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<document id="E9261E3249E9ED2DC6D09E9210F18D5C" ID-DOI="10.5281/zenodo.184994" ID-GBIF-Dataset="d0919e71-6d1b-4899-ac73-536f20733624" ID-ISSN="1175-5326" ID-Zenodo-Dep="184994" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460054369551" checkinUser="plazi" docAuthor="Pombal, José P., Siqueira, Carla C., Dorigo, Thiago A., Vrcibradic, Davor &amp; Rocha, Carlos Frederico D." docDate="2008" docId="181287A1A5364518C8D7FC628F12F971" docLanguage="en" docName="zt01938p068.pdf" docOrigin="Zootaxa 1938" docStyle="DocumentStyle:6581673A57F01A3145754A1E615EDFF0.4:Zootaxa.2007-2008.journal_article" docStyleId="6581673A57F01A3145754A1E615EDFF0" docStyleName="Zootaxa.2007-2008.journal_article" docStyleVersion="4" docTitle="Holoaden pholeter Pombal, Siqueira, Dorigo, Vrcibradic &amp; Rocha, 2008, sp. nov." docType="treatment" docVersion="8" lastPageNumber="65" masterDocId="E42BFFD9A537451CC840FFB88B28FFD3" masterDocTitle="A third species of the rare frog genus Holoaden (Terrarana, Strabomantidae) from a montane rainforest area of southeastern Brazil" masterLastPageNumber="68" masterPageNumber="61" pageNumber="62" updateTime="1698234573113" updateUser="plazi">
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<mods:title id="B982DEECBD3E9B6BBD263548FA1E514F">A third species of the rare frog genus Holoaden (Terrarana, Strabomantidae) from a montane rainforest area of southeastern Brazil</mods:title>
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<mods:namePart id="69EE60CEC7291F37CF84862981598F0F">Pombal, José P.</mods:namePart>
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<mods:namePart id="71A3F9ACBE3D3460951138357A3C3A3D">Siqueira, Carla C.</mods:namePart>
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<mods:namePart id="717E2F3998A2ADCA525825392E846CC0">Dorigo, Thiago A.</mods:namePart>
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<mods:namePart id="2199BC1FB75857D7F15DF3930E60304D">Vrcibradic, Davor</mods:namePart>
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<mods:namePart id="3966BF151CDBFAC81159E45B67C004E7">Rocha, Carlos Frederico D.</mods:namePart>
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<mods:title id="6CC7DAF040D436237B73A2A18A99D062">Zootaxa</mods:title>
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<mods:date id="8523B5C8C73C7770BB3132054274DE8F">2008</mods:date>
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<mods:number id="FE2AC7B300BCB5157C7B9393DA36B6E7">1938</mods:number>
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<treatment id="181287A1A5364518C8D7FC628F12F971" ID-DOI="http://doi.org/10.5281/zenodo.6231300" ID-GBIF-Taxon="124874249" ID-Zenodo-Dep="6231300" LSID="urn:lsid:plazi:treatment:181287A1A5364518C8D7FC628F12F971" httpUri="http://treatment.plazi.org/id/181287A1A5364518C8D7FC628F12F971" lastPageId="4" lastPageNumber="65" pageId="1" pageNumber="62">
<subSubSection id="D8A1653CA536451DC8D7FC628A2CFBC4" pageId="1" pageNumber="62" type="nomenclature">
<paragraph id="900436B7A536451DC8D7FC628AF5FC27" blockId="1.[151,477,986,1047]" box="[151,477,986,1012]" pageId="1" pageNumber="62">
<heading id="CB4C81DBA536451DC8D7FC628AF5FC27" bold="true" box="[151,477,986,1012]" fontSize="11" level="1" pageId="1" pageNumber="62" reason="1">
<emphasis id="A2CFEAA5A536451DC8D7FC628AF5FC27" bold="true" box="[151,477,986,1012]" pageId="1" pageNumber="62">
<taxonomicName id="57BB4D34A536451DC8D7FC628A51FC27" box="[151,377,986,1012]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="pholeter" status="sp. nov.">
<emphasis id="A2CFEAA5A536451DC8D7FC628A51FC27" bold="true" box="[151,377,986,1012]" italics="true" pageId="1" pageNumber="62">Holoaden pholeter</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B9FC57DEA536451DC9C0FC628AF5FC27" box="[384,477,986,1012]" pageId="1" pageNumber="62" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="900436B7A536451DC8D7FC458A2CFBC4" blockId="1.[151,477,986,1047]" box="[151,260,1021,1047]" pageId="1" pageNumber="62">
<figureCitation id="08802A32A536451DC8D7FC458A2CFBC4" box="[151,260,1021,1047]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURE 3" captionStartId-0="2.[151,254,1055,1079]" captionStartId-1="2.[151,254,1615,1639]" captionStartId-2="3.[151,255,1971,1995]" captionTargetBox-0="[161,1425,183,1029]" captionTargetBox-1="[387,1206,1154,1570]" captionTargetBox-2="[183,1436,388,1926]" captionTargetId-0="figure@2.[161,1426,183,1029]" captionTargetId-1="figure@2.[378,1209,1135,1588]" captionTargetId-2="figure@3.[138,1450,359,1944]" captionTargetPageId-0="2" captionTargetPageId-1="2" captionTargetPageId-2="3" captionText-0="FIGURE 1. Holoaden pholeter sp. nov., holotype (MNRJ 51475, adult female, SVL 44.6 mm), in life." captionText-1="FIGURE 2. Holoaden pholeter sp. nov., holotype (MNRJ 51475, adult female, SVL 44.6 mm), dorsal and ventral views." captionText-2="FIGURE 3. Holoaden pholeter sp. nov., holotype (MNRJ 51475). Dorsal and lateral views of head; ventral views of hand and foot; scale bars represent 5 mm." httpUri-0="https://zenodo.org/record/184995/files/figure.png" httpUri-1="https://zenodo.org/record/184996/files/figure.png" httpUri-2="https://zenodo.org/record/184997/files/figure.png" pageId="1" pageNumber="62">Figs. 1–3</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="D8A1653CA536451DC8D7FBE88A41FA8E" pageId="1" pageNumber="62" type="materials_examined">
<paragraph id="900436B7A536451DC8D7FBE88EB8FB31" blockId="1.[151,1436,1101,1973]" pageId="1" pageNumber="62">
<typeStatus id="4F008815A536451DC8D7FBE88A21FBB9" box="[151,265,1104,1130]" pageId="1" pageNumber="62" type="holotype">
<emphasis id="A2CFEAA5A536451DC8D7FBE88A21FBB9" bold="true" box="[151,265,1104,1130]" pageId="1" pageNumber="62">Holotype</emphasis>
</typeStatus>
.
<collectionCode id="F6AAAE72A536451DC957FBE88A4DFBB9" box="[279,357,1104,1130]" httpUri="http://grbio.org/cool/zi1i-a0b5" name="Museu Nacional/Universidade Federal de Rio de Janeiro" pageId="1" pageNumber="62">MNRJ</collectionCode>
51475, adult female, collected on a forested mountain slope (
<geoCoordinate id="F58F5070A536451DCC77FBE88FE0FBB9" box="[1079,1224,1101,1130]" direction="south" orientation="latitude" pageId="1" pageNumber="62" precision="15" value="-22.373056">22°22’23” S</geoCoordinate>
;
<geoCoordinate id="F58F5070A536451DCC94FBE88E47FBB9" box="[1236,1391,1101,1130]" direction="west" orientation="longitude" pageId="1" pageNumber="62" precision="15" value="-42.554443">42°33’16” W</geoCoordinate>
; ca.
<quantity id="57439B52A536451DC8D7FBC08BC7FB41" box="[151,239,1144,1170]" metricMagnitude="3" metricUnit="m" metricValue="1.2" pageId="1" pageNumber="62" unit="m" value="1200.0">1200 m</quantity>
above sea level) in the district of Theodoro de Oliveira, municipality of Nova Friburgo, Rio de Janeiro State,
<collectingCountry id="E8AC7627A536451DC89FFB188A02FB69" box="[223,298,1184,1210]" name="Brazil" pageId="1" pageNumber="62">Brazil</collectingCountry>
, on
<date id="E4051077A536451DC919FB188924FB69" box="[345,524,1184,1210]" pageId="1" pageNumber="62" value="2008-03-17">17 March 2008</date>
by Thiago A. Dorigo. The area is located within the Parque Estadual dos Três Picos, a recently created Conservation Unit that encompasses much of the Serra dos Órgãos mountain range.
</paragraph>
<paragraph id="900436B7A536451DC886FB488A41FA8E" blockId="1.[151,1436,1101,1973]" pageId="1" pageNumber="62">
<typeStatus id="4F008815A536451DC886FB488A6BFAD9" box="[198,323,1264,1290]" pageId="1" pageNumber="62" type="paratype">
<emphasis id="A2CFEAA5A536451DC886FB488A6BFAD9" bold="true" box="[198,323,1264,1290]" pageId="1" pageNumber="62">Paratypes</emphasis>
</typeStatus>
.
<collectionCode id="F6AAAE72A536451DC915FB488A8BFAD9" box="[341,419,1264,1290]" httpUri="http://grbio.org/cool/zi1i-a0b5" name="Museu Nacional/Universidade Federal de Rio de Janeiro" pageId="1" pageNumber="62">MNRJ</collectionCode>
53482 (male) and
<collectionCode id="F6AAAE72A536451DCACDFB4889F4FAD9" box="[653,732,1264,1290]" httpUri="http://grbio.org/cool/zi1i-a0b5" name="Museu Nacional/Universidade Federal de Rio de Janeiro" pageId="1" pageNumber="62">MNRJ</collectionCode>
53483 (female), both collected at the same locality of the
<typeStatus id="4F008815A536451DC8D7FAA38A29FAE6" box="[151,257,1307,1333]" pageId="1" pageNumber="62" type="holotype">holotype</typeStatus>
, on a nearby mountain slope (
<geoCoordinate id="F58F5070A536451DCA29FAA389D3FAE6" box="[617,763,1304,1333]" direction="south" orientation="latitude" pageId="1" pageNumber="62" precision="15" value="-22.366388">22°21’59” S</geoCoordinate>
;
<geoCoordinate id="F58F5070A536451DCB4AFAA3888EFAE6" box="[778,934,1304,1333]" direction="west" orientation="longitude" pageId="1" pageNumber="62" precision="15" value="-42.55861">42°33’31” W</geoCoordinate>
; ca.
<quantity id="57439B52A536451DCB9DFAA38F10FAE6" box="[989,1080,1307,1333]" metricMagnitude="3" metricUnit="m" metricValue="1.4" pageId="1" pageNumber="62" unit="m" value="1400.0">1400 m</quantity>
a.s.l.), on
<date id="E4051077A536451DCCF7FAA38E5EFAE6" box="[1207,1398,1307,1333]" pageId="1" pageNumber="62" value="2008-08-14">14 August 2008</date>
by Carla C. Siqueira.
</paragraph>
</subSubSection>
<subSubSection id="D8A1653CA536451DC886FAD28EBDF866" pageId="1" pageNumber="62" type="diagnosis">
<paragraph id="900436B7A536451DC886FAD288CBF9F6" blockId="1.[151,1436,1101,1973]" pageId="1" pageNumber="62">
<emphasis id="A2CFEAA5A536451DC886FAD28A17FA57" bold="true" box="[198,319,1386,1412]" pageId="1" pageNumber="62">Diagnosis</emphasis>
. Dorsum densely glandular, toes free of webbing, fingers and toes with rounded tips not expanded into disks, large and relatively wide head, absence of a tympanum, big forward-directed eyes, and rhomboid or round (when dilated) pupils.
<taxonomicName id="57BB4D34A536451DCA3FFA038873FA07" box="[639,859,1467,1492]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="pholeter">
<emphasis id="A2CFEAA5A536451DCA3FFA038873FA07" box="[639,859,1467,1492]" italics="true" pageId="1" pageNumber="62">Holoaden pholeter</emphasis>
</taxonomicName>
is a comparatively large
<taxonomicName id="57BB4D34A536451DCC3EFA038FDAFA07" box="[1150,1266,1467,1492]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="genus">
<emphasis id="A2CFEAA5A536451DCC3EFA038FDAFA07" box="[1150,1266,1467,1492]" italics="true" pageId="1" pageNumber="62">Holoaden</emphasis>
</taxonomicName>
species with a large head, glandular texture on dorsum, limbs long and slender, overall dorsal coloration deeply dark, and ventral body surface dark with a light abdominal blotch of variable size.
</paragraph>
<paragraph id="900436B7A536451DC886F98A8EBDF866" blockId="1.[151,1436,1101,1973]" pageId="1" pageNumber="62">
<emphasis id="A2CFEAA5A536451DC886F98A89A0F99F" bold="true" box="[198,648,1586,1612]" pageId="1" pageNumber="62">Comparisons with the other species</emphasis>
. The new species differs from
<taxonomicName id="57BB4D34A536451DCBBDF98B8FEEF99F" box="[1021,1222,1587,1612]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="bradei">
<emphasis id="A2CFEAA5A536451DCBBDF98B8FEEF99F" box="[1021,1222,1587,1612]" italics="true" pageId="1" pageNumber="62">Holoaden bradei</emphasis>
</taxonomicName>
in its larger body size (maximum
<quantity id="57439B52A536451DC914F9E38A8FF9A6" box="[340,423,1627,1653]" metricMagnitude="-2" metricUnit="m" metricValue="3.7" pageId="1" pageNumber="62" unit="mm" value="37.0">37 mm</quantity>
SVL in
<taxonomicName id="57BB4D34A536451DCA4BF9E38953F9A7" box="[523,635,1627,1652]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="bradei">
<emphasis id="A2CFEAA5A536451DCA4BF9E38953F9A7" box="[523,635,1627,1652]" italics="true" pageId="1" pageNumber="62">H. bradei</emphasis>
</taxonomicName>
;
<bibRefCitation id="F42A4B46A536451DCACBF9E3882FF9A6" author="Lutz" box="[651,775,1627,1653]" pageId="1" pageNumber="62" refString="Lutz, B. (1958) Anfibios novos e raros das Serras Costeiras do Brasil (New or rare frogs from the Coastal Ranges of Brazil). Eleutherodactylus venancioi n. sp., E. hoehnei n. sp., Holoaden bradei n. sp. e H. luderwaldti Mir. Rib., 1920. Memorias do Instituto Oswaldo Cruz, 56, 373 - 399." type="journal article" year="1958">Lutz 1958</bibRefCitation>
), longer limbs, wider head, palmar and plantar tubercles and tips of digits leaden gray in life (yellow in
<taxonomicName id="57BB4D34A536451DCA8FF93B8868F94F" box="[719,832,1667,1692]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="bradei">
<emphasis id="A2CFEAA5A536451DCA8FF93B8868F94F" box="[719,832,1667,1692]" italics="true" pageId="1" pageNumber="62">H. bradei</emphasis>
</taxonomicName>
;
<bibRefCitation id="F42A4B46A536451DCB12F93B88E7F94E" author="Lutz" box="[850,975,1667,1693]" pageId="1" pageNumber="62" refString="Lutz, B. (1958) Anfibios novos e raros das Serras Costeiras do Brasil (New or rare frogs from the Coastal Ranges of Brazil). Eleutherodactylus venancioi n. sp., E. hoehnei n. sp., Holoaden bradei n. sp. e H. luderwaldti Mir. Rib., 1920. Memorias do Instituto Oswaldo Cruz, 56, 373 - 399." type="journal article" year="1958">Lutz 1958</bibRefCitation>
), and dorsal coloration uniformly dark purplish-brown (upper surfaces olive brown with irregular soot-black suffusions in
<taxonomicName id="57BB4D34A536451DCC28F9138FF0F917" box="[1128,1240,1707,1732]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="bradei">
<emphasis id="A2CFEAA5A536451DCC28F9138FF0F917" box="[1128,1240,1707,1732]" italics="true" pageId="1" pageNumber="62">H. bradei</emphasis>
</taxonomicName>
;
<bibRefCitation id="F42A4B46A536451DCCA8F9138E43F916" author="Lutz" box="[1256,1387,1707,1733]" pageId="1" pageNumber="62" refString="Lutz, B. (1958) Anfibios novos e raros das Serras Costeiras do Brasil (New or rare frogs from the Coastal Ranges of Brazil). Eleutherodactylus venancioi n. sp., E. hoehnei n. sp., Holoaden bradei n. sp. e H. luderwaldti Mir. Rib., 1920. Memorias do Instituto Oswaldo Cruz, 56, 373 - 399." type="journal article" year="1958">Lutz, 1958</bibRefCitation>
; see color picture in
<bibRefCitation id="F42A4B46A536451DC912F96B891DF93E" author="Haddad" box="[338,565,1747,1773]" pageId="1" pageNumber="62" refString="Haddad, C. F. B., Toledo, L. F., &amp; Prado, C. P. A. (2008) Anfibios da Mata Atlantica - Guia dos anfibios anuros da Mata Atlantica. Editora Neotropica, Sao Paulo, 243 pp." type="book" year="2008">
Haddad
<emphasis id="A2CFEAA5A536451DC9F5F96B8AC1F93F" box="[437,489,1747,1772]" italics="true" pageId="1" pageNumber="62">et al</emphasis>
. 2008
</bibRefCitation>
, page 80).
<taxonomicName id="57BB4D34A536451DCAF7F96B88BCF93F" box="[695,916,1747,1772]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="pholeter">
<emphasis id="A2CFEAA5A536451DCAF7F96B88BCF93F" box="[695,916,1747,1772]" italics="true" pageId="1" pageNumber="62">Holoaden pholeter</emphasis>
</taxonomicName>
is morphologically more similar to
<taxonomicName id="57BB4D34A536451DCD7CF96B8BD1F8C7" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="luederwaldti">
<emphasis id="A2CFEAA5A536451DCD7CF96B8BD1F8C7" italics="true" pageId="1" pageNumber="62">H. luederwaldti</emphasis>
</taxonomicName>
, but is distinguished from the latter by its moderately bulging dorsal glands (dorsal glands markedly bulging and more conspicuous in
<taxonomicName id="57BB4D34A536451DCA6AF89B89CBF8EF" box="[554,739,1827,1852]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="luederwaldti">
<emphasis id="A2CFEAA5A536451DCA6AF89B89CBF8EF" box="[554,739,1827,1852]" italics="true" pageId="1" pageNumber="62">H. luederwaldti</emphasis>
</taxonomicName>
; see figures
<quantity id="57439B52A536451DCB3CF89B88EAF8EE" box="[892,962,1827,1853]" metricMagnitude="-2" metricUnit="m" metricValue="3.81" metricValueMax="5.08" metricValueMin="2.54" pageId="1" pageNumber="62" unit="in" value="1.5" valueMax="2.0" valueMin="1.0">1-2 in</quantity>
Caramaschi &amp;
<bibRefCitation id="F42A4B46A536451DCC3DF89B8E33F8EE" author="Pombal" box="[1149,1307,1827,1853]" pageId="1" pageNumber="62" refString="Caramashi, U. &amp; Pombal Jr, J. P. (2006) Notas sobre as series-tipo de Holoaden bradei B. Lutz e Holoaden luederwaldti Miranda-Ribeiro (Anura, Brachycephalidae). Revista Brasileira de Zoologia, 23, 1261 - 1263." type="journal article" year="2006">Pombal 2006</bibRefCitation>
and figure
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<bibRefCitation id="F42A4B46A536451DC8A0F8F38AE2F8B6" author="Hedges" box="[224,458,1867,1893]" pageId="1" pageNumber="62" refString="Hedges, S. B., Duellman, W. E. &amp; Heinicke, M. P. (2008) New World direct-developing frogs (Anura: Terrana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182." type="journal article" year="2008">
Hedges
<emphasis id="A2CFEAA5A536451DC903F8F38A52F8B7" box="[323,378,1867,1892]" italics="true" pageId="1" pageNumber="62">et al</emphasis>
. 2008
</bibRefCitation>
), darker dorsal coloration (dorsal surfaces brown to reddish brown in
<taxonomicName id="57BB4D34A536451DCD60F8F38BC8F85F" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="luederwaldti">
<emphasis id="A2CFEAA5A536451DCD60F8F38BC8F85F" italics="true" pageId="1" pageNumber="62">H. luederwaldti</emphasis>
</taxonomicName>
; see figure
<quantity id="57439B52A536451DC930F8CB8A86F85E" box="[368,430,1907,1933]" metricMagnitude="0" metricUnit="m" metricValue="2.4892" pageId="1" pageNumber="62" unit="in" value="98.0">98 in</quantity>
<bibRefCitation id="F42A4B46A536451DC9F7F8CB89B5F85E" author="Hedges" box="[439,669,1907,1933]" pageId="1" pageNumber="62" refString="Hedges, S. B., Duellman, W. E. &amp; Heinicke, M. P. (2008) New World direct-developing frogs (Anura: Terrana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182." type="journal article" year="2008">
Hedges
<emphasis id="A2CFEAA5A536451DCA58F8CB8966F85F" box="[536,590,1907,1932]" italics="true" pageId="1" pageNumber="62">et al</emphasis>
. 2008
</bibRefCitation>
), and ventral surfaces dark (slightly lighter than dorsum) with a light blotch on abdomen (ventral surfaces uniformly cream, distinctly lighter than dorsum in
<taxonomicName id="57BB4D34A536451DCC8DF8238EACF867" box="[1229,1412,1947,1972]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="1" pageNumber="62" phylum="Chordata" rank="species" species="luederwaldti">
<emphasis id="A2CFEAA5A536451DCC8DF8238EACF867" box="[1229,1412,1947,1972]" italics="true" pageId="1" pageNumber="62">H. luederwaldti</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<caption id="C4C4663FA535451EC8D7FBA78FCAFBE4" box="[151,1250,1055,1079]" httpUri="https://zenodo.org/record/184995/files/figure.png" pageId="2" pageNumber="63" targetBox="[161,1425,183,1029]" targetPageId="2">
<paragraph id="900436B7A535451EC8D7FBA78FCAFBE4" blockId="2.[151,1250,1055,1079]" box="[151,1250,1055,1079]" pageId="2" pageNumber="63">
<emphasis id="A2CFEAA5A535451EC8D7FBA78A30FBE4" bold="true" box="[151,280,1055,1079]" pageId="2" pageNumber="63">FIGURE 1.</emphasis>
<taxonomicName id="57BB4D34A535451EC95FFB988AC0FBE4" box="[287,488,1056,1079]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="2" pageNumber="63" phylum="Chordata" rank="species" species="pholeter" status="sp. nov.">
<emphasis id="A2CFEAA5A535451EC95FFB988AC0FBE4" box="[287,488,1056,1079]" italics="true" pageId="2" pageNumber="63">Holoaden pholeter</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B9FC57DEA535451EC9AEFBA7896CFBE4" box="[494,580,1055,1079]" pageId="2" pageNumber="63" rank="species">
<emphasis id="A2CFEAA5A535451EC9AEFBA7896CFBE4" bold="true" box="[494,580,1055,1079]" pageId="2" pageNumber="63">sp. nov.</emphasis>
</taxonomicNameLabel>
, holotype (MNRJ 51475, adult female, SVL 44.6 mm), in life.
</paragraph>
</caption>
<caption id="C4C4663FA535451EC8D7F9F78EBFF9B5" box="[151,1431,1615,1639]" httpUri="https://zenodo.org/record/184996/files/figure.png" pageId="2" pageNumber="63" targetBox="[387,1206,1154,1570]" targetPageId="2">
<paragraph id="900436B7A535451EC8D7F9F78EBFF9B5" blockId="2.[151,1431,1615,1639]" box="[151,1431,1615,1639]" pageId="2" pageNumber="63">
<emphasis id="A2CFEAA5A535451EC8D7F9F78A39F9B4" bold="true" box="[151,273,1615,1639]" pageId="2" pageNumber="63">FIGURE 2</emphasis>
.
<taxonomicName id="57BB4D34A535451EC95DF9E88ACDF9B4" box="[285,485,1616,1639]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="2" pageNumber="63" phylum="Chordata" rank="species" species="pholeter" status="sp. nov.">
<emphasis id="A2CFEAA5A535451EC95DF9E88ACDF9B4" box="[285,485,1616,1639]" italics="true" pageId="2" pageNumber="63">Holoaden pholeter</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B9FC57DEA535451EC9ABF9F78917F9B4" box="[491,575,1615,1639]" pageId="2" pageNumber="63" rank="species">
<emphasis id="A2CFEAA5A535451EC9ABF9F78917F9B4" bold="true" box="[491,575,1615,1639]" pageId="2" pageNumber="63">sp. nov.</emphasis>
</taxonomicNameLabel>
, holotype (MNRJ 51475, adult female, SVL 44.6 mm), dorsal and ventral views.
</paragraph>
</caption>
<subSubSection id="D8A1653CA5354518C886F92C8808FD69" lastPageId="4" lastPageNumber="65" pageId="2" pageNumber="63" type="description">
<paragraph id="900436B7A535451FC886F92C8AB7FE81" blockId="2.[151,1437,1684,2031]" lastBlockId="3.[151,1436,152,338]" lastPageId="3" lastPageNumber="64" pageId="2" pageNumber="63">
<emphasis id="A2CFEAA5A535451EC886F92C8AC7F97D" bold="true" box="[198,495,1684,1710]" pageId="2" pageNumber="63">
Description of
<typeStatus id="4F008815A535451EC9C3F92C8AC7F97D" box="[387,495,1684,1710]" pageId="2" pageNumber="63" type="holotype">holotype</typeStatus>
</emphasis>
. Body robust; head large, wider than long, and wider than body; snout nearly semicircular in dorsal view and slightly protruding in lateral view; eyes large, protuberant, directed slightly forward; pupil round; nostrils protuberant, directed dorsolaterally; canthus rostralis not well marked and slightly concave; loreal region concave; tympanum absent; a glandular supratympanic fold extending from corner of eye to insertion of front limb; tongue large; vomerine teeth in two short series behind and between choanae; choanae circular; a single small toothlike process in front of lower jaw with a small single socket between premaxillae; numerous teeth on maxilla and premaxillae. Arms slender; forearms and fingers long and slender; finger lengths II&lt;IV&lt;I&lt;III; finger tips rounded, without expanded discs; fingers not webbed nor fringed; subarticular tubercles single, slightly elliptical or rounded; inner metatarsal tubercle large, ovoid; outer metatarsal tubercle large, slightly rounded; few supranumerary tubercles. Legs short, slender; toes long, slender; toe lengths I~II&lt;III~V&lt;IV; toes tips rounded, without expanded discs; toes not webbed or fringed; subarticular tubercles slightly elliptical; inner metatarsal tubercle large, ovoid; outer metatarsal tubercle medium-sized, slightly rounded. Head and dorsum glandular; loreal region, fore limbs and hind limbs smooth; undersurfaces smooth.
</paragraph>
<caption id="C4C4663FA534451FC8D7F80B897CF83F" httpUri="https://zenodo.org/record/184997/files/figure.png" pageId="3" pageNumber="64" targetBox="[183,1436,388,1926]" targetPageId="3">
<paragraph id="900436B7A534451FC8D7F80B897CF83F" blockId="3.[151,1436,1971,2028]" pageId="3" pageNumber="64">
<emphasis id="A2CFEAA5A534451FC8D7F80B8A33F818" bold="true" box="[151,283,1971,1995]" pageId="3" pageNumber="64">FIGURE 3.</emphasis>
<taxonomicName id="57BB4D34A534451FC964F80B8AC7F819" box="[292,495,1971,1994]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="3" pageNumber="64" phylum="Chordata" rank="species" species="pholeter" status="sp. nov.">
<emphasis id="A2CFEAA5A534451FC964F80B8AC7F819" box="[292,495,1971,1994]" italics="true" pageId="3" pageNumber="64">Holoaden pholeter</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B9FC57DEA534451FC9B8F80B8967F818" box="[504,591,1971,1995]" pageId="3" pageNumber="64" rank="species">
<emphasis id="A2CFEAA5A534451FC9B8F80B8967F818" bold="true" box="[504,591,1971,1995]" pageId="3" pageNumber="64">sp. nov.</emphasis>
</taxonomicNameLabel>
, holotype (MNRJ 51475). Dorsal and lateral views of head; ventral views of hand and foot; scale bars represent 5 mm.
</paragraph>
</caption>
<paragraph id="900436B7A5334518C886FF2F8940FE81" blockId="4.[151,1437,151,1698]" pageId="4" pageNumber="65">
<emphasis id="A2CFEAA5A5334518C886FF2F8A8AFF62" bold="true" box="[198,418,151,177]" pageId="4" pageNumber="65">
Color of
<typeStatus id="4F008815A5334518C977FF2F8A8AFF62" box="[311,418,151,177]" pageId="4" pageNumber="65" type="holotype">holotype</typeStatus>
</emphasis>
. In life, upper surfaces and flanks dark purplish-brown; eyes slightly lighter than dorsum (
<figureCitation id="08802A32A5334518C899FF788A37FF09" box="[217,287,192,218]" captionStart="FIGURE 1" captionStartId="2.[151,254,1055,1079]" captionTargetBox="[161,1425,183,1029]" captionTargetId="figure@2.[161,1426,183,1029]" captionTargetPageId="2" captionText="FIGURE 1. Holoaden pholeter sp. nov., holotype (MNRJ 51475, adult female, SVL 44.6 mm), in life." httpUri="https://zenodo.org/record/184995/files/figure.png" pageId="4" pageNumber="65">Fig. 1</figureCitation>
); throat and ventral surfaces slightly lighter than dorsum, with a whitish patch on abdomen; palmar and plantar tubercles and tips of digits leaden gray; in close-up view, dorsal surfaces and eyes have a pattern of numerous very small light dots on a dark brown background. In preservative, upper surfaces and flanks dark purplish-gray, almost black.
</paragraph>
<paragraph id="900436B7A5334518C886FEE78862FE19" blockId="4.[151,1437,151,1698]" pageId="4" pageNumber="65">
<emphasis id="A2CFEAA5A5334518C886FEE78927FEAA" bold="true" box="[198,527,351,377]" pageId="4" pageNumber="65">
Measurements of
<typeStatus id="4F008815A5334518C9E4FEE78927FEAA" box="[420,527,351,377]" pageId="4" pageNumber="65" type="holotype">holotype</typeStatus>
</emphasis>
. Snout-vent length 44.6; head length 15.2; head width 16.3; eye diameter 5.4; interorbital distance 5.9; eye-nostril distance 4.6; internasal distance 3.6; forearm length 12.9; hand length 11.2; femur length 19.9; tibia length 19.0; foot length 17.7.
</paragraph>
<paragraph id="900436B7A5334518C886FE6F8808FD69" blockId="4.[151,1437,151,1698]" pageId="4" pageNumber="65">
<emphasis id="A2CFEAA5A5334518C886FE6F8A11FE22" bold="true" box="[198,313,471,497]" pageId="4" pageNumber="65">Variation</emphasis>
. Measurements for the male and female
<typeStatus id="4F008815A5334518CB74FE60888CFE21" box="[820,932,472,498]" pageId="4" pageNumber="65" type="paratype">paratypes</typeStatus>
are, respectively: snout-vent length 41.6, 47.7; head length 14.7, 18.4; head width 16.4, 19.3; eye diameter 4.7, 6.6; interorbital distance 4.2, 6.2; eyenostril distance 3.7, 4.4; internasal distance 3.1, 3.4; forearm length 11.9, 12.8; hand length 11.0, 12.0; femur length 17.7, 20.6; tibia length 18.0, 19.3; foot length 17.7, 19.5. The forearm in the male
<typeStatus id="4F008815A5334518CCFBFDE88E08FDB9" box="[1211,1312,592,618]" pageId="4" pageNumber="65" type="paratype">paratype</typeStatus>
is slightly more robust and longer than in the two females. The light abdominal blotch is variable in size, being smallest in the male
<typeStatus id="4F008815A5334518C960FD188AADFD69" box="[288,389,672,698]" pageId="4" pageNumber="65" type="paratype">paratype</typeStatus>
and largest in the female
<typeStatus id="4F008815A5334518CAF3FD188834FD69" box="[691,796,672,698]" pageId="4" pageNumber="65" type="paratype">paratype</typeStatus>
.
</paragraph>
</subSubSection>
<subSubSection id="D8A1653CA5334518C886FD7F8F13FCD9" pageId="4" pageNumber="65" type="etymology">
<paragraph id="900436B7A5334518C886FD7F8F13FCD9" blockId="4.[151,1437,151,1698]" pageId="4" pageNumber="65">
<emphasis id="A2CFEAA5A5334518C886FD7F8A63FD32" bold="true" box="[198,331,711,737]" pageId="4" pageNumber="65">Etymology</emphasis>
. The specific epithet “
<taxonomicName id="57BB4D34A5334518CA11FD70899FFD31" box="[593,695,712,738]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="4" pageNumber="65" phylum="Chordata" rank="species" species="pholeter">pholeter</taxonomicName>
” is a Greek word, herein used in apposition, which means “one who lurks in a hole”, in allusion to the cryptic habits of the species (see below).
</paragraph>
</subSubSection>
<subSubSection id="D8A1653CA5334518C886FCAF8F12F971" pageId="4" pageNumber="65" type="distribution">
<paragraph id="900436B7A5334518C886FCAF8AB7FC51" blockId="4.[151,1437,151,1698]" pageId="4" pageNumber="65">
<emphasis id="A2CFEAA5A5334518C886FCAF8A77FCE2" bold="true" box="[198,351,791,817]" pageId="4" pageNumber="65">Distribution</emphasis>
.
<taxonomicName id="57BB4D34A5334518C92EFCA08963FCE2" box="[366,587,792,817]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="4" pageNumber="65" phylum="Chordata" rank="species" species="pholeter">
<emphasis id="A2CFEAA5A5334518C92EFCA08963FCE2" box="[366,587,792,817]" italics="true" pageId="4" pageNumber="65">Holoaden pholeter</emphasis>
</taxonomicName>
is known only from the
<typeStatus id="4F008815A5334518CB2DFCA08888FCE1" box="[877,928,792,818]" pageId="4" pageNumber="65">type</typeStatus>
locality, at
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a.s.l., on the eastern portion of the Serra dos Órgãos (a subdivision of the Serra do Mar mountain range), within the Atlantic Forest domain (fig. 4).
</paragraph>
<paragraph id="900436B7A5334518C886FC378903FA59" blockId="4.[151,1437,151,1698]" pageId="4" pageNumber="65">
<emphasis id="A2CFEAA5A5334518C886FC378AA0FC7A" bold="true" box="[198,392,911,937]" pageId="4" pageNumber="65">Natural history</emphasis>
. All three specimens of
<taxonomicName id="57BB4D34A5334518CAF6FC2888BFFC7A" box="[694,919,912,937]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="4" pageNumber="65" phylum="Chordata" rank="species" species="pholeter">
<emphasis id="A2CFEAA5A5334518CAF6FC2888BFFC7A" box="[694,919,912,937]" italics="true" pageId="4" pageNumber="65">Holoaden pholeter</emphasis>
</taxonomicName>
were collected at night. The
<typeStatus id="4F008815A5334518CCBDFC288E4BFC79" box="[1277,1379,912,938]" pageId="4" pageNumber="65" type="holotype">holotype</typeStatus>
was found when it emerged from the leaf litter that was accumulated among tree roots after the site was disturbed by one of us (TAD).
<typeStatus id="4F008815A5334518C9CDFC588ADBFC29" box="[397,499,992,1018]" pageId="4" pageNumber="65" type="paratype">Paratype</typeStatus>
MNRJ 53483 was collected on the leaf litter, whereas MNRJ 53482 was found shortly afterwards, hidden inside a hole at the base of a tree, ca.
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from where the female was found. The hole where it was found (ca.
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high,
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deep, and
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wide) was adjacent to a second hole, quite similar in dimensions and structure, which may have been used by the female. The floor of these holes consisted of bare soil with a shallow depression (on which the male was nested). Although there was abundant leaf litter on the forest floor surrounding this double den, its interior was clean of litter, as was the small area within
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of its entrance. Judging by these observations, it is possible that this shelter was “cleaned up” by its occupant(s). We cannot assure that
<taxonomicName id="57BB4D34A5334518CA2DFB408866FAC2" box="[621,846,1272,1297]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="4" pageNumber="65" phylum="Chordata" rank="species" species="pholeter">
<emphasis id="A2CFEAA5A5334518CA2DFB408866FAC2" box="[621,846,1272,1297]" italics="true" pageId="4" pageNumber="65">Holoaden pholeter</emphasis>
</taxonomicName>
(and maybe other congeners) actually “prepare” such dens, but if they do, this would represent a unique behavior among terraranan frogs. Nevertheless, our observations, along with those of
<bibRefCitation id="F42A4B46A5334518CA6CFAF08994FAB1" author="Lutz" box="[556,700,1352,1378]" pageId="4" pageNumber="65" refString="Lutz, B. (1958) Anfibios novos e raros das Serras Costeiras do Brasil (New or rare frogs from the Coastal Ranges of Brazil). Eleutherodactylus venancioi n. sp., E. hoehnei n. sp., Holoaden bradei n. sp. e H. luderwaldti Mir. Rib., 1920. Memorias do Instituto Oswaldo Cruz, 56, 373 - 399." type="journal article" year="1958">Lutz (1958)</bibRefCitation>
, suggest that
<taxonomicName id="57BB4D34A5334518CB21FAF088FDFAB2" box="[865,981,1352,1377]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="4" pageNumber="65" phylum="Chordata" rank="genus">
<emphasis id="A2CFEAA5A5334518CB21FAF088FDFAB2" box="[865,981,1352,1377]" italics="true" pageId="4" pageNumber="65">Holoaden</emphasis>
</taxonomicName>
spp. are reclusive animals, frequently hiding in various
<typeStatus id="4F008815A5334518C925FAC88A8BFA59" box="[357,419,1392,1418]" pageId="4" pageNumber="65">types</typeStatus>
of shelters.
</paragraph>
<paragraph id="900436B7A5334518C885FA20899CF9F9" blockId="4.[151,1437,151,1698]" pageId="4" pageNumber="65">
Upon capture, MNRJ 53483 released a copious amount of a colorless and very sticky secretion. It is likely that the numerous, conspicuous dorsal glands of
<taxonomicName id="57BB4D34A5334518CA93FA78886EFA0A" box="[723,838,1472,1497]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="4" pageNumber="65" phylum="Chordata" rank="genus">
<emphasis id="A2CFEAA5A5334518CA93FA78886EFA0A" box="[723,838,1472,1497]" italics="true" pageId="4" pageNumber="65">Holoaden</emphasis>
</taxonomicName>
are responsible for secreting that substance, possibly as a defense, though there is currently no information on other species of the genus regarding such behavior nor the chemical nature of such secretions.
</paragraph>
<paragraph id="900436B7A5334518C885F9808F12F971" blockId="4.[151,1437,151,1698]" pageId="4" pageNumber="65">
The
<typeStatus id="4F008815A5334518C8BBF9808A49F981" box="[251,353,1592,1618]" pageId="4" pageNumber="65" type="holotype">holotype</typeStatus>
and
<typeStatus id="4F008815A5334518C9DBF9808928F981" box="[411,512,1592,1618]" pageId="4" pageNumber="65" type="paratype">paratype</typeStatus>
MNRJ 53483 contained large unpigmented eggs, as is believed to occur in all Terrarana, a clade of direct-developing frogs (
<bibRefCitation id="F42A4B46A5334518CAEBF9D888A4F9A9" author="Hedges" box="[683,908,1632,1658]" pageId="4" pageNumber="65" refString="Hedges, S. B., Duellman, W. E. &amp; Heinicke, M. P. (2008) New World direct-developing frogs (Anura: Terrana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182." type="journal article" year="2008">
Hedges
<emphasis id="A2CFEAA5A5334518CB4CF9D88817F9AA" box="[780,831,1632,1657]" italics="true" pageId="4" pageNumber="65">et al</emphasis>
. 2008
</bibRefCitation>
). Similar eggs were reported for
<taxonomicName id="57BB4D34A5334518CD48F9D88E50F9AA" box="[1288,1400,1632,1657]" class="Amphibia" family="Strabomantidae" genus="Holoaden" kingdom="Animalia" order="Anura" pageId="4" pageNumber="65" phylum="Chordata" rank="species" species="bradei">
<emphasis id="A2CFEAA5A5334518CD48F9D88E50F9AA" box="[1288,1400,1632,1657]" italics="true" pageId="4" pageNumber="65">H. bradei</emphasis>
</taxonomicName>
by
<bibRefCitation id="F42A4B46A5334518C8D7F9308A0AF971" author="Lutz" box="[151,290,1672,1698]" pageId="4" pageNumber="65" refString="Lutz, B. (1958) Anfibios novos e raros das Serras Costeiras do Brasil (New or rare frogs from the Coastal Ranges of Brazil). Eleutherodactylus venancioi n. sp., E. hoehnei n. sp., Holoaden bradei n. sp. e H. luderwaldti Mir. Rib., 1920. Memorias do Instituto Oswaldo Cruz, 56, 373 - 399." type="journal article" year="1958">Lutz (1958)</bibRefCitation>
who observed parental care and direct development in that species.
</paragraph>
</subSubSection>
</treatment>
</document>
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