: 1; 1FiL: 3.2; 2FiL: 3.3.
</paragraph>
<paragraph id="6F3436ACFFF9FF91BC226E9EFD34FBA8" blockId="5.[264,1325,284,1876]" pageId="5" pageNumber="64">
<emphasis id="5DFFEABEFFF9FF91BC226E9EFE5FFD10" bold="true" box="[312,427,684,710]" pageId="5" pageNumber="64">Variation</emphasis>
—All specimens have a similar dorsal colour pattern, which is more or less conspicuous. It is most distinct in MHNLS 17045 (a male), and the juvenile (MHNLS 17047) and is less defined (but still detectable) in the female
<typeStatus id="B030880EFFF9FF91BEFA6ECEFBB2FCC0" box="[992,1094,764,790]" pageId="5" pageNumber="64" type="holotype">holotype</typeStatus>
and male MHNLS 17046. The oblique lateral stripe occurs in all specimens, although varying in shape (always short), and never well defined. It is best defined in MHNLS 17045, or fragmented into three small white spots, as in the
<typeStatus id="B030880EFFF9FF91BFD46F46FCCCFC58" box="[718,824,884,910]" pageId="5" pageNumber="64" type="holotype">holotype</typeStatus>
. Males differ from the single female in at least three external characters (these may be due to sexual dimorphism rather than individual variation): First, the belly of adult males is immaculate white, whereas that of the adult female is immaculate yellow; second, the presence of short vocal slits, extending from near tongue insertion to nearly the end of tongue; and third, the presence of a larger lingual process, longer than wide; this last character was also observed in
<taxonomicName id="A88B4D2FFFF9FF91B9BA680EFD48FBA8" authority="Barrio-Amoros et al. 2004" authorityName="Barrio-Amoros et al." authorityYear="2004" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="5" pageNumber="64" phylum="Chordata" rank="species" species="wothuja">
<emphasis id="5DFFEABEFFF9FF91B9BA680EFE92FBAB" italics="true" pageId="5" pageNumber="64">Colostethus wothuja</emphasis>
(
<bibRefCitation id="0B1A4B5DFFF9FF91BC6F6856FD40FBA8" author="Barrio-Amoros" box="[373,692,1124,1150]" pageId="5" pageNumber="66" refString="Barrio-Amoros, C. L., Fuentes O., &amp; Rivas, G. (2004) Two new species of Colostethus (Anura: Dendrobatidae) from the Venezuelan Guayana. Salamandra, Rheinbach, 40 (3 / 4), 1 - 18." type="journal article" year="2004">
Barrio­Amorós
<emphasis id="5DFFEABEFFF9FF91BF286856FD99FBAB" box="[562,621,1124,1149]" italics="true" pageId="5" pageNumber="64">et al.</emphasis>
2004
</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="6F3436ACFFF9FF91BC2268BEFB63FA90" blockId="5.[264,1325,284,1876]" pageId="5" pageNumber="64">
In preservative,
<typeStatus id="B030880EFFF9FF91BF1968BEFD9DFB70" box="[515,617,1164,1190]" pageId="5" pageNumber="64" type="holotype">holotype</typeStatus>
and MHNLS 17046 dorsal ground colour became dark brown with diffuse markings; MHNLS 17045 and juvenile (MHNLS 17047) pale gray with conspicuous dark gray markings; flanks dark brown to dark gray. Black arm band (
<emphasis id="5DFFEABEFFF9FF91BC086936FEA7FACB" box="[274,339,1284,1309]" italics="true" pageId="5" pageNumber="64">sensu</emphasis>
<bibRefCitation id="0B1A4B5DFFF9FF91BC476936FD86FAC8" author="Grant" box="[349,626,1284,1310]" pageId="5" pageNumber="66" refString="Grant, T., &amp; F. Castro. (1998) The cloud forest Colostethus (Anura, Dendrobatidae) of a region of Cordillera Occidental of Colombia. Journal of Herpetology, 32 (3), 378 - 392." type="journal article" year="1998">Grant and Castro 1998</bibRefCitation>
) absent, but in all specimens except the juvenile, a black longitudinal stripe on the anterior surface of the arms and forearms is present.
</paragraph>
<paragraph id="6F3436ACFFF9FF91BC226966FB6DFA30" blockId="5.[264,1325,284,1876]" pageId="5" pageNumber="64">Males (MHNLS 17045 and 17046) with dirty white throat (white with profusion of melanophores) and larger white spots especially at edges of throat; anterior part of chest also with melanophores in MHNLS 17045 but not in MHNLS 17046; posterior part of chest, belly and ventral surfaces of the thighs immaculate white. Testes white.</paragraph>
</subSubSection>
<subSubSection id="27916527FFF9FF92BC2269C6FC03FD48" lastPageId="6" lastPageNumber="65" pageId="5" pageNumber="64" type="distribution">
<paragraph id="6F3436ACFFF9FF91BC2269C6FBA8F882" blockId="5.[264,1325,284,1876]" pageId="5" pageNumber="64">
<emphasis id="5DFFEABEFFF9FF91BC2269C6FE25F9D8" bold="true" box="[312,465,1524,1550]" pageId="5" pageNumber="64">Distribution</emphasis>
—The species is known only from one locality at Aprada tepui, a flattopped sandstone mesa (tepui) of Guayana highlands in
<collectingCountry id="179C763CFFF9FF91BEB16A2EFBD3F9E0" box="[939,1063,1564,1590]" name="Venezuela" pageId="5" pageNumber="64">Venezuela</collectingCountry>
, at 660 masl (
<figureCitation id="F7B02A29FFF9FF91B9C86A2EFAE8F9E0" box="[1234,1308,1564,1590]" captionStart="FIGURE 2" captionStartId="4.[264,368,1485,1509]" captionTargetBox="[409,1177,393,1473]" captionTargetId="figure@4.[409,1177,392,1475]" captionTargetPageId="4" captionText="FIGURE 2. View of the type locality (Cueva de El Fantasma, Aprada tepui) from inside. Note the size of the two helicopters at the entrance. This is the first geographic report and photograph of such immense “ cave ”. Photo by Charles Brewer­Carías." httpUri="https://zenodo.org/record/171519/files/figure.png" pageId="5" pageNumber="64">Fig. 2</figureCitation>
). This locality lies in medium to tall, evergreen, basimontane and lower montane forests (
<bibRefCitation id="0B1A4B5DFFF9FF91BC0A6A5EFDD2F950" author="Huber" box="[272,550,1644,1670]" pageId="5" pageNumber="67" refString="Huber, O. &amp; Alarcon, C. (1988) Mapa de Vegetacion de Venezuela, 1: 2.000.000. Ministerio del Ambiente y de los Recursos Naturales Renovables and The Nature Conservancy, Caracas, Venezuela." type="book chapter" year="1988">Huber &amp; Alarcón 1988</bibRefCitation>
). To found the
<typeStatus id="B030880EFFF9FF91BFC26A5EFCFEF950" box="[728,778,1644,1670]" pageId="5" pageNumber="64">type</typeStatus>
series inside a “cave” must not be significant, as this “Cueva de El Fantasma” (
<figureCitation id="F7B02A29FFF9FF91BF966AA6FD27F978" box="[652,723,1684,1710]" captionStart="FIGURE 2" captionStartId="4.[264,368,1485,1509]" captionTargetBox="[409,1177,393,1473]" captionTargetId="figure@4.[409,1177,392,1475]" captionTargetPageId="4" captionText="FIGURE 2. View of the type locality (Cueva de El Fantasma, Aprada tepui) from inside. Note the size of the two helicopters at the entrance. This is the first geographic report and photograph of such immense “ cave ”. Photo by Charles Brewer­Carías." httpUri="https://zenodo.org/record/171519/files/figure.png" pageId="5" pageNumber="64">Fig. 2</figureCitation>
) is a huge, collapsed steep gorge and is not strictly a cave. The maximum altitude of Aprada is at
<quantity id="A8739B49FFF9FF91BE276A8DFC6FF90F" box="[829,923,1727,1753]" metricMagnitude="3" metricUnit="m" metricValue="2.5" pageId="5" pageNumber="64" unit="m" value="2500.0">2500 m</quantity>
, with a tepui surface of
<quantity id="A8739B49FFF9FF91B9D86A8DFAD6F90F" box="[1218,1314,1727,1753]" metricMagnitude="3" metricUnit="m" metricValue="4.37" pageId="5" pageNumber="64" unit="km" value="4.37">4.37 km</quantity>
2 (
<bibRefCitation id="0B1A4B5DFFF9FF91BC0A6AD8FE56F8D2" author="Huber" box="[272,418,1770,1796]" pageId="5" pageNumber="67" refString="Huber, O. (1995) Vegetation, 97 - 160 pp. In: Flora of the Venezuelan Guayana. Vol I: Introduction. (P. E. Berry. B. K. Holst &amp; K. Yatskievych (Eds.), Missouri Botanical Gardens, St. Louis. Timber Press." type="book chapter" year="1995">Huber 1995</bibRefCitation>
), although its slopes embraces at
<quantity id="A8739B49FFF9FF91BE2C6AD8FC65F8D2" box="[822,913,1770,1796]" metricMagnitude="3" metricUnit="m" metricValue="1.0" pageId="5" pageNumber="64" unit="m" value="1000.0">1000 m</quantity>
of altitude a wider area including the Chimantá massif and even the whole Gran Sabana area. The species could be an endemic, or more widely expanded through uplands of the Gran Sabana.
</paragraph>
<paragraph id="6F3436ACFFFAFF92BC226D2EFEBEFE28" blockId="6.[264,1324,284,1870]" pageId="6" pageNumber="65">
Aprada tepui was first explored by an expedition leaded by C. Brewer­Carías on
<date id="1B35106CFFFAFF92BC126D76FE12FE88" box="[264,486,324,350]" pageId="6" pageNumber="65" value="1978-02-25">February 25, 1978</date>
, accompanied by Roy McDiarmid, as herpetologist, Luis José Joly as enthomologist, Julian Steyermark as botanist and G.C.K. Dunsterville and his wife Nora as orchidologists (
<bibRefCitation id="0B1A4B5DFFFAFF92BCDA6DA6FD68FE78" author="Dunsterville" box="[448,668,404,430]" pageId="6" pageNumber="66" refString="Dunsterville, G. C. K. &amp; Dunsterville, N. (1979) Tepui Top hopping by helicopter. American Orchid Society Bulletin, 48 (3), 222 - 228." type="journal article" year="1979">Dunsterville 1979</bibRefCitation>
). This paper becomes the first written reference about this huge &quot;cave&quot; at N
<geoCoordinate id="0ABF506BFFFAFF92BF146D8EFD47FE00" box="[526,691,444,470]" direction="west" orientation="longitude" pageId="6" pageNumber="65" precision="1" value="-5.458">05º 27.480W</geoCoordinate>
62º 27.588 on the northwestern slope of the Aprada tepui.
</paragraph>
<paragraph id="6F3436ACFFFAFF92BC226E39FC03FD48" blockId="6.[264,1324,284,1870]" pageId="6" pageNumber="65">
<emphasis id="5DFFEABEFFFAFF92BC226E39FE08FDF3" bold="true" box="[312,508,523,549]" pageId="6" pageNumber="65">Natural history</emphasis>
—This species is a fast­moving frog that lives along creeks and in quiet pools along small streams on the slopes of Aprada­tepui. An undescribed species of
<taxonomicName id="A88B4D2FFFFAFF92BC126E6EFE28FDA3" box="[264,476,604,629]" class="Amphibia" family="Eleutherodactylidae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="genus">
<emphasis id="5DFFEABEFFFAFF92BC126E6EFE28FDA3" box="[264,476,604,629]" italics="true" pageId="6" pageNumber="65">Eleutherodactylus</emphasis>
</taxonomicName>
and the lizard
<taxonomicName id="A88B4D2FFFFAFF92BFA86E6EFB7FFDA0" box="[690,1163,604,630]" class="Reptilia" family="Gymnophthalmidae" genus="Neusticurus" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="rudis">
<emphasis id="5DFFEABEFFFAFF92BFA86E6EFC7FFDA3" box="[690,907,604,629]" italics="true" pageId="6" pageNumber="65">Neusticurus rudis</emphasis>
(Gymnophtalmidae)
</taxonomicName>
were found syntopically. Tadpoles and call of the new species are unknown.
</paragraph>
</subSubSection>
<subSubSection id="27916527FFFAFF92BC226E9EFD64FCC0" pageId="6" pageNumber="65" type="etymology">
<paragraph id="6F3436ACFFFAFF92BC226E9EFD64FCC0" blockId="6.[264,1324,284,1870]" pageId="6" pageNumber="65">
<emphasis id="5DFFEABEFFFAFF92BC226E9EFE4BFD10" bold="true" box="[312,447,684,710]" pageId="6" pageNumber="65">Etymology</emphasis>
—I dedicate this frog to the indefatigable explorer of the Venezuelan Guayana, Charles Brewer­Carías, in recognition for his endless support of biological research in the
<collectingCountry id="179C763CFFFAFF92BCA06ECEFDFBFCC0" box="[442,527,764,790]" name="Guyana" pageId="6" pageNumber="65">Guiana</collectingCountry>
highlands.
</paragraph>
</subSubSection>
<subSubSection id="27916527FFFAFF93BC226F16FC0FFDD7" lastPageId="7" lastPageNumber="66" pageId="6" pageNumber="65" type="discussion">
<paragraph id="6F3436ACFFFAFF92BC226F16FC2DFAC8" blockId="6.[264,1324,284,1870]" pageId="6" pageNumber="65">
<emphasis id="5DFFEABEFFFAFF92BC226F16FE48FCE8" bold="true" box="[312,444,804,830]" pageId="6" pageNumber="65">Discussion</emphasis>
—Currently, knowledge of Guayanan
<taxonomicName id="A88B4D2FFFFAFF92BEB06F16FBC1FCEB" box="[938,1077,804,829]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="genus">
<emphasis id="5DFFEABEFFFAFF92BEB06F16FBC1FCEB" box="[938,1077,804,829]" italics="true" pageId="6" pageNumber="65">Colostethus</emphasis>
</taxonomicName>
is increasing, and several characters have been observed recurrently for most of the species. The most apparent is the presence of a MLP (Grant et al. 1997); although it is not exclusive of Guayanan
<taxonomicName id="A88B4D2FFFFAFF92BC966FAEFDE2FC63" box="[396,534,924,949]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="genus">
<emphasis id="5DFFEABEFFFAFF92BC966FAEFDE2FC63" box="[396,534,924,949]" italics="true" pageId="6" pageNumber="65">Colostethus</emphasis>
</taxonomicName>
, it seems most of the species share it. Furthermore, all species known to have a MLP (
<emphasis id="5DFFEABEFFFAFF92BF286FF6FC49FC0B" box="[562,957,964,989]" italics="true" pageId="6" pageNumber="65">
<taxonomicName id="A88B4D2FFFFAFF92BF286FF6FCD6FC0B" box="[562,802,964,989]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="breweri">Colostethus breweri</taxonomicName>
<taxonomicName id="A88B4D2FFFFAFF92BE376FF6FC49FC0B" box="[813,957,964,989]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="parkerae">C. parkerae</taxonomicName>
</emphasis>
,
<emphasis id="5DFFEABEFFFAFF92BECA6FF6FB08FC0B" box="[976,1276,964,989]" italics="true" pageId="6" pageNumber="65">
<taxonomicName id="A88B4D2FFFFAFF92BECA6FF6FB93FC0B" box="[976,1127,964,989]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="praderioi">C. praderioi</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFAFF92B9626FF6FB08FC0B" box="[1144,1276,964,989]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="roraima">C. roraima</taxonomicName>
</emphasis>
,
<taxonomicName id="A88B4D2FFFFAFF92B80A6FF6FEAFFBD3" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="shrevei">
<emphasis id="5DFFEABEFFFAFF92B80A6FF6FEAFFBD3" italics="true" pageId="6" pageNumber="65">C. shrevei</emphasis>
</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFAFF92BC6E6FDEFDBFFBD3" box="[372,587,1004,1029]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="tamacuarensis">
<emphasis id="5DFFEABEFFFAFF92BC6E6FDEFDBFFBD3" box="[372,587,1004,1029]" italics="true" pageId="6" pageNumber="65">C. tamacuarensis</emphasis>
</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFAFF92BF7E6FDEFCFEFBD3" box="[612,778,1004,1029]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="tepuyensis">
<emphasis id="5DFFEABEFFFAFF92BF7E6FDEFCFEFBD3" box="[612,778,1004,1029]" italics="true" pageId="6" pageNumber="65">C. tepuyensis</emphasis>
</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFAFF92BE396FDEFC54FBD3" box="[803,928,1004,1029]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="triunfo">
<emphasis id="5DFFEABEFFFAFF92BE396FDEFC54FBD3" box="[803,928,1004,1029]" italics="true" pageId="6" pageNumber="65">C. triunfo</emphasis>
</taxonomicName>
,
<emphasis id="5DFFEABEFFFAFF92BEA26FDEFB14FBD3" box="[952,1248,1004,1029]" italics="true" pageId="6" pageNumber="65">
<taxonomicName id="A88B4D2FFFFAFF92BEA26FDEFBB7FBD3" box="[952,1091,1004,1029]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="wothuja">C. wothuja</taxonomicName>
and C. sp.
</emphasis>
from Sarisariñama (Barrio­Amorós and Brewer­Carías,
<emphasis id="5DFFEABEFFFAFF92BE7C6826FC31FBFB" box="[870,965,1044,1069]" italics="true" pageId="6" pageNumber="65">in press</emphasis>
), also share other characters, such as a short oblique lateral stripe, absence of dorsolateral and ventrolateral stripes, moderate to scant webbing, and in at least two species,
<taxonomicName id="A88B4D2FFFFAFF92BE896856FBF1FBAB" box="[915,1029,1124,1149]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="triunfo">
<emphasis id="5DFFEABEFFFAFF92BE896856FBF1FBAB" box="[915,1029,1124,1149]" italics="true" pageId="6" pageNumber="65">C. triunfo</emphasis>
</taxonomicName>
and
<taxonomicName id="A88B4D2FFFFAFF92B9246856FB4BFBAB" box="[1086,1215,1124,1149]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="wothuja">
<emphasis id="5DFFEABEFFFAFF92B9246856FB4BFBAB" box="[1086,1215,1124,1149]" italics="true" pageId="6" pageNumber="65">C. wothuja</emphasis>
</taxonomicName>
, absence of palatine bones. I assume that most of the species of the Venezuelan Guayana (and probably the rest of the
<collectingCountry id="179C763CFFFAFF92BF066886FD84FB18" box="[540,624,1204,1230]" name="Guyana" pageId="6" pageNumber="65">Guiana</collectingCountry>
Shield) that do not belong to the
<taxonomicName id="A88B4D2FFFFAFF92BEEE6886FB79FB1B" box="[1012,1165,1204,1229]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="trilineatus">
<emphasis id="5DFFEABEFFFAFF92BEEE6886FB79FB1B" box="[1012,1165,1204,1229]" italics="true" pageId="6" pageNumber="65">C. trilineatus</emphasis>
</taxonomicName>
group (
<emphasis id="5DFFEABEFFFAFF92B9F36886FADEFB1B" box="[1257,1322,1204,1229]" italics="true" pageId="6" pageNumber="65">sensu</emphasis>
Morales 2002), will have a MLP (which can be larger in males) and absence of palatine bones, which suggest they might form a monophyletic group.
</paragraph>
<paragraph id="6F3436ACFFFAFF92BC22691EFE1EFA40" blockId="6.[264,1324,284,1870]" pageId="6" pageNumber="65">
The remaining species are not known to have the MLP, but it is probable to occur in
<emphasis id="5DFFEABEFFFAFF92B80A691EFBD2FABB" italics="true" pageId="6" pageNumber="65">
<taxonomicName id="A88B4D2FFFFAFF92B80A691EFE5EFABB" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="ayarzaguenai">C. ayarzaguenai</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFAFF92BCD86966FD70FABB" box="[450,644,1364,1389]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="guanayensis">C. guanayensis</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFAFF92BF816966FC89FABB" box="[667,893,1364,1389]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="murisipanensis">C. murisipanensis</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFAFF92BE8F6966FBD2FABB" box="[917,1062,1364,1389]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="parimae">C. parimae</taxonomicName>
</emphasis>
, and
<taxonomicName id="A88B4D2FFFFAFF92B9676966FADFFABB" box="[1149,1323,1364,1389]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="sanmartini">
<emphasis id="5DFFEABEFFFAFF92B9676966FADFFABB" box="[1149,1323,1364,1389]" italics="true" pageId="6" pageNumber="65">C. sanmartini</emphasis>
</taxonomicName>
(unpublished data).
</paragraph>
<paragraph id="6F3436ACFFFAFF93BC226996FE57FE88" blockId="6.[264,1324,284,1870]" lastBlockId="7.[264,1323,284,513]" lastPageId="7" lastPageNumber="66" pageId="6" pageNumber="65">
The geography of the MLP is of interest. Grant
<emphasis id="5DFFEABEFFFAFF92BE6A6996FC58FA6B" box="[880,940,1444,1469]" italics="true" pageId="6" pageNumber="65">et al.</emphasis>
(1997) recognized eight species (only four named at that time) of
<taxonomicName id="A88B4D2FFFFAFF92BFC869FEFCA8FA33" box="[722,860,1484,1509]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="genus">
<emphasis id="5DFFEABEFFFAFF92BFC869FEFCA8FA33" box="[722,860,1484,1509]" italics="true" pageId="6" pageNumber="65">Colostethus</emphasis>
</taxonomicName>
having a MLP. I mentioned above Venezuelan species currently known to have a MLP, which are 11 (plus two more without proper name). Of all species with the MLP, only three (
<taxonomicName id="A88B4D2FFFFAFF92BE976A2EFBBCF9E3" box="[909,1096,1564,1589]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="atopoglossus">
<emphasis id="5DFFEABEFFFAFF92BE976A2EFBBCF9E3" box="[909,1096,1564,1589]" italics="true" pageId="6" pageNumber="65">C. atopoglossus</emphasis>
</taxonomicName>
,
<taxonomicName id="A88B4D2FFFFAFF92B94C6A2EFB0DF9E3" box="[1110,1273,1564,1589]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="lacrimosus">
<emphasis id="5DFFEABEFFFAFF92B94C6A2EFB0DF9E3" box="[1110,1273,1564,1589]" italics="true" pageId="6" pageNumber="65">C. lacrimosus</emphasis>
</taxonomicName>
and
<taxonomicName id="A88B4D2FFFFAFF92BC126A76FE28F98B" box="[264,476,1604,1630]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="chocoensis">
<emphasis id="5DFFEABEFFFAFF92BC126A76FED7F98B" box="[264,291,1604,1629]" italics="true" pageId="6" pageNumber="65">C.</emphasis>
aff.
<emphasis id="5DFFEABEFFFAFF92BC416A76FE28F98B" box="[347,476,1604,1629]" italics="true" pageId="6" pageNumber="65">chocoensis</emphasis>
</taxonomicName>
from
<collectingCountry id="179C763CFFFAFF92BF3F6A76FD7DF988" box="[549,649,1604,1630]" name="Ecuador" pageId="6" pageNumber="65">Ecuador</collectingCountry>
) are known from the Chocoan bioregion (Pacific versant of
<collectingCountry id="179C763CFFFAFF92BC366A5EFE78F950" box="[300,396,1644,1670]" name="Panama" pageId="6" pageNumber="65">Panama</collectingCountry>
,
<collectingCountry id="179C763CFFFAFF92BC816A5EFDFAF950" box="[411,526,1644,1670]" name="Colombia" pageId="6" pageNumber="65">Colombia</collectingCountry>
and
<collectingCountry id="179C763CFFFAFF92BF556A5EFD41F950" box="[591,693,1644,1670]" name="Ecuador" pageId="6" pageNumber="65">Ecuador</collectingCountry>
), and the rest (13) occur at the
<collectingCountry id="179C763CFFFAFF92B95A6A5EFB61F950" box="[1088,1173,1644,1670]" name="Guyana" pageId="6" pageNumber="65">Guiana</collectingCountry>
Shield. The distance between the closest Guayanan and Chocoan species, respectively
<taxonomicName id="A88B4D2FFFFAFF92B96E6AA6FB1EF97B" box="[1140,1258,1684,1709]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="shrevei">
<emphasis id="5DFFEABEFFFAFF92B96E6AA6FB1EF97B" box="[1140,1258,1684,1709]" italics="true" pageId="6" pageNumber="65">C. shrevei</emphasis>
</taxonomicName>
from Venezuelan Amazonas state and
<taxonomicName id="A88B4D2FFFFAFF92BF8E6A8EFCCEF903" box="[660,826,1724,1749]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="lacrimosus">
<emphasis id="5DFFEABEFFFAFF92BF8E6A8EFCCEF903" box="[660,826,1724,1749]" italics="true" pageId="6" pageNumber="65">C. lacrimosus</emphasis>
</taxonomicName>
from Valle del Cauca in
<collectingCountry id="179C763CFFFAFF92B96A6A8EFB12F900" box="[1136,1254,1724,1750]" name="Colombia" pageId="6" pageNumber="65">Colombia</collectingCountry>
, is of about
<quantity id="A8739B49FFFAFF92BC4A6AD6FE4EF928" box="[336,442,1764,1790]" metricMagnitude="6" metricUnit="m" metricValue="1.2" pageId="6" pageNumber="65" unit="km" value="1200.0">1200 km</quantity>
, separated by Los Llanos, the Cordillera Oriental and Central de
<collectingCountry id="179C763CFFFAFF92B9AA6AD6FADCF928" box="[1200,1320,1764,1790]" name="Colombia" pageId="6" pageNumber="65">Colombia</collectingCountry>
. No relationship can be inferred between the Chocoan and Guayanan groups of
<taxonomicName id="A88B4D2FFFFAFF92B9BA6B3EFADFF8F3" box="[1184,1323,1804,1829]" class="Amphibia" family="Dendrobatidae" genus="Colostethus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="genus">
<emphasis id="5DFFEABEFFFAFF92B9BA6B3EFADFF8F3" box="[1184,1323,1804,1829]" italics="true" pageId="6" pageNumber="65">Colostethus</emphasis>
</taxonomicName>
having MLP. Grant
<emphasis id="5DFFEABEFFFAFF92BCEE6B06FDDBF89B" box="[500,559,1844,1869]" italics="true" pageId="6" pageNumber="65">et al.</emphasis>
(1997) considered the MLP to be a dendrobatid plesiomorphy. I consider here the presence of the MLP in both biogeographically separated groups, as a convergence.
</paragraph>
<paragraph id="6F3436ACFFFBFF93BC226D5DFC0FFDD7" blockId="7.[264,1323,284,513]" pageId="7" pageNumber="66">
Due to its restricted distribution to a single tepui of
<quantity id="A8739B49FFFBFF93BE826D5DFC0CFE5F" box="[920,1016,367,393]" metricMagnitude="3" metricUnit="m" metricValue="4.37" pageId="7" pageNumber="66" unit="km" value="4.37">4.37 km</quantity>
2, I consider the species to be potentially endangered and sensitive to natural or artificial catastrophes, as was predicted by
<bibRefCitation id="0B1A4B5DFFFBFF93BCBA6D8DFD5CFE0F" author="Barrio-Amoros" box="[416,680,447,473]" pageId="7" pageNumber="66" refString="Barrio-Amoros, C. L. (2001) State of knowledge on the declination of amphibians in Venezuela. FROGLOG, 47, 2 - 4." type="journal article" year="2001">Barrio­Amorós (2001)</bibRefCitation>
for other tepui endemics. I should locate the species in the category VU D2 of the IUCN, following
<bibRefCitation id="0B1A4B5DFFFBFF93BE026DD5FC03FDD7" box="[792,1015,487,513]" pageId="7" pageNumber="67" refString="Young, B. E., Stuart, S. N., Chanson, J. S., Cox, N. A. &amp; Boucher, T. M. (2004) Dissapearing jewels: The status of New World Amphibians. NatureServe, Arlington, Virginia." type="book">Young et al (2004)</bibRefCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>