treatments-xml/data/C9/AC/51/C9AC519FE0737BF5FAA4CEFB27662860.xml
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<mods:title id="698A7941DF6A486C8A3EDD888217DCD8">Systematics of the Rhinellamargaritifera complex (Anura, Bufonidae) from western Ecuador and Panama with insights in the biogeography of Rhinellaalata</mods:title>
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<mods:namePart id="FDEB62371428199E3581FE40722A4726">dos Santos, Sueny P.</mods:namePart>
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<mods:namePart id="C70270CC22C1FA00B24B88B808F65FAB">Ibanez, Roberto</mods:namePart>
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<mods:namePart id="1E41B4D5B89FA418C117DFB255E62E4F">Ron, Santiago R.</mods:namePart>
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<treatment id="C9AC519FE0737BF5FAA4CEFB27662860" ID-GBIF-Taxon="152059885" LSID="urn:lsid:plazi:treatment:C9AC519FE0737BF5FAA4CEFB27662860" httpUri="http://treatment.plazi.org/id/C9AC519FE0737BF5FAA4CEFB27662860" lastPageId="15" lastPageNumber="124" pageId="10" pageNumber="119">
<subSubSection id="D5648F7972EB0196A632CDE90AF116E6" pageId="10" pageNumber="119" type="multiple">
<paragraph id="4AAC230358EE0F67238D2DDEDC172994" pageId="10" pageNumber="119">Taxon classification Animalia Anura Bufonidae</paragraph>
</subSubSection>
<subSubSection id="5215DCBB88EE41C69F69E68F4D2EDB8B" pageId="10" pageNumber="119" type="nomenclature">
<paragraph id="2B68F6D6E42BEF1267D3E699C314692D" pageId="10" pageNumber="119">
<taxonomicName id="8B3314225315C283226B4FB94EC4BC16" ID-CoL="4S7M7" authority="Thominot, 1884" authorityName="Thominot" authorityYear="1884" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="10" pageNumber="119" phylum="Chordata" rank="species" species="alata">Rhinella alata (Thominot, 1884)</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="795235102D0BCED1D8C45F8855FDC007" pageId="10" pageNumber="119" type="reference_group">
<paragraph id="EE48FF5C7AB9357277CC28E5B792DC86" pageId="10" pageNumber="119">
<taxonomicName id="50F78E03688CBE62CD0BCEECAA8F2FB8" class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo alatus" order="Anura" pageId="10" pageNumber="119" phylum="Chordata" rank="species" species="alatus">Bufo alatus</taxonomicName>
Thominot, 1884. Holotype: MNHN 84285, adult male from Obispo, Panama.
</paragraph>
</subSubSection>
<subSubSection id="1B5D08D638A7C18617286180397B972E" lastPageId="11" lastPageNumber="120" pageId="10" pageNumber="119" type="diagnosis">
<paragraph id="617FB040B98891595332A769EC1F2F32" pageId="10" pageNumber="119">Diagnosis.</paragraph>
<paragraph id="1D9256E0C1FD8F875AF03AA7B5256742" pageId="10" pageNumber="119">
<taxonomicName id="7F59CC1A462FAEA882DB19418F0E243D" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="10" pageNumber="119" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
is a small-sized (Table 2; Figs 8 and 9) species of
<taxonomicName id="93736A34A6FA7ADA293C3BB06C021028" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella" order="Anura" pageId="10" pageNumber="119" phylum="Chordata" rank="genus">Rhinella</taxonomicName>
having the following combination of characters: (1) average SVL of females 44.25 mm (SD = 4.36, n = 39), males 36.83 mm (SD = 2.31, n = 49); (2) bony knob at angle of jaws absent, corner of mouth angular; (3) supraorbital crests low and thick, continuous with preorbital crests; usually with crenulate texture on vertical surfaces; (4) supratympanic crests concave and small; their posterior edge usually next to the anterior border of parotoid glands; (5) canthus rostralis present but inconspicuous, sometimes continuous with preorbital crests; (6) parietal crests usually present, ill-defined; (7) heel reaching posterior margin of eye when hindlimbs adpressed; (8) vertebral apophyses no protruding; (9) snout subacuminate in dorsal view, from rounded to protruding in profile; (10) skin on dorsum bearing a mixture of warts, pustules, and minute tubercles; (11) mid-dorsal line from snout to vent often present; (12) spiculate tubercles on external border of shank, evident especially on females; (13) dorsolateral row of sharply pointed, conical tubercles between posterior border of parotoid glands and groin; (14) tympanic membrane and tympanic annulus distinct; moderately large, ovoid to round; (15) parotoid glands small, elongated posteriorly; (16) upper eyelid warty; (17) tarsal fold absent; (18) digits slender and long, with small knobs at tip; lateral fringes present; finger lengths 3&gt; 4&gt; 2&gt; 1; toe lengths 4&gt; 5&gt; 3&gt; 2&gt; 1; (19) nuptial pads present.
</paragraph>
<paragraph id="08B707A8B9366E4F2FFEBF11572D91A6" pageId="11" pageNumber="120">
<taxonomicName id="9535C704A8B42ABF9F77D0288FB3029B" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">
<pageBreakToken id="FEC8C61C4241937A81E0BE467E9CED2A" pageId="11" pageNumber="120" start="start">Rhinella</pageBreakToken>
alata
</taxonomicName>
is most similar to
<taxonomicName id="817E290981BF9CDC1B2E820BB02E9341" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella acutirostris" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="acutirostris">Rhinella acutirostris</taxonomicName>
. Both species differ from other members of the
<taxonomicName id="04A41E2C1D62A9A0A1F48273286BAE78" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella margaritifera" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="margaritifera">Rhinella margaritifera</taxonomicName>
group by the absence of protruding vertebral apophyses, canthus rostralis not raised, snout projected, and low cranial crests.
<taxonomicName id="F364D02FDCDC507A3235ABDEE7536BC1" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella acutirostris" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="acutirostris">Rhinella acutirostris</taxonomicName>
differs from
<taxonomicName id="1D0BD3E7CBE7D873B24B7FE6D43A52F5" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
in having a bony knob at the angle of jaws (bony knob absent in
<taxonomicName id="C5DF9B23465C3E74BC38E01BD0F89D47" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
[
<bibRefCitation id="B0E5470B10849649BE78E1258C3224E8" author="Hoogmoed, MS" editor="Rocek, Z" journalOrPublisher="Prague" pageId="19" pageNumber="128" pagination="147 - 150" title="Biosystematic studies of the Bufo &quot; typhonius &quot; group. A preliminary progress report." volumeTitle="Studies in Herpetology" year="1986">Hoogmoed 1986</bibRefCitation>
,
<bibRefCitation id="FA78C4FC7BFB9B5B55F879DC7C71450F" author="Loetters, S" journalOrPublisher="Spixiana" pageId="20" pageNumber="129" pagination="293 - 303" title="A new toad of the Bufotyphonius complex from humid montane forests of Bolivia." volume="23" year="2000">
<normalizedToken id="341462FE381A40FB0015E44AF573148D" originalValue="Lötters">Loetters</normalizedToken>
and
<normalizedToken id="D9911F27E6DAC6846AE0CD526A1E69C0" originalValue="Köhler">Koehler</normalizedToken>
2000
</bibRefCitation>
]).
<taxonomicName id="04F50F5E45E3302118255703D1597E88" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
differs from the holotype of
<taxonomicName id="5FFEF62389D427BC3D54018D48048F44" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella proboscidea" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="proboscidea">Rhinella proboscidea</taxonomicName>
(ZSM 1145/0) in having a less protruding snout and skin on dorsum bearing a mixture of warts, pustules, and minute tubercles (smooth skin in
<taxonomicName id="5BE73D460352134FC2B75B2CE9D63BE3" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella proboscidea" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="proboscidea">Rhinella proboscidea</taxonomicName>
).
<taxonomicName id="E96C6C300A8B85F2F7B78FC47F22CF9B" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella dapsilis" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="dapsilis">Rhinella dapsilis</taxonomicName>
is much larger than
<taxonomicName id="5D70D05D198E72D708106B5AB3A26EE5" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
(
<taxonomicName id="525FD6C019FD090CCBBC17ACED919D5A" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella dapsilis" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="dapsilis">Rhinella dapsilis</taxonomicName>
holotype SVL = 77 mm, adult male;
<bibRefCitation id="B927B0D3C7956236A4F968048026810B" author="Myers, GS" journalOrPublisher="Boletim do Museu Nacional" pageId="20" pageNumber="129" pagination="1 - 24" title="Notes on some new or little-known Brazilian amphibians, with an examination of the history of the Plata salamander, Ensatinaplatensis." volume="35" year="1945">Myers and Carvalho 1945</bibRefCitation>
) and has a fleshy proboscis in the snout (proboscis absent in
<taxonomicName id="6F1DBBF3FD4819C6D3BD1196E3D6E355" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
).
<taxonomicName id="1BB7BBA271F7E86B29A976C905AAB450" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
differs from
<taxonomicName id="8F5F3416D4583EB1CB9278782C349211" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella yunga" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="yunga">Rhinella yunga</taxonomicName>
in having tympanic membrane and annulus distinct (tympanic membrane and annulus absent in
<taxonomicName id="FABBA69E6B1ACCDDB8EF9F9C9C012241" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella yunga" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="yunga">Rhinella yunga</taxonomicName>
;
<bibRefCitation id="86ECAC76B9C5625EDA0D7E07A506899B" author="Moravec, J" journalOrPublisher="ZooKeys" pageId="20" pageNumber="129" pagination="35 - 56" title="A new species of the Rhinellamargaritifera species group (Anura, Bufonidae) from the montane forest of the Selva Central, Peru." url="10.3897/zookeys.371.6580" volume="17" year="2014">Moravec et al. 2014</bibRefCitation>
).
<taxonomicName id="1643484C4E9DD9855E6FCF558BAA6E74" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella hoogmoedi" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="hoogmoedi">Rhinella hoogmoedi</taxonomicName>
,
<taxonomicName id="D072A13040426B9FF27A86FEEADC2C0C" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella magnussoni" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="magnussoni">Rhinella magnussoni</taxonomicName>
,
<taxonomicName id="062E233616DC079C856DFB9A0F665250" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella martyi" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="martyi">Rhinella martyi</taxonomicName>
,
<taxonomicName id="11BCFDD7D040CC91BAF56C30E66D6573" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella paraguayensis" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="paraguayensis">Rhinella paraguayensis</taxonomicName>
,
<taxonomicName id="DFF2541CF73384C8E4B65A9E42C39744" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella scitula" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="scitula">Rhinella scitula</taxonomicName>
,
<taxonomicName id="FD1A6D7F54915100EDBE90B707888C8C" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella sclerocephala" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="sclerocephala">Rhinella sclerocephala</taxonomicName>
, and
<taxonomicName id="7168D27A5560C6AAF40C53954E207FE8" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella stanlaii" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="stanlaii">Rhinella stanlaii</taxonomicName>
have a bony knob at angle of jaws (
<bibRefCitation id="540C13CA7CA030125728641ABE795113" author="Caramaschi, U" journalOrPublisher="Papeis Avulsos de Zoologia" pageId="18" pageNumber="127" pagination="251 - 259" title="A new species of Rhinella Fitzinger, 1826 from the Atlantic rain forest, eastern Brazil (Amphibia, Anura, Bufonidae)." url="10.1590/S0031-10492006002300001" volume="46" year="2006">Caramaschi and Pombal 2006</bibRefCitation>
,
<bibRefCitation id="5FF110F299A8DCDCEC01E878FBADD866" author="Lima, AP" journalOrPublisher="Zootaxa" pageId="20" pageNumber="129" pagination="1 - 15" title="A new species of Rhinella (Anura: Bufonidae) from Brazilian Amazon." volume="1663" year="2007">Lima et al. 2007</bibRefCitation>
,
<bibRefCitation id="1195D6FA377078B558C82BB73CCBEF2F" author="Fouquet, A" journalOrPublisher="Zootaxa" pageId="19" pageNumber="128" pagination="17 - 32" title="Description of two new species of Rhinella (Anura: Bufonidae) from the lowlands of the Guiana Shield." volume="1663" year="2007 a">Fouquet et al. 2007a</bibRefCitation>
,
<bibRefCitation id="3BC83B6F7803DD8342D2ABA490AA4FA3" author="Avila, RW" journalOrPublisher="Zootaxa" pageId="18" pageNumber="127" pagination="57 - 68" title="A new species of the Rhinellamargaritifera group (Anura: Bufonidae) from Brazilian Pantanal." volume="2339" year="2010">
<normalizedToken id="6F4C51A6D52B5A402956CDE074DDED2A" originalValue="Ávila">Avila</normalizedToken>
et al. 2010
</bibRefCitation>
,
<bibRefCitation id="C002729B028AAF89E63A4180768C23C2" author="Caramaschi, U" journalOrPublisher="Boletim do Museu Nacional" pageId="18" pageNumber="127" pagination="1 - 16" title="Nova especie do complexo de Bufomargaritifer (Laurenti, 1768) do Estado do Mato Grosso do Sul, Brasil (Amphibia, Anura, Bufonidae)." volume="501" year="2003">Caramaschi and Niemeyer 2003</bibRefCitation>
,
<bibRefCitation id="55B09F45D4C02EEF3FE9740F43F133CA" author="Mijares-Urrutia, A" journalOrPublisher="Herpetologica" pageId="20" pageNumber="129" pagination="523 - 531" title="A new toad of the Bufomargaritifer complex (Amphibia: Bufonidae) from northwestern Venezuela." volume="57" year="2001">Mijares-Arrutia and Arends-R 2001</bibRefCitation>
,
<bibRefCitation id="F42D2654AE6AB656FCC3B401EEE6D03A" author="Loetters, S" journalOrPublisher="Spixiana" pageId="20" pageNumber="129" pagination="293 - 303" title="A new toad of the Bufotyphonius complex from humid montane forests of Bolivia." volume="23" year="2000">
<normalizedToken id="72DE7102E97303FD4D8B56A6A19EC6D4" originalValue="Lötters">Loetters</normalizedToken>
and
<normalizedToken id="98DF40EAA83F8F09FADC4FC7944E38A7" originalValue="Köhler">Koehler</normalizedToken>
2000
</bibRefCitation>
; bony knob absent in
<taxonomicName id="F471633D7BBA538AE31BBFAE41C48469" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
).
<taxonomicName id="81201A7F7EDFABE182E3F4C93D0FBE24" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
differs from
<taxonomicName id="E68EED3FE0C7534A9B1934BA310FE23F" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella castaneotica" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="castaneotica">Rhinella castaneotica</taxonomicName>
,
<taxonomicName id="EAAB20A49505E42EF2E059CDF1589E8D" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella margaritifera" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="margaritifera">Rhinella margaritifera</taxonomicName>
(sensu stricto) and
<taxonomicName id="8F30B6CEE45FA17F4B6B852DF7C897EC" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella roqueana" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="roqueana">Rhinella roqueana</taxonomicName>
, by the absence of protruding vertebral apophyses (present in
<taxonomicName id="1C987446ED11AF8BFB78FA8D3041DF34" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella castaneotica" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="castaneotica">Rhinella castaneotica</taxonomicName>
[
<bibRefCitation id="E8A79D2183E44BC9A4ED7605EBA3CF9F" author="Caldwell, JP" journalOrPublisher="Papeis Avulsos de Zoologia" pageId="18" pageNumber="127" pagination="389 - 400" title="A new species of toad in the genus Bufo from Para, Brazil, with an unusual breeding site." volume="37" year="1991">Caldwell 1991</bibRefCitation>
],
<taxonomicName id="46788D55686E772F24CF108930D4CB9E" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella margaritifera" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="margaritifera">Rhinella margaritifera</taxonomicName>
[
<bibRefCitation id="C114BD0FDA943153FDA75E655DFC89B7" author="Lavilla, EO" journalOrPublisher="Zootaxa" pageId="19" pageNumber="128" pagination="251 - 264" title="The identity of Ranamargaritifera Laurenti, 1768 (Anura, Bufonidae)." url="10.11646/zootaxa.3646.3.4" volume="3646" year="2013">Lavilla et al. 2013</bibRefCitation>
], and
<taxonomicName id="69D804A43F2C2438C349E6C73ABFCCB2" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella roqueana" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="roqueana">Rhinella roqueana</taxonomicName>
[
<bibRefCitation id="6833A0D7CFE5ABB56DD92BF50EAE0129" author="Melin, DE" journalOrPublisher="Handlingar. Serien B, Matematiska och Naturvetenskapliga Skrifter" pageId="20" pageNumber="129" pagination="1 - 71" title="Contributions to the knowledge of the Amphibia of South America. Goeteborgs Kungl. Vetenskaps-och Vitterhets-samhaelles." volume="1" year="1941">Melin 1941</bibRefCitation>
]).
</paragraph>
<paragraph id="6EF5756F294CA7AE2F4579197BF3A6C0" pageId="11" pageNumber="120">
<taxonomicName id="C7B6FD449389295E0D55241372B98B55" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
is most closely related to populations of
<taxonomicName id="DAB3440B56A9AC0313759E5102651FEC" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella margaritifera" order="Anura" pageId="11" pageNumber="120" phylum="Chordata" rank="species" species="margaritifera">Rhinella margaritifera</taxonomicName>
from the upper Amazon basin in Ecuador and Peru. They can be easily distinguished by differences in body size (Fig. 5; see morphometric comparisons section) and relative size of cranial crests (Fig. 6).
</paragraph>
</subSubSection>
<subSubSection id="1391CB8492D3BA8C53DBE72446EA7167" pageId="11" pageNumber="120" type="holotype">
<paragraph id="E7DD7925DC3BE45A6E160B82914FF770" pageId="11" pageNumber="120">Holotype.</paragraph>
<paragraph id="7C04683E53FB9FCDB0D5FD38910046CB" pageId="11" pageNumber="120">
The holotype is an adult male with SVL = 39.2 mm (Fig. 11). Descriptions of the holotype have been provided by
<bibRefCitation id="2CFE0EEDD4DCC311AE1C9371E7ADBC7A" author="Leavitt, BB" journalOrPublisher="Copeia" pageId="19" pageNumber="128" pagination="7 - 8" title="On three races of Bufotyphonius." url="10.2307/1436176" volume="1933" year="1933">Leavitt (1933)</bibRefCitation>
and
<bibRefCitation id="6D8FA045A15F352E67A54EE06E7A3F44" author="Hoogmoed, MS" journalOrPublisher="Societat Catalana d'Ictiologia i Herpetologia" pageId="19" pageNumber="128" pagination="167 - 180" title="South American bufonids (Amphibia: Anura: Bufonidae), an enigma for taxonomists." volume="2" year="1989">Hoogmoed (1989)</bibRefCitation>
. The bony knob at angle of jaws and vertebral apophyses are absent. The crests are low and thick. There is a dorsolateral row of conical tubercles from the posterior border of the parotoid gland to the groin. There is a clear mid-dorsal line from the snout to the vent. The tympanum is rounded.
</paragraph>
</subSubSection>
<subSubSection id="E066DD0561529BC1D2369742B66C1CDB" lastPageId="13" lastPageNumber="122" pageId="11" pageNumber="120" type="variation">
<paragraph id="71F0CD2DE6542ACA1B2CA83DBA46B791" pageId="11" pageNumber="120">Variation.</paragraph>
<paragraph id="C6D63F5F5C51438BA53AAA8D06E251A4" lastPageId="12" lastPageNumber="121" pageId="11" pageNumber="120">
Variation in dorsal and ventral coloration of preserved specimens is shown in Figures 12 and 13. Background dorsal coloration varies from light gray (QCAZ 37244, AMNH 88689), light brown (QCAZ 14607, AMNH 104454) to dark gray (QCAZ 6733) or dark brown (QCAZ 11598, AMNH 52744), with irregular black and yellowish marks (QCAZ 4444, AMNH 88690). Some specimens
<pageBreakToken id="943C2B1BCAEACF83F15AB684DEF18320" pageId="12" pageNumber="121" start="start">have</pageBreakToken>
nearly uniform brown dorsum without marks (QCAZ 31603, 10296, AMNH 10296). A clear mid-dorsal line is often present (e.g. QCAZ 3502, QCAZ 12233).
</paragraph>
<caption id="B6A952CD0CDC8474E8F4BCE81F3A2751" pageId="12" pageNumber="121">
<paragraph id="1CA8C2A7A2517E823E399315E84ACECB" pageId="12" pageNumber="121">
Figure 12.
<taxonomicName id="DA1F095D032DDD64D7AE9F8552960306" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="12" pageNumber="121" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
from Ecuador showing variation in dorsal and ventral coloration of preserved specimens. Left to right, males: QCAZ 6733 (SVL 38.23 mm), QCAZ 10279 (SVL 35.08 mm); females, QCAZ 11598 (SVL 42.13 mm), QCAZ 14607 (SVL 50.95 mm), QCAZ 10439 (SVL 47.06 mm). See Appendix 1 for locality data. Not shown at the same scale.
</paragraph>
</caption>
<caption id="540B6D2D4F83991C0FEC0F19B3A06C4A" pageId="12" pageNumber="121">
<paragraph id="9125BF66489FFE345E04F6EA83C3FEBD" pageId="12" pageNumber="121">
Figure 13.
<taxonomicName id="D5222E26DB1897A45CB0081AE892556E" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="12" pageNumber="121" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
from Panama showing variation in dorsal and ventral coloration of preserved specimens. Left to right, male: AMNH 89459 (SVL 37.54 mm); females, AMNH 88694 (SVL 41.21 mm), AMNH 55476 (SVL 41.19 mm), AMNH 104454 (SVL 49.69 mm), AMNH 88689 (SVL 42.75 mm), AMNH 20896 (SVL 42.98 mm). See Appendix 1 for locality data. Not shown at the same scale.
</paragraph>
</caption>
<paragraph id="75BE2AF2348F78DFE1F79A2D804BBC99" lastPageId="13" lastPageNumber="122" pageId="12" pageNumber="121">
Ventral surfaces of preserved specimens have a cream to yellowish-cream background color with irregular darker marks arranged in diverse patterns; marks can
<pageBreakToken id="AFD43E811F03D0872444B5B2AF5DD11C" pageId="13" pageNumber="122" start="start">be</pageBreakToken>
light gray (QCAZ 6734, AMNH 88689), light brown (QCAZ 6732, AMNH 104454), dark gray (QCAZ 31606) or dark brown (QCAZ 6733, AMNH 89459), and vary from being restricted to the anterior half of the body (QCAZ 31604, AMNH 89459) to being present over the entire venter (QCAZ 4445, AMNH 88694). A longitudinal mid-ventral cream thin stripe can be present in the gular region (QCAZ 31602, 31606) or from the gular region to the mid-venter (QCAZ 6731, 11598).
</paragraph>
<paragraph id="53990DCC6BC66893DD28062891772657" pageId="13" pageNumber="122">Head shape in dorsal view varies from elongated (QCAZ 11598, AMNH 89459) to subtriangular (QCAZ 4447, AMNH 55475); in lateral view it varies from rounded (QCAZ 31605, AMNH 52749) to protruding (QCAZ 11393, AMNH 55475). Canthal region coloration varies from light gray or light brown to dark gray or dark brown. In some individuals the area below the eye and tympanum is yellowish cream (QCAZ 4447, AMNH 20896) or brown (QCAZ 31603, AMNH 88694) and differs from the color of the dorsum. Cloacal tubercles vary from yellowish cream (QCAZ 4441, AMNH 20896), to gray (QCAZ 31606) or brown (QCAZ 31602, AMNH 88695).</paragraph>
</subSubSection>
<subSubSection id="A199E1764E23B4432EAD49A491B95E13" lastPageId="14" lastPageNumber="123" pageId="13" pageNumber="122" type="color in life">
<paragraph id="6A8447B3A4E3B4D97FB3BAE1102ED562" pageId="13" pageNumber="122">Color in life.</paragraph>
<paragraph id="2AA60BBE3300FDCC502A10FFE4D0ABC0" pageId="13" pageNumber="122">Based on digital photograph of an adult female QCAZ 50568 (Fig. 8). Dark brown dorsum with irregular light brown and yellowish marks; there is a clear mid-dorsal line. Dorsal surfaces of tights and shanks are dark brown with transversal brown bands. Dorsal surfaces of forelimbs are dark brown with irregular light brown marks. Dark brown tubercles are abundant on the dorsum. Ventral surfaces vary from light brown to dark brown, with some irregularly distributed white and orange spots. The fingertips and the subarticular tubercles on fingers and toes are red-orange. Canthal region and tympanum are dark brown; iris greenish yellow with black reticulation.</paragraph>
<paragraph id="34163D2E9E3F7216BE723599F30311C8" pageId="14" pageNumber="123">
<pageBreakToken id="FC93E40C98CB278E72FCEE34D286EC79" pageId="14" pageNumber="123" start="start">Based</pageBreakToken>
on a digital photography of an adult male QCAZ 37248 (Fig. 8). Light brown dorsum with black spots and light brown and light gray marks. Dorsal surfaces of tights, shanks and forelimbs are light brown with transversal dark brown bands. Brown tubercles are abundant on the dorsum. Ventral surfaces are dark brown with irregularly distributed yellowish marks; the posterior part of the venter is cream. The subarticular tubercles of palms, soles, and fingertips are red-orange. Canthal region and tympanum are dark brown; iris greenish yellow with black reticulation.
</paragraph>
</subSubSection>
<subSubSection id="E41A3ED1DABA1D5FE711F5FAD2D31A12" lastPageId="15" lastPageNumber="124" pageId="14" pageNumber="123" type="distribution and ecology">
<paragraph id="0930A729D08FB2873BD944CEDDFBE585" pageId="14" pageNumber="123">Distribution and ecology.</paragraph>
<paragraph id="FFDB7A9423A60976169280B2FFEE7577" pageId="14" pageNumber="123">
<taxonomicName id="8054A3D4819EC5BA36F5D685973A43DA" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="14" pageNumber="123" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
has been recorded at 37 localities in the Ecuadorian
<normalizedToken id="173B99E82D91A661E876346144014110" originalValue="Chocó">Choco</normalizedToken>
(
<normalizedToken id="6D3DC7E94B1DA0F5E41A4B5785835C29" originalValue="Cañar">Canar</normalizedToken>
, Carchi, El Oro, Esmeraldas,
<normalizedToken id="7E48DC9CF07A2B08CA16FA7BAA693082" originalValue="Manabí">Manabi</normalizedToken>
, Pichincha, and Santo Domingo Provinces; Fig. 1), one locality in the Colombian
<normalizedToken id="144B92A242473997A76D7108398E3473" originalValue="Chocó">Choco</normalizedToken>
(Barbacoas,
<normalizedToken id="1EE19E920543F4E1FB37267406079911" originalValue="Nariño">Narino</normalizedToken>
; see Taxonomic remarks) and 35 localities in Panama (Comarca Guna Yala and Provinces
<normalizedToken id="442C8AD3425D86F64C2EEA8E298907A2" originalValue="Coclé">Cocle</normalizedToken>
,
<normalizedToken id="98627B4748B4FB68877E83F9A745FEBB" originalValue="Colón">Colon</normalizedToken>
,
<normalizedToken id="910A75CE728C9D04DDB619F3BC38DBE3" originalValue="Darién">Darien</normalizedToken>
and Panama; Fig. 2). It has a wide elevation range, from 19 to 1500 m above sea level.
</paragraph>
<paragraph id="2DEEC67896CA4C40AD3F039022889E46" pageId="14" pageNumber="123">
The examined specimens from Chocoan populations contain 21 gravid females (average SVL = 45.37 mm, SD = 4.05 mm): QCAZ 4262, QCAZ 4441, QCAZ 4442, QCAZ 4443, QCAZ 7065, QCAZ 10296, QCAZ 11597, QCAZ 11598 collected in January; QCAZ 50568 collected in February; QCAZ 11392, QCAZ 31601, QCAZ 31603, QCAZ 31605 collected in April; QCAZ 25023 collected in June; QCAZ 10439 collected in August; QCAZ 14607 collected in November; QCAZ 10301 collected in December. This suggests year round reproductive activity with a peak between January and April, a period that corresponds to the rainy season in the Ecuadorian
<normalizedToken id="88F30C9A52367EA412FBE67721EDDECF" originalValue="Chocó">Choco</normalizedToken>
.
</paragraph>
<paragraph id="D14DF89E69C47EDE23664AC614CC2A03" pageId="14" pageNumber="123">
In Panamanian populations gravid females were found in January (AMNH 104454), September (AMNH 55461), November (AMNH 88689), and December (AMNH 53699). In central Panama,
<taxonomicName id="29E6E7AF9B188F953C08DB912268CFA2" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="14" pageNumber="123" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
breeds in ponds and pools along permanent streams or swamps. Reproduction is explosive and most takes place from the middle of the rainy season to early dry season (
<bibRefCitation id="0568ECA47C12AAD1D23DB1791C028EE2" author="Wells, KD" journalOrPublisher="Biotropica" pageId="21" pageNumber="130" pagination="301 - 7" title="Reproductive behavior and male mating success in a Neotropical toad, Bufotyphonius." url="10.2307/2387922" volume="11" year="1979">Wells 1979</bibRefCitation>
,
<bibRefCitation id="3CC524620F47EC051702961D42544CC3" author="Ibanez, DR" journalOrPublisher="Editorial Mizrachi and Pujol, Panama" pageId="19" pageNumber="128" title="Los Anfibios del Monumento Natural Barro Colorado, Parque Nacional Soberania y Areas Adyacentes / The Amphibians of Barro Colorado Nature Monument, Soberania National Park and Adjacent Areas" year="1999">
<normalizedToken id="69DF87582BF31C197769EDAEC8B6B332" originalValue="Ibáñez">Ibanez</normalizedToken>
et al. 1999
</bibRefCitation>
). Choruses last less than 24 hours with males usually calling at night and oviposition occurring by day, especially in the early afternoon (
<bibRefCitation id="64588DEEA813F4C9DC86707CD53197D0" author="Wells, KD" journalOrPublisher="Biotropica" pageId="21" pageNumber="130" pagination="301 - 7" title="Reproductive behavior and male mating success in a Neotropical toad, Bufotyphonius." url="10.2307/2387922" volume="11" year="1979">Wells 1979</bibRefCitation>
). Otherwise, individuals are primarily diurnal, found active on the leaf litter of the forest floor during daytime, and often found asleep on leaves of low vegetation at night (
<bibRefCitation id="6416E6D58C9673205073CDDAEDE87DC0" author="Ibanez, DR" journalOrPublisher="Editorial Mizrachi and Pujol, Panama" pageId="19" pageNumber="128" title="Los Anfibios del Monumento Natural Barro Colorado, Parque Nacional Soberania y Areas Adyacentes / The Amphibians of Barro Colorado Nature Monument, Soberania National Park and Adjacent Areas" year="1999">
<normalizedToken id="50D6E82AE1BC401EFCABB7B7CE574E21" originalValue="Ibáñez">Ibanez</normalizedToken>
et al. 1999
</bibRefCitation>
). Diet is specialized on ants (
<bibRefCitation id="0D4467CA6A4928661FC762B03EBA53CD" author="Toft, CA" journalOrPublisher="Journal of Herpetology" pageId="21" pageNumber="130" pagination="139 - 144" title="Feeding ecology of Panamanian litter anurans: patterns in diet and foraging mode." url="10.2307/1563372" volume="15" year="1981">Toft 1981</bibRefCitation>
).
</paragraph>
<paragraph id="944707D9F6C8B90498DF91AB6579FCCF" pageId="14" pageNumber="123">
Most of the Ecuadorian specimens are from Reserva Mayronga and Reserva
<normalizedToken id="4242697EB6119C6D1DC349C1B6222C0E" originalValue="Ecológica">Ecologica</normalizedToken>
Cotacachi-Cayapas. They were found in the leaf litter of secondary forest and in agricultural lands. Some adults were observed at night within the forest in vegetation above the ground and some were found in amplexus (QCAZ 10271, QCAZ 10274, QCAZ 10275 in November 1996, and QCAZ 31604, QCAZ 31605 in February 1996). All the specimens collected in Reserva
<normalizedToken id="42AA51AD8AABC0DB8E09DB7927903C4D" originalValue="Ecológica">Ecologica</normalizedToken>
Cotacachi-Cayapas were found in secondary forest. At some collecting sites, the forest has been cleared for cacao plantations (QCAZ specimen database).
</paragraph>
<paragraph id="68313670958E0C39ADBCEFFD6DA12DEC" lastPageId="15" lastPageNumber="124" pageId="14" pageNumber="123">
According to the classification of
<bibRefCitation id="3CFB693D27A768EF4B79C40746E27EDB" author="Sierra, R" journalOrPublisher="Proyecto INEFAN / GEF-BIRF, Wildlife Conservation Society y Ecociencia, Quito" pageId="20" pageNumber="129" title="Mapa de vegetacion del Ecuador Continental 1: 1 ' 000.000." year="1999">Sierra et al. (1999)</bibRefCitation>
the vegetation types for Ecuadorian localities are: (1) Lowland Evergreen Forest of Coastal Range, characterized by abundant epiphytes, climbers and herbaceous plants, with a canopy of 30 m (e.g. Reserva La Chiquita, Durango); (2) Semideciduous Lowland Forest of Coastal Range, defined by
<pageBreakToken id="5D781ADB6A53BCADBBF1BB4F352252BE" pageId="15" pageNumber="124" start="start">the</pageBreakToken>
presence of broad canopy trees up to 20 m and curved shafts; the tree stratum is characterized by the presence of spiny, deciduous species with epiphytes while the forest floor has herbaceous plants (e.g. Bilsa, La Tortuga); (3) Evergreen Foothill Forest of Coastal Range, characterized by a canopy that can reach 30 m or more and trunks of trees covered with orchids, bromeliads, ferns and aroids (e.g. Manta Real, Alto Tambo); (4) Deciduous Lowland Forest of Costal Range, characterized by losing leaves during part of the year with a great varieties of cactus and thorny plants; the most conspicuous trees are the family
<taxonomicName id="99F3175C89A4B99506A654D37EECE68B" family="Bombacaceae" lsidName="Rhinella" pageId="15" pageNumber="124" rank="family">Bombacaceae</taxonomicName>
have curved trunks and broad crown. (e.g. El Progreso); (5) Semideciduos Foothill Forest of Coastal Range, characterized by having slightly dispersed vegetation, with trees over 20 m and dense herbaceous layers of ferns (e.g. Valle Hermoso).
</paragraph>
<paragraph id="0C4DCC7E7BC98C19248815FD8077887A" pageId="15" pageNumber="124">
The main vegetation types for Panamanian localities are (following
<bibRefCitation id="D7D2AACC7387B234FB87704C82FCA596" author="Hogan, MC" editor="Cleveland, CJ" journalOrPublisher="Environmental Information Coalition, National Council for Science and the Environment" pageId="19" pageNumber="128" title="Ecoregions of Panama." url="http://www.eoearth.org/view/article/51cbed867896bb431f692a98/" volumeTitle="Encyclopedia of Earth" year="2010">Hogan 2010</bibRefCitation>
): (1) Isthmian-Atlantic Moist Forests, characterized by tall tropical evergreen forest with buttressed canopy trees reaching 40 m and with an extremely rich epiphyte flora (e.g. Cruces Trail, Punta
<normalizedToken id="692F3975802E5E91E45F05D0B8440739" originalValue="Rincón">Rincon</normalizedToken>
); (2) Eastern Panamanian Montane Forest, at elevations from 500 to 1800 m above sea level, includes marshes, swamp forests, semi-deciduous tropical moist forests, premontane wet forest, cloud forests and elfin forests (e.g. Cana, Cerro Tacarcuna); (3)
<normalizedToken id="C0B922481BC70456B793680C8687DBE8" originalValue="Chocó-Darién">Choco-Darien</normalizedToken>
Moist Forests, at elevations between 0 and 1000 m above sea level, between the Pacific Ocean and the western range of the Andes (e.g. Dad Nakue Dubpir, Udirbi).
</paragraph>
</subSubSection>
<subSubSection id="E0D7292DF29D9007408BE51358A71473" pageId="15" pageNumber="124" type="taxonomic remarks">
<paragraph id="B952B8BE9E81A69DBE5E9ABB75E6DCC0" pageId="15" pageNumber="124">Taxonomic remarks.</paragraph>
<paragraph id="03542191D614EC4D49E7A9E02009A3AB" pageId="15" pageNumber="124">
Based on morphological characters,
<bibRefCitation id="F4991BFEBBE6B846E97421EC5CDF2D5A" author="Velez-Rodriguez, CM" journalOrPublisher="Universidad Austral de Chile, Valdivia, Chile" pageId="21" pageNumber="130" title="Sistematica de los sapos neotropicales pertenecientes al grupo Bufotyphonius (Amphibia: Bufonidae)" year="2004">
<normalizedToken id="0CD2F96D57C5C0EBE101003BA9391270" originalValue="Vélez-Rodriguez">Velez-Rodriguez</normalizedToken>
(2004)
</bibRefCitation>
ascribed four populations from Panama and Colombia to
<taxonomicName id="1EC91FC89C72ED35A334AA0648C56BF8" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="15" pageNumber="124" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
: Isthmus of Panama (Panama; 15 males, 10 females); Parque Nacional Los
<normalizedToken id="C74264F0441D587EDA58B3EABA33AFC2" originalValue="Katíos">Katios</normalizedToken>
(Colombia; 12 males, 15 females); Gorgona and
<normalizedToken id="EB50F7711BECFA2F6BA2C63D001EF820" originalValue="Güape">Gueape</normalizedToken>
Island (Colombia; 7 males, 8 females); Municipio Restrepo (Colombia; 7 males, 8 females). Based on data from
<bibRefCitation id="2BC87D76755D022B98012F07CD725973" author="Velez-Rodriguez, CM" journalOrPublisher="Universidad Austral de Chile, Valdivia, Chile" pageId="21" pageNumber="130" title="Sistematica de los sapos neotropicales pertenecientes al grupo Bufotyphonius (Amphibia: Bufonidae)" year="2004">
<normalizedToken id="63C327BCF4E8A2AB1E7C3851FCC0BD1B" originalValue="Vélez-Rodriguez">Velez-Rodriguez</normalizedToken>
(2004)
</bibRefCitation>
, these populations differ from the holotype of
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and populations of
<taxonomicName id="9E54D428CB05A669B218C8471AB22EF2" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="15" pageNumber="124" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
in Ecuador and Panama (in parentheses) in having: (1) a canthus rostralis protruding in females and ill-defined in males (inconspicuous in males and females), (2) parietal crests well defined in females, ill-defined in males (ill-defined in males and females), (3) vertebral apophyses slightly visible externally (absent). The differences suggest that those specimens are not
<taxonomicName id="3A1D9485277FF80A45F93252A57D6F6C" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="15" pageNumber="124" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
and may belong to a different species. Alternatively, differences between
<taxonomicName id="BC76523C2C50D6F02E786E4115740D35" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="15" pageNumber="124" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
described by
<bibRefCitation id="39023D7DC89CC9D24B92059E72315C71" author="Velez-Rodriguez, CM" journalOrPublisher="Universidad Austral de Chile, Valdivia, Chile" pageId="21" pageNumber="130" title="Sistematica de los sapos neotropicales pertenecientes al grupo Bufotyphonius (Amphibia: Bufonidae)" year="2004">
<normalizedToken id="3091A93D6DC298D82B7B50FD64B9742A" originalValue="Vélez-Rodriguez">Velez-Rodriguez</normalizedToken>
(2004)
</bibRefCitation>
and our study could be an artifact resulting from the use of distinct terminology for similar character states.
</paragraph>
<paragraph id="996551E8311688100324491F0945878D" pageId="15" pageNumber="124">
In contrast,
<bibRefCitation id="5375FF3BC4D31BEEF90A4DF783A2A2D0" author="Mueses-Cisneros, JJ" journalOrPublisher="Herpetotropicos" pageId="20" pageNumber="129" pagination="39 - 54" title="Fauna anfibia de la reserva natural biotopo selva humeda, Barbacoas, Narino, Colombia." volume="7" year="2012">Mueses-Cisneros and Moreno-Quintero (2012)</bibRefCitation>
reported two species of the
<taxonomicName id="6C6D7DE783DAB3410E51A2FBB27B3759" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella margaritifera" order="Anura" pageId="15" pageNumber="124" phylum="Chordata" rank="species" species="margaritifera">Rhinella margaritifera</taxonomicName>
group form Barbacoas,
<normalizedToken id="994302430082B198D6CBC8DCC5B962DC" originalValue="Nariño">Narino</normalizedToken>
, Colombia (
<taxonomicName id="C9E33E6DA896F8FAE6C944230275B1AF" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella" order="Anura" pageId="15" pageNumber="124" phylum="Chordata" rank="genus">Rhinella</taxonomicName>
sp. 9 and
<taxonomicName id="96510DFD4BD406780906C4147A3F3B2A" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella" order="Anura" pageId="15" pageNumber="124" phylum="Chordata" rank="genus">Rhinella</taxonomicName>
sp. 10). Two photographs of live individuals (pp. 45) show morphological features that fall within the observed variation of
<taxonomicName id="56570FEF6D2CA3C2F865465C10537FD2" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="15" pageNumber="124" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
. We tentatively assign them to
<taxonomicName id="CD3A21B688E089EC166F62BE19388748" class="Amphibia" family="Bufonidae" genus="Rhinella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rhinella alata" order="Anura" pageId="15" pageNumber="124" phylum="Chordata" rank="species" species="alata">Rhinella alata</taxonomicName>
but direct specimen examination is required to confirm this identification.
</paragraph>
</subSubSection>
</treatment>
</document>