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<document ID-DOI="10.1093/zoolinnean/zlac057" ID-GBIF-Dataset="a6a1a213-903b-4c6f-b317-50fc21657684" ID-ISSN="0024-4082" ID-Zenodo-Dep="7695481" ID-ZooBank="27C78E3C-CD39-4BA9-99D0-778D850368C7" checkinTime="1677744247128" checkinUser="plazi" docAuthor="Novaes-E-Fagundes, Gabriel, Lyra, Mariana L., Loredam, Vinicius S. A., Carvalho, Thiago R., Haddad, Célio F. B., Rodrigues, Miguel T., Baldo, Diego, Barrasso, Diego A., Loebmann, Daniel, Ávila, Robson W., Brusquetti, Francisco, Prudente, Ana L. C., Wheeler, Ward C., Orrico, Victor Goyannes Dill &amp; Peloso, Pedro" docDate="2023" docId="03D26765400B1D2ED7644454EBE30000" docLanguage="en" docName="zlac057.pdf" docOrigin="Zoological Journal of the Linnean Society 197" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-.journal_article" docStyleVersion="7" docTitle="Elachistocleis Parker 1927" docType="treatment" docVersion="2" lastPageNumber="555" masterDocId="FFEB1F1D40021D24D45F4756EF4E0678" masterDocTitle="A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis)" masterLastPageNumber="568" masterPageNumber="545" pageNumber="554" updateTime="1677842110288" updateUser="ExternalLinkService">
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<mods:title>A tale of two bellies: systematics of the oval frogs (Anura: Microhylidae: Elachistocleis)</mods:title>
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<mods:namePart>Novaes-E-Fagundes, Gabriel</mods:namePart>
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<mods:namePart>Lyra, Mariana L.</mods:namePart>
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<mods:namePart>Loredam, Vinicius S. A.</mods:namePart>
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<mods:namePart>Carvalho, Thiago R.</mods:namePart>
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<mods:namePart>Haddad, Célio F. B.</mods:namePart>
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<mods:namePart>Rodrigues, Miguel T.</mods:namePart>
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<mods:namePart>Baldo, Diego</mods:namePart>
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<mods:namePart>Barrasso, Diego A.</mods:namePart>
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<mods:namePart>Loebmann, Daniel</mods:namePart>
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<mods:namePart>Ávila, Robson W.</mods:namePart>
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<mods:namePart>Brusquetti, Francisco</mods:namePart>
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<mods:namePart>Orrico, Victor Goyannes Dill</mods:namePart>
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<subSubSection box="[827,1443,770,795]" pageId="9" pageNumber="554" type="nomenclature">
<paragraph blockId="9.[827,1443,770,795]" box="[827,1443,770,795]" pageId="9" pageNumber="554">
<heading box="[827,1443,770,795]" centered="true" fontSize="9" level="2" pageId="9" pageNumber="554" reason="2">
TAXONOMY OF
<taxonomicName ID-CoL="632QW" ID-ENA="143566" authorityName="Parker" authorityYear="1927" box="[997,1190,771,794]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="genus">
<emphasis box="[997,1190,771,794]" italics="true" pageId="9" pageNumber="554">ELACHISTOCLEIS</emphasis>
</taxonomicName>
A COMPLEX MATTER
</heading>
</paragraph>
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<subSubSection lastPageId="10" lastPageNumber="555" pageId="9" pageNumber="554" type="description">
<paragraph blockId="9.[827,1444,810,1906]" pageId="9" pageNumber="554">
<bibRefCitation author="Parker HW" box="[827,986,810,832]" pageId="9" pageNumber="554" pagination="1 - 6" refId="ref17854" refString="Parker HW. 1927. The brevicipitid frogs allied to the genus Gastrophryne. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 187: 1 - 6." type="journal article" year="1927">Parker (1927)</bibRefCitation>
erected
<taxonomicName authorityName="Parker" authorityYear="1927" box="[1086,1244,810,831]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="genus">
<emphasis box="[1086,1244,810,831]" italics="true" pageId="9" pageNumber="554">Elachistocleis</emphasis>
</taxonomicName>
to allocate
<taxonomicName authority="Schneider, 1799" authorityName="Schneider" authorityYear="1799" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="species" species="ovalis">
<emphasis italics="true" pageId="9" pageNumber="554">Rana ovalis</emphasis>
<bibRefCitation author="Schneider JG" box="[905,1092,841,863]" pageId="9" pageNumber="554" refId="ref18956" refString="Schneider JG. 1799. Historia amphibiorum naturalis et literarariae. Fasciculus primus. Continens ranas, calamitas, bufones, salamandras et hydros in genera et species descriptos notisque suis distinctos. Jena: Friederici Frommanni." type="book" year="1799">Schneider, 1799</bibRefCitation>
</taxonomicName>
and
<taxonomicName baseAuthorityName="Schneider" baseAuthorityYear="1799" box="[1154,1354,841,863]" class="Amphibia" family="Microhylidae" genus="Engystoma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="species" species="ovale">
<emphasis box="[1154,1354,841,863]" italics="true" pageId="9" pageNumber="554">Engystoma ovale</emphasis>
</taxonomicName>
<taxonomicName authority="(Guerin-Meneville, 1838)" baseAuthorityName="Guerin-Meneville" baseAuthorityYear="1838" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="species" species="bicolor">
<emphasis box="[1362,1442,841,862]" italics="true" pageId="9" pageNumber="554">bicolor</emphasis>
(
<bibRefCitation author="Guerin-Meneville F-E" box="[836,1118,872,894]" pageId="9" pageNumber="554" refId="ref16247" refString="Guerin-Meneville F-E. 1838. Iconographie du regne Animal de G. CuVier ou representation d'apres nature de l'une des especes les plus remarquables et souVent non enVore figurees, de chaque genre d'animaux, aVec un texte descriptiff mis au courant de la science, Vol. 3 (Part - Reptiles). Paris: J. B. Balliere." type="book" year="1838">Guérin-Méneville, 1838</bibRefCitation>
)
</taxonomicName>
. As usual at the time, the descriptions of both species were brief and did not designate
<typeStatus box="[939,988,934,955]" pageId="9" pageNumber="554">type</typeStatus>
specimens.
<bibRefCitation author="Lavilla EO &amp; Vaira M &amp; Ferrari L" box="[1117,1327,933,955]" pageId="9" pageNumber="554" pagination="269 - 284" refId="ref17160" refString="Lavilla EO, Vaira M, Ferrari L. 2003. A new species of Elachistocleis (Anura: Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis oValis - E. bicolor controversy. Amphibia-Reptilia 24: 269 - 284." type="journal article" year="2003">
Lavilla
<emphasis box="[1201,1253,933,955]" italics="true" pageId="9" pageNumber="554">et al.</emphasis>
(2003)
</bibRefCitation>
suggested that these names apply to complexes of species, because species of this genus are morphologically similar, with few obvious external characters that can be used as reliable diagnostic characters (i.e. non-overlapping and fixed). Ultimately, as mentioned throughout our contribution, diagnoses in this genus rely heavily on the ventral colour pattern, head proportions and the presence/absence and extension of body lines, such as a mid-dorsal white line and lines on the hidden surface of hindlimbs (
<bibRefCitation author="Parker HW" box="[1072,1215,1240,1262]" pageId="9" pageNumber="554" pagination="1 - 6" refId="ref17854" refString="Parker HW. 1927. The brevicipitid frogs allied to the genus Gastrophryne. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 187: 1 - 6." type="journal article" year="1927">Parker, 1927</bibRefCitation>
;
<bibRefCitation author="Lavilla EO &amp; Vaira M &amp; Ferrari L" box="[1227,1437,1240,1262]" pageId="9" pageNumber="554" pagination="269 - 284" refId="ref17160" refString="Lavilla EO, Vaira M, Ferrari L. 2003. A new species of Elachistocleis (Anura: Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis oValis - E. bicolor controversy. Amphibia-Reptilia 24: 269 - 284." type="journal article" year="2003">
Lavilla
<emphasis box="[1314,1370,1240,1261]" italics="true" pageId="9" pageNumber="554">et al.</emphasis>
, 2003
</bibRefCitation>
;
<bibRefCitation author="Caramaschi U" box="[827,1031,1270,1292]" pageId="9" pageNumber="554" pagination="1 - 30" refId="ref14635" refString="Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30." type="journal article" year="2010">Caramaschi, 2010</bibRefCitation>
;
<bibRefCitation author="Pereyra LC &amp; Akmentins MS &amp; Laufer G &amp; Vaira M" box="[1044,1263,1270,1292]" pageId="9" pageNumber="554" pagination="525 - 544" refId="ref18066" refString="Pereyra LC, Akmentins MS, Laufer G, Vaira M. 2013. A new species of Elachistocleis (Anura: Microhylidae) from north-western Argentina. Zootaxa 3694: 525 - 544." type="journal article" year="2013">
Pereyra
<emphasis box="[1140,1196,1270,1292]" italics="true" pageId="9" pageNumber="554">et al.</emphasis>
, 2013
</bibRefCitation>
).
</paragraph>
<paragraph blockId="9.[827,1444,810,1906]" lastBlockId="10.[145,762,197,1905]" lastPageId="10" lastPageNumber="555" pageId="9" pageNumber="554">
<bibRefCitation author="Caramaschi U" box="[851,1068,1301,1323]" pageId="9" pageNumber="554" pagination="1 - 30" refId="ref14635" refString="Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30." type="journal article" year="2010">Caramaschi (2010)</bibRefCitation>
and
<bibRefCitation author="Piva A &amp; Caramaschi U &amp; Albuquerque NR" box="[1127,1322,1301,1323]" pageId="9" pageNumber="554" pagination="143 - 154" refId="ref18209" refString="Piva A, Caramaschi U, Albuquerque NR. 2017. A new species of Elachistocleis (Anura: Microhylidae) from the Brazilian Pantanal. Phyllomedusa. Journal of Herpetology 16: 143 - 154." type="journal article" year="2017">
Piva
<emphasis box="[1186,1242,1301,1323]" italics="true" pageId="9" pageNumber="554">et al.</emphasis>
(2017)
</bibRefCitation>
suggested the ventral colour pattern as a diagnostic character for two internal species groups: a group with immaculate bellies (uniformly clear, free of markings) and another with maculate bellies. We found that the groups proposed by the above-cited authors are not monophyletic. For example,
<taxonomicName authorityName="Nunes-de-Almeida &amp; Toledo" authorityYear="2012" box="[1138,1442,1485,1507]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="species" species="muiraquitan">
<emphasis box="[1138,1442,1485,1507]" italics="true" pageId="9" pageNumber="554">Elachistocleis muiraquitan</emphasis>
</taxonomicName>
(immaculate-bellied) is not related to the remaining immaculate-bellied species; instead, it was recovered in the
<taxonomicName baseAuthorityName="Daudin" baseAuthorityYear="1802" box="[898,1080,1577,1599]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="species" species="surinamensis">
<emphasis box="[898,916,1578,1599]" italics="true" pageId="9" pageNumber="554">E</emphasis>
.
<emphasis box="[928,1080,1578,1599]" italics="true" pageId="9" pageNumber="554">surinamensis</emphasis>
</taxonomicName>
species group, a clade composed of species with predominantly maculate bellies. Furthermore, whereas
<taxonomicName box="[1092,1260,1639,1660]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="species" species="panamensis">
<emphasis box="[1092,1110,1639,1660]" italics="true" pageId="9" pageNumber="554">E</emphasis>
.
<emphasis box="[1123,1260,1639,1660]" italics="true" pageId="9" pageNumber="554">panamensis</emphasis>
</taxonomicName>
has a maculate belly (
<bibRefCitation author="Dunn ER &amp; Trapido H &amp; Evans H" box="[899,1101,1669,1691]" pageId="9" pageNumber="554" pagination="1 - 8" refId="ref15330" refString="Dunn ER, Trapido H, Evans H. 1948. A new species of the microhylid frog genus Chiasmocleis from Panama. American Museum NoVitates 1376: 1 - 8." type="journal article" year="1948">
Dunn
<emphasis box="[973,1031,1669,1691]" italics="true" pageId="9" pageNumber="554">et al.</emphasis>
, 1948
</bibRefCitation>
;
<bibRefCitation author="Nelson CE" box="[1115,1266,1669,1691]" pageId="9" pageNumber="554" pagination="895 - 898" refId="ref17614" refString="Nelson CE. 1972. Distribution and biology of Chiasmocleis panamensis (Amphibia: Microhylidae). Copeia 1972: 895 - 898." type="journal article" year="1972">Nelson, 1972</bibRefCitation>
),
<taxonomicName box="[1289,1393,1669,1691]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="species" species="araios">
<emphasis box="[1289,1307,1670,1691]" italics="true" pageId="9" pageNumber="554">E</emphasis>
.
<emphasis box="[1321,1393,1670,1691]" italics="true" pageId="9" pageNumber="554">araios</emphasis>
</taxonomicName>
has an immaculate belly (
<bibRefCitation author="Sanchez-Nivicela JC &amp; Peloso PLV &amp; Urgiles VL &amp; Munoz MH &amp; Sagredo Y &amp; Paez N &amp; Ron S" box="[1086,1427,1700,1722]" pageId="9" pageNumber="554" pagination="323 - 340" refId="ref18871" refString="Sanchez-Nivicela JC, Peloso PLV, Urgiles VL, Yanez- Munoz MH, Sagredo Y, Paez N, Ron S. 2020. Description and phylogenetic relationships of a new trans-Andean species of Elachistocleis Parker 1927 (Amphibia, Anura, Microhylidae). Zootaxa 4779: 323 - 340." type="journal article" year="2020">
Sánchez-Nivicela
<emphasis box="[1298,1357,1700,1721]" italics="true" pageId="9" pageNumber="554">et al.</emphasis>
, 2020
</bibRefCitation>
). Given their phylogenetic position as successive sistertaxa to the remaining species of the genus (but see our comments on the position of
<taxonomicName box="[1151,1252,1792,1813]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="species" species="araios">
<emphasis box="[1151,1169,1792,1813]" italics="true" pageId="9" pageNumber="554">E</emphasis>
.
<emphasis box="[1181,1252,1792,1813]" italics="true" pageId="9" pageNumber="554">araios</emphasis>
</taxonomicName>
), the description of
<taxonomicName box="[855,959,1823,1844]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="species" species="araios">
<emphasis box="[855,873,1823,1844]" italics="true" pageId="9" pageNumber="554">E</emphasis>
.
<emphasis box="[887,959,1823,1844]" italics="true" pageId="9" pageNumber="554">araios</emphasis>
</taxonomicName>
already rendered the immaculate-bellied species group as paraphyletic. Also, it implies that the optimization of the ventral pattern of the
<taxonomicName authorityName="Parker" authorityYear="1927" box="[1287,1442,1884,1905]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="9" pageNumber="554" phylum="Chordata" rank="genus">
<emphasis box="[1287,1442,1884,1905]" italics="true" pageId="9" pageNumber="554">Elachistocleis</emphasis>
</taxonomicName>
ancestor is ambiguous, given our topology and that of
<bibRefCitation author="Sanchez-Nivicela JC &amp; Peloso PLV &amp; Urgiles VL &amp; Munoz MH &amp; Sagredo Y &amp; Paez N &amp; Ron S" box="[145,495,228,250]" pageId="10" pageNumber="555" pagination="323 - 340" refId="ref18871" refString="Sanchez-Nivicela JC, Peloso PLV, Urgiles VL, Yanez- Munoz MH, Sagredo Y, Paez N, Ron S. 2020. Description and phylogenetic relationships of a new trans-Andean species of Elachistocleis Parker 1927 (Amphibia, Anura, Microhylidae). Zootaxa 4779: 323 - 340." type="journal article" year="2020">
Sánchez-Nivicela
<emphasis box="[355,413,228,250]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
(2020)
</bibRefCitation>
. More importantly, we found intraspecific variation of the ventral pattern in some species, such as
<taxonomicName baseAuthorityName="Guerin-Meneville" baseAuthorityYear="1838" box="[408,521,289,311]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="bicolor">
<emphasis box="[408,426,290,311]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[441,521,289,310]" italics="true" pageId="10" pageNumber="555">bicolor</emphasis>
</taxonomicName>
and
<taxonomicName box="[582,761,289,311]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="nigrogularis">
<emphasis box="[582,600,290,311]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[615,761,289,310]" italics="true" pageId="10" pageNumber="555">nigrogularis</emphasis>
</taxonomicName>
(
<figureCitation box="[154,233,320,342]" captionStart="Figure 1" captionStartId="5.[163,244,1810,1832]" captionTargetBox="[226,1379,195,1770]" captionTargetId="graphics-25@5.[226,1378,239,1770]" captionTargetPageId="5" captionText="Figure 1. Species delimitation in Elachistocleis. The tree shown is from the maximum likelihood (ML) analysis with only 16S + COI from only Elachistocleis samples (tree rooted in E. araios). Numbers on branches are SH-aLRT/UFBoot support values. Asterisks represent support above 80/95. The terminals of the tree are named as: original labelling of" figureDoi="http://doi.org/10.5281/zenodo.7695483" httpUri="https://zenodo.org/record/7695483/files/figure.png" pageId="10" pageNumber="555">Figs 1</figureCitation>
,
<figureCitation box="[252,266,320,341]" captionStart="Figure 4" captionStartId="12.[144,224,1106,1128]" captionTargetBox="[145,1428,195,1067]" captionTargetId="figure-410@12.[145,1428,195,1067]" captionTargetPageId="12" captionText="Figure 4. Variation of ventral colour pattern among sample vouchers of different species of Elachistocleis of the bicolor group. A, B, E. helianneae; MPEG 29416, 29419. C, E. haroi; IIBPH 2947. DF, E. bicolor; CFBH 10117, AAGUFU 5953, IIBPH 0285. G, H, E. nigrogularis; CAS 231811, 245101." figureDoi="http://doi.org/10.5281/zenodo.7695495" httpUri="https://zenodo.org/record/7695495/files/figure.png" pageId="10" pageNumber="555">4</figureCitation>
). Ontogenetic variation of the ventral pattern has also been recently reported for
<taxonomicName authority="(Bueno-Villafane et al., 2020)" baseAuthorityName="Bueno-Villafane" baseAuthorityYear="2020" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="haroi">
<emphasis box="[665,683,351,372]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[698,761,350,371]" italics="true" pageId="10" pageNumber="555">haroi</emphasis>
(
<bibRefCitation author="Bueno-Villafane D &amp; Caballlero-Gini A &amp; Ferreira M &amp; Netto F &amp; Fernandez Rios D &amp; Brusquetti F" box="[153,469,381,403]" pageId="10" pageNumber="555" pagination="191 - 200" refId="ref14581" refString="Bueno-Villafane D, Caballlero-Gini A, Ferreira M, Netto F, Fernandez Rios D, Brusquetti F. 2020. Ontogenetic changes in the ventral colouration of post metamorphic Elachistocleis haroi Pereyra, Akmentins, Laufer, Vaira, 2013 (Anura: Microhylidae). Amphibia-Reptilia 41: 191 - 200." type="journal article" year="2020">
Bueno-Villafañe
<emphasis box="[345,401,381,403]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2020
</bibRefCitation>
)
</taxonomicName>
. Therefore, the available phylogenetic and morphological evidence emphatically rejects the existence of groups diagnosed solely by the colour of their bellies.
</paragraph>
<paragraph blockId="10.[145,762,197,1905]" pageId="10" pageNumber="555">
It is noteworthy that this discussion seems far from settled. Ventral patterns have been recorded idiosyncratically in the literature.
<bibRefCitation author="Sanchez-Nivicela JC &amp; Peloso PLV &amp; Urgiles VL &amp; Munoz MH &amp; Sagredo Y &amp; Paez N &amp; Ron S" pageId="10" pageNumber="555" pagination="323 - 340" refId="ref18871" refString="Sanchez-Nivicela JC, Peloso PLV, Urgiles VL, Yanez- Munoz MH, Sagredo Y, Paez N, Ron S. 2020. Description and phylogenetic relationships of a new trans-Andean species of Elachistocleis Parker 1927 (Amphibia, Anura, Microhylidae). Zootaxa 4779: 323 - 340." type="journal article" year="2020">
Sánchez-Nivicela
<emphasis box="[145,211,596,617]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
(2020)
</bibRefCitation>
cite
<taxonomicName baseAuthorityName="MirandaRibeiro" baseAuthorityYear="1920" box="[374,496,596,617]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="cesarii">
<emphasis box="[374,392,596,617]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[412,496,596,617]" italics="true" pageId="10" pageNumber="555">cesarii</emphasis>
</taxonomicName>
as having uniform, immaculate, ventral colouration, although
<bibRefCitation author="Toledo LF &amp; Loebmann D &amp; Haddad CFB" pageId="10" pageNumber="555" pagination="50 - 60" refId="ref19506" refString="Toledo LF, Loebmann D, Haddad CFB. 2010. Revalidation and redescription of Elachistocleis cesarii (Miranda-Ribeiro, 1920) (Anura: Microhylidae). Zootaxa 2418: 50 - 60." type="journal article" year="2010">
Toledo
<emphasis box="[145,206,657,679]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
(2010)
</bibRefCitation>
when resurrecting the species clearly state that it has ventral colouration white or yellow with grey marks and reticulations, and their figure 3d depicts a specimen with a maculate belly.
<bibRefCitation author="Jowers MJ &amp; Othman SN &amp; Borzee A &amp; Rivas GA &amp; Ramirez S &amp; Auguste RJ &amp; Downie JR &amp; Read M &amp; Murphy JC" pageId="10" pageNumber="555" pagination="189 - 206" refId="ref16761" refString="Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity &amp; EVolution 21: 189 - 206." type="journal article" year="2021">
Jowers
<emphasis box="[705,761,749,771]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
(2021)
</bibRefCitation>
state that
<taxonomicName baseAuthorityName="MirandaRibeiro" baseAuthorityYear="1920" box="[347,455,780,801]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="cesarii">
<emphasis box="[347,365,780,801]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[379,455,780,801]" italics="true" pageId="10" pageNumber="555">cesarii</emphasis>
</taxonomicName>
has a uniform belly. This same was said of
<taxonomicName authorityName="Kwet &amp; Di-Bernardo" authorityYear="1998" box="[340,520,810,832]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="erythrogaster">
<emphasis box="[340,358,811,832]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[370,520,810,831]" italics="true" pageId="10" pageNumber="555">erythrogaster</emphasis>
</taxonomicName>
and
<taxonomicName baseAuthorityName="Guerin-Meneville" baseAuthorityYear="1838" box="[576,683,810,832]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="bicolor">
<emphasis box="[576,594,811,832]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[606,683,810,831]" italics="true" pageId="10" pageNumber="555">bicolor</emphasis>
</taxonomicName>
. While
<taxonomicName baseAuthorityName="MirandaRibeiro" baseAuthorityYear="1920" box="[145,249,842,863]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="cesarii">
<emphasis box="[145,163,842,863]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[175,249,842,863]" italics="true" pageId="10" pageNumber="555">cesarii</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Kwet &amp; Di-Bernardo" authorityYear="1998" box="[305,484,841,863]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="erythrogaster">
<emphasis box="[305,323,842,863]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[334,484,841,862]" italics="true" pageId="10" pageNumber="555">erythrogaster</emphasis>
</taxonomicName>
are maculate (the latter comparatively less maculate),
<taxonomicName baseAuthorityName="Guerin-Meneville" baseAuthorityYear="1838" box="[486,592,872,893]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="bicolor">
<emphasis box="[486,504,872,893]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[515,592,872,893]" italics="true" pageId="10" pageNumber="555">bicolor</emphasis>
</taxonomicName>
bellies usually present no markings whatsoever (but see discussion on
<taxonomicName baseAuthorityName="Guerin-Meneville" baseAuthorityYear="1838" box="[182,295,933,955]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="bicolor">
<emphasis box="[182,200,934,955]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[215,295,933,954]" italics="true" pageId="10" pageNumber="555">bicolor</emphasis>
</taxonomicName>
below). Given the historical use of the ventral pattern in the taxonomy of
<taxonomicName authorityName="Parker" authorityYear="1927" box="[557,715,964,985]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[557,715,964,985]" italics="true" pageId="10" pageNumber="555">Elachistocleis</emphasis>
</taxonomicName>
, we attempted to verify as many specimens as possible for this trait. We have not extensively checked our vouchers for other morphological diagnostic traits (male throat colour; presence and shape of the femoral stripe; post-commissural gland). Nevertheless, our preliminary data suggest that none of the above-cited morphological traits is as reliable as suggested in the literature [see also comments in
<bibRefCitation author="Nunes I &amp; Canedo C &amp; Carvalho RR" box="[536,761,1209,1231]" pageId="10" pageNumber="555" pagination="30 - 34" refId="ref17712" refString="Nunes I, Canedo C, Carvalho RR. 2010. Advertisement call and geographic distribution of Elachistocleis piauiensis Caramaschi &amp; Jim, 1983 (Amphibia, Microhylidae), with notes on the presence of post-commissural gland in the genus. South American Journal of Herpetology 5: 30 - 34." type="journal article" year="2010">
Nunes
<emphasis box="[620,679,1209,1231]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
(2010)
</bibRefCitation>
and
<bibRefCitation author="Marinho P &amp; Carvalho TR &amp; Bang DL &amp; Teixeira BFDV &amp; Azarak PA &amp; Campos CEC &amp; Giaretta AA" box="[195,434,1240,1262]" pageId="10" pageNumber="555" pagination="357 - 375" refId="ref17282" refString="Marinho P, Carvalho TR, Bang DL, Teixeira BFDV, Azarak PA, Campos CEC, Giaretta AA. 2018. Advertisement calls, intraspecific variation and species diagnosis of six Brazilian species of Elachistocleis (Anura: Microhylidae: Gastrophryninae). Zootaxa 4521: 357 - 375." type="journal article" year="2018">
Marinho
<emphasis box="[300,356,1240,1261]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
(2018)
</bibRefCitation>
].
</paragraph>
<paragraph blockId="10.[145,762,197,1905]" lastBlockId="10.[809,1425,197,495]" pageId="10" pageNumber="555">
Data from other sources are scarce. Tadpoles of few species are described (e.g.
<bibRefCitation author="Rossa-Feres D de &amp; Nomura F" pageId="10" pageNumber="555" pagination="1 - 26" refId="ref18621" refString="Rossa-Feres D de C, Nomura F. 2006. Characterization and taxonomic key for tadpoles (Amphibia: Anura) from the northwestern region of Sao Paulo State, Brazil. Biota Neotropica 6: 1 - 26." type="journal article" year="2006">Rossa-Feres &amp; Nomura, 2006</bibRefCitation>
;
<bibRefCitation author="Pereyra LC &amp; Akmentins MS &amp; Laufer G &amp; Vaira M" box="[211,428,1332,1353]" pageId="10" pageNumber="555" pagination="525 - 544" refId="ref18066" refString="Pereyra LC, Akmentins MS, Laufer G, Vaira M. 2013. A new species of Elachistocleis (Anura: Microhylidae) from north-western Argentina. Zootaxa 3694: 525 - 544." type="journal article" year="2013">
Pereyra
<emphasis box="[306,361,1332,1353]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2013
</bibRefCitation>
;
<bibRefCitation author="Schulze A &amp; Jansen M &amp; Kohler G" box="[439,655,1331,1354]" pageId="10" pageNumber="555" pagination="1 - 111" refId="ref18994" refString="Schulze A, Jansen M, Kohler G. 2015. Tadpole diversity of Bolivia's lowland anuran communities: molecular identification, morphological characterisation, and ecological assignment. Zootaxa 4016: 1 - 111." type="journal article" year="2015">
Schulze
<emphasis box="[534,589,1332,1353]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2015
</bibRefCitation>
;
<bibRefCitation author="Ferreira JS &amp; Weber LN" pageId="10" pageNumber="555" pagination="472 - 484" refId="ref15532" refString="Ferreira JS, Weber LN. 2021. A survey of the external morphology, internal oral morphology, chondrocranium and hyobranchial apparatus of Elachistocleis larvae Parker, 1927 (Anura, Microhylidae). Journal of Morphology 282: 472 - 484." type="journal article" year="2021">Ferreira &amp; Weber, 2021</bibRefCitation>
), but it is difficult to confidently assign these descriptions to lineages recovered because most lack an association with available DNA sequences. Moreover, there is no comprehensive understanding of variation, although
<bibRefCitation author="Gomez VI &amp; Kehr AI" box="[363,599,1485,1507]" pageId="10" pageNumber="555" pagination="1001 - 1006" refId="ref16050" refString="Gomez VI, Kehr AI. 2012. The effect of chemical signal of predatory fish and water bug on the morphology and development of Elachistocleis bicolor tadpoles (Anura: Microhylidae). Biologia 67: 1001 - 1006." type="journal article" year="2012">Gómez &amp; Kehr (2012)</bibRefCitation>
reported some variability in larval morphology related to chemical cues in the presence of predators. Advertisement calls of several
<taxonomicName authorityName="Parker" authorityYear="1927" box="[274,441,1577,1598]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[274,441,1577,1598]" italics="true" pageId="10" pageNumber="555">Elachistocleis</emphasis>
</taxonomicName>
have been described (e.g.
<bibRefCitation author="Nelson CE" box="[145,296,1608,1630]" pageId="10" pageNumber="555" pagination="895 - 898" refId="ref17614" refString="Nelson CE. 1972. Distribution and biology of Chiasmocleis panamensis (Amphibia: Microhylidae). Copeia 1972: 895 - 898." type="journal article" year="1972">Nelson, 1972</bibRefCitation>
;
<bibRefCitation author="Duellman WE" box="[311,497,1608,1630]" pageId="10" pageNumber="555" pagination="1 - 52" refId="ref15194" refString="Duellman WE. 1997. Amphibians of La Escalera region, southeastern Venezuela: taxonomy, ecology, and biogeography. Scientific Papers Natural History Museum the UniVersity of Kansas 2: 1 - 52." type="journal article" year="1997">Duellman, 1997</bibRefCitation>
;
<bibRefCitation author="Kwet A &amp; Di-Bernardo M" pageId="10" pageNumber="555" pagination="7 - 18" refId="ref17074" refString="Kwet A, Di-Bernardo M. 1998. Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and EnVironment 33: 7 - 18." type="journal article" year="1998">Kwet &amp; Di-Bernardo, 1998</bibRefCitation>
;
<bibRefCitation author="Lavilla EO &amp; Vaira M &amp; Ferrari L" box="[216,436,1638,1660]" pageId="10" pageNumber="555" pagination="269 - 284" refId="ref17160" refString="Lavilla EO, Vaira M, Ferrari L. 2003. A new species of Elachistocleis (Anura: Microhylidae) from the Andean Yungas of Argentina, with comments on the Elachistocleis oValis - E. bicolor controversy. Amphibia-Reptilia 24: 269 - 284." type="journal article" year="2003">
Lavilla
<emphasis box="[307,366,1638,1660]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2003
</bibRefCitation>
;
<bibRefCitation author="Nunes I &amp; Canedo C &amp; Carvalho RR" box="[451,665,1638,1660]" pageId="10" pageNumber="555" pagination="30 - 34" refId="ref17712" refString="Nunes I, Canedo C, Carvalho RR. 2010. Advertisement call and geographic distribution of Elachistocleis piauiensis Caramaschi &amp; Jim, 1983 (Amphibia, Microhylidae), with notes on the presence of post-commissural gland in the genus. South American Journal of Herpetology 5: 30 - 34." type="journal article" year="2010">
Nunes
<emphasis box="[535,594,1638,1660]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2010
</bibRefCitation>
; Toledo, 2010;
<bibRefCitation author="Marinho P &amp; Carvalho TR &amp; Bang DL &amp; Teixeira BFDV &amp; Azarak PA &amp; Campos CEC &amp; Giaretta AA" box="[219,467,1669,1691]" pageId="10" pageNumber="555" pagination="357 - 375" refId="ref17282" refString="Marinho P, Carvalho TR, Bang DL, Teixeira BFDV, Azarak PA, Campos CEC, Giaretta AA. 2018. Advertisement calls, intraspecific variation and species diagnosis of six Brazilian species of Elachistocleis (Anura: Microhylidae: Gastrophryninae). Zootaxa 4521: 357 - 375." type="journal article" year="2018">
Marinho
<emphasis box="[332,393,1669,1691]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2018
</bibRefCitation>
;
<bibRefCitation author="Pansonato A &amp; Mudrek JR &amp; Nunes CA &amp; Strussmann C" box="[484,754,1669,1691]" pageId="10" pageNumber="555" pagination="92 - 96" refId="ref17820" refString="Pansonato A, Mudrek JR, Nunes CA, Strussmann C. 2018. Advertisement calls of topotypes of Elachistocleis matogrosso (Anura: Microhylidae). Salamandra 54: 92 - 96." type="journal article" year="2018">
Pansonato
<emphasis box="[619,680,1669,1691]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2018
</bibRefCitation>
;
<bibRefCitation author="Jowers MJ &amp; Othman SN &amp; Borzee A &amp; Rivas GA &amp; Ramirez S &amp; Auguste RJ &amp; Downie JR &amp; Read M &amp; Murphy JC" box="[145,355,1700,1721]" pageId="10" pageNumber="555" pagination="189 - 206" refId="ref16761" refString="Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity &amp; EVolution 21: 189 - 206." type="journal article" year="2021">
Jowers
<emphasis box="[231,287,1700,1721]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2021
</bibRefCitation>
). Some of these are associated with vouchers present in our phylogeny (see:
<bibRefCitation author="Marinho P &amp; Carvalho TR &amp; Bang DL &amp; Teixeira BFDV &amp; Azarak PA &amp; Campos CEC &amp; Giaretta AA" pageId="10" pageNumber="555" pagination="357 - 375" refId="ref17282" refString="Marinho P, Carvalho TR, Bang DL, Teixeira BFDV, Azarak PA, Campos CEC, Giaretta AA. 2018. Advertisement calls, intraspecific variation and species diagnosis of six Brazilian species of Elachistocleis (Anura: Microhylidae: Gastrophryninae). Zootaxa 4521: 357 - 375." type="journal article" year="2018">
Marinho
<emphasis box="[700,755,1730,1752]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2018
</bibRefCitation>
;
<bibRefCitation author="Jowers MJ &amp; Othman SN &amp; Borzee A &amp; Rivas GA &amp; Ramirez S &amp; Auguste RJ &amp; Downie JR &amp; Read M &amp; Murphy JC" box="[216,436,1761,1783]" pageId="10" pageNumber="555" pagination="189 - 206" refId="ref16761" refString="Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity &amp; EVolution 21: 189 - 206." type="journal article" year="2021">
Jowers
<emphasis box="[306,366,1761,1783]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2021
</bibRefCitation>
). These are not adequately distributed in our topology and most are described from a few specimens and localities. Published data have already shown that even slight increases in sample size may affect recorded variation (see:
<bibRefCitation author="Marinho P &amp; Carvalho TR &amp; Bang DL &amp; Teixeira BFDV &amp; Azarak PA &amp; Campos CEC &amp; Giaretta AA" box="[594,866,197,1905]" pageId="10" pageNumber="555" pagination="357 - 375" refId="ref17282" refString="Marinho P, Carvalho TR, Bang DL, Teixeira BFDV, Azarak PA, Campos CEC, Giaretta AA. 2018. Advertisement calls, intraspecific variation and species diagnosis of six Brazilian species of Elachistocleis (Anura: Microhylidae: Gastrophryninae). Zootaxa 4521: 357 - 375." type="journal article" year="2018">
Marinho
<emphasis box="[699,755,1883,1905]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2018
</bibRefCitation>
). Finally, some localities present at least two sympatric species of
<taxonomicName authorityName="Parker" authorityYear="1927" box="[1048,1205,227,248]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1048,1205,227,248]" italics="true" pageId="10" pageNumber="555">Elachistocleis</emphasis>
</taxonomicName>
. For instance, both
<taxonomicName box="[809,985,258,280]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="nigrogularis">
<emphasis box="[809,827,259,280]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[841,985,258,279]" italics="true" pageId="10" pageNumber="555">nigrogularis</emphasis>
</taxonomicName>
and
<taxonomicName baseAuthorityName="Daudin" baseAuthorityYear="1802" box="[1044,1232,259,280]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="surinamensis">
<emphasis box="[1044,1062,259,280]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[1076,1232,259,280]" italics="true" pageId="10" pageNumber="555">surinamensis</emphasis>
</taxonomicName>
are syntopically found at the
<typeStatus box="[963,1014,289,310]" pageId="10" pageNumber="555">type</typeStatus>
locality of
<taxonomicName authority="(Jowers et al., 2021)" baseAuthorityName="Jowers" baseAuthorityYear="2021" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="nigrogularis">
<emphasis box="[1147,1165,290,311]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[1180,1326,289,310]" italics="true" pageId="10" pageNumber="555">nigrogularis</emphasis>
(
<bibRefCitation author="Jowers MJ &amp; Othman SN &amp; Borzee A &amp; Rivas GA &amp; Ramirez S &amp; Auguste RJ &amp; Downie JR &amp; Read M &amp; Murphy JC" pageId="10" pageNumber="555" pagination="189 - 206" refId="ref16761" refString="Jowers MJ, Othman SN, Borzee A, Rivas GA, Sanchez- Ramirez S, Auguste RJ, Downie JR, Read M, Murphy JC. 2021. Unraveling unique island colonization events in Elachistocleis frogs: phylogeography, cryptic divergence, and taxonomical implications. Organisms DiVersity &amp; EVolution 21: 189 - 206." type="journal article" year="2021">
Jowers
<emphasis box="[809,867,319,341]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2021
</bibRefCitation>
)
</taxonomicName>
, and both
<taxonomicName baseAuthorityName="Guerin-Meneville" baseAuthorityYear="1838" box="[1069,1180,319,341]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="bicolor">
<emphasis box="[1069,1087,320,341]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[1102,1180,319,340]" italics="true" pageId="10" pageNumber="555">bicolor</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Kwet &amp; Di-Bernardo" authorityYear="1998" box="[1239,1424,319,341]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="erythrogaster">
<emphasis box="[1239,1257,320,341]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[1271,1424,319,340]" italics="true" pageId="10" pageNumber="555">erythrogaster</emphasis>
</taxonomicName>
are found at the
<typeStatus box="[997,1046,351,372]" pageId="10" pageNumber="555">type</typeStatus>
locality of
<taxonomicName authority="(Kwet &amp; Di-Bernardo, 1998)" baseAuthorityName="Kwet &amp; Di-Bernardo" baseAuthorityYear="1998" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="species" species="erythrogaster">
<emphasis box="[1169,1187,351,372]" italics="true" pageId="10" pageNumber="555">E</emphasis>
.
<emphasis box="[1200,1350,350,371]" italics="true" pageId="10" pageNumber="555">erythrogaster</emphasis>
(
<bibRefCitation author="Kwet A &amp; Di-Bernardo M" pageId="10" pageNumber="555" pagination="7 - 18" refId="ref17074" refString="Kwet A, Di-Bernardo M. 1998. Elachistocleis erythrogaster, a new microhylid species from Rio Grande do Sul, Brazil. Studies on Neotropical Fauna and EnVironment 33: 7 - 18." type="journal article" year="1998">Kwet &amp; Di-Bernardo, 1998</bibRefCitation>
)
</taxonomicName>
(also see
<figureCitation box="[1193,1261,381,403]" captionStart="Figure 3" captionStartId="11.[163,243,1675,1697]" captionTargetBox="[297,1305,198,1632]" captionTargetId="figure-26@11.[298,1308,195,1635]" captionTargetPageId="11" captionText="Figure 3. Geographic location of sampled specimens (symbols) and type-localities of the nominal species (numbers). 1, Elachistocleis araios; 2, E. bicolor (here illustrated as the city of Buenos Aires; see Discussion); 3, E. bumbameuboi; 4, E. carƲalhoi; 5, E. cesarii; 6, E. corumbaensis; 7, E. erythrogaster; 8, E. haroi; 9, E. helianneae; 10, E. magna; 11, E. matogrosso; 12, E. muiraquitan; 13, E. nigrogularis; 14, E. panamensis; 15, E. pearsei; 16, E. piauiensis; 17, E. sikuani; 18, E. skotogaster; 19, E. surinamensis; 20, E. surumu; 21, E. tinigua." figureDoi="http://doi.org/10.5281/zenodo.7695491" httpUri="https://zenodo.org/record/7695491/files/figure.png" pageId="10" pageNumber="555">Fig. 3</figureCitation>
). This makes it even more difficult to associate larvae with adults and advertisement calls of unvouchered specimens to named species without molecular data.
</paragraph>
<paragraph blockId="10.[809,933,551,575]" box="[809,933,551,575]" pageId="10" pageNumber="555">
<heading box="[809,933,551,575]" centered="true" fontSize="9" level="2" pageId="10" pageNumber="555" reason="2">
<taxonomicName box="[809,933,551,575]" pageId="10" pageNumber="555" rank="subSpecies" species="fine" subSpecies="line">
<emphasis box="[809,933,551,575]" italics="true" pageId="10" pageNumber="555">A fine line</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph blockId="10.[809,1426,592,1656]" pageId="10" pageNumber="555">
A white or light yellow mid-dorsal line is present in many individuals of
<taxonomicName authorityName="Parker" authorityYear="1927" box="[1039,1194,622,643]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1039,1194,622,643]" italics="true" pageId="10" pageNumber="555">Elachistocleis</emphasis>
</taxonomicName>
, and other genera of Gastrophryninae, such as
<taxonomicName authority="Mehely, 1904" authorityName="Mehely" authorityYear="1904" box="[1114,1424,653,675]" class="Amphibia" family="Microhylidae" genus="Chiasmocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1114,1266,653,674]" italics="true" pageId="10" pageNumber="555">Chiasmocleis</emphasis>
Méhelÿ, 1904
</taxonomicName>
(e.g.
<bibRefCitation author="Peloso PLV &amp; Sturaro MJ &amp; Forlani MC &amp; Gaucher P &amp; Motta AP &amp; Wheeler WC" box="[862,1067,684,706]" pageId="10" pageNumber="555" pagination="1 - 112" refId="ref17941" refString="Peloso PLV, Sturaro MJ, Forlani MC, Gaucher P, Motta AP, Wheeler WC. 2014. Phylogeny, taxonomic revision, and character evolution of the genera Chiasmocleis and Syncope (Anura, Microhylidae) in Amazonia, with descriptions of three new species. Bulletin of the American Museum of Natural History 386: 1 - 112." type="journal article" year="2014">
Peloso
<emphasis box="[942,999,684,705]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2014
</bibRefCitation>
),
<taxonomicName authority="Mocquard, 1904" authorityName="Mocquard" authorityYear="1904" box="[1089,1425,684,706]" class="Amphibia" family="Microhylidae" genus="Ctenophryne" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1089,1234,684,705]" italics="true" pageId="10" pageNumber="555">Ctenophryne</emphasis>
Mocquard, 1904
</taxonomicName>
(e.g.
<bibRefCitation author="Duellman WE" box="[865,1056,714,736]" pageId="10" pageNumber="555" refId="ref15230" refString="Duellman WE. 1978. The Biology of an Equatorial Herpetofauna in Amazonian Ecuador. Lawrence: University of Kansas." type="book" year="1978">Duellman, 1978</bibRefCitation>
),
<taxonomicName authorityName="Carvalho" authorityYear="1954" box="[1077,1253,714,735]" class="Amphibia" family="Microhylidae" genus="Hamptophryne" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1077,1253,714,735]" italics="true" pageId="10" pageNumber="555">Hamptophryne</emphasis>
</taxonomicName>
A.L.
<bibRefCitation author="de Carvalho AL" pageId="10" pageNumber="555" pagination="1 - 22" refId="ref14789" refString="de Carvalho AL. 1954. A preliminary synopsis of the genera of American microhylid frogs. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 555: 1 - 22." type="journal article" year="1954">Carvalho, 1954</bibRefCitation>
(e.g.
<bibRefCitation author="Parker HW" box="[924,1068,745,767]" pageId="10" pageNumber="555" pagination="1 - 6" refId="ref17854" refString="Parker HW. 1927. The brevicipitid frogs allied to the genus Gastrophryne. Occasional Papers of the Museum of Zoology. UniVersity of Michigan 187: 1 - 6." type="journal article" year="1927">Parker, 1927</bibRefCitation>
;
<bibRefCitation author="Duellman WE" box="[1082,1263,745,767]" pageId="10" pageNumber="555" refId="ref15230" refString="Duellman WE. 1978. The Biology of an Equatorial Herpetofauna in Amazonian Ecuador. Lawrence: University of Kansas." type="book" year="1978">Duellman, 1978</bibRefCitation>
),
<taxonomicName authorityName="Keferstein" authorityYear="1867" box="[1285,1425,745,766]" class="Amphibia" family="Microhylidae" genus="Hypopachus" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1285,1425,745,766]" italics="true" pageId="10" pageNumber="555">Hypopachus</emphasis>
</taxonomicName>
[e.g.
<bibRefCitation author="Cope ED" box="[869,1015,776,797]" pageId="10" pageNumber="555" pagination="5 - 525" refId="ref15012" refString="Cope ED. 1889. Batrachia of North America. Bulletin of the United States National Museum 34: 5 - 525." type="journal article" year="1889">Cope (1889)</bibRefCitation>
; see also figure
<quantity box="[1224,1272,776,798]" metricMagnitude="-1" metricUnit="m" metricValue="1.27" pageId="10" pageNumber="555" unit="in" value="5.0">5 in</quantity>
<bibRefCitation author="Greenbaum E &amp; Smith EN &amp; de Sa RO" pageId="10" pageNumber="555" pagination="265 - 277" refId="ref16211" refString="Greenbaum E, Smith EN, de Sa RO. 2011. Molecular systematics of the Middle American genus Hypopachus (Anura: Microhylidae). Molecular Phylogenetics and EVolution 61: 265 - 277." type="journal article" year="2011">
Greenbaum
<emphasis box="[809,861,806,828]" italics="true" pageId="10" pageNumber="555">et al</emphasis>
. (2011)
</bibRefCitation>
],
<taxonomicName authority="Miranda-Ribeiro, 1924" authorityName="Miranda-Ribeiro" authorityYear="1924" box="[977,1371,806,828]" class="Amphibia" family="Microhylidae" genus="Dasypops" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[977,1090,807,828]" italics="true" pageId="10" pageNumber="555">Dasypops</emphasis>
Miranda-Ribeiro, 1924
</taxonomicName>
and
<taxonomicName authority="Cope, 1870" authorityName="Cope" authorityYear="1870" box="[809,1101,837,858]" class="Amphibia" family="Microhylidae" genus="Stereocyclops" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[809,965,837,858]" italics="true" pageId="10" pageNumber="555">Stereocyclops</emphasis>
Cope, 1870
</taxonomicName>
(PP, personal observation). Many authors have noticed important intraspecific or intrapopulational variation (i.e. being present in some individuals and absent in others) in this character in other genera (e.g.
<taxonomicName authority=", Peloso et al., 2014" authorityName="Peloso" authorityYear="2014" box="[1028,1418,960,982]" class="Amphibia" family="Microhylidae" genus="Chiasmocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1028,1185,960,981]" italics="true" pageId="10" pageNumber="555">Chiasmocleis</emphasis>
,
<bibRefCitation author="Peloso PLV &amp; Sturaro MJ &amp; Forlani MC &amp; Gaucher P &amp; Motta AP &amp; Wheeler WC" box="[1201,1418,960,982]" pageId="10" pageNumber="555" pagination="1 - 112" refId="ref17941" refString="Peloso PLV, Sturaro MJ, Forlani MC, Gaucher P, Motta AP, Wheeler WC. 2014. Phylogeny, taxonomic revision, and character evolution of the genera Chiasmocleis and Syncope (Anura, Microhylidae) in Amazonia, with descriptions of three new species. Bulletin of the American Museum of Natural History 386: 1 - 112." type="journal article" year="2014">
Peloso
<emphasis box="[1285,1346,960,981]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2014
</bibRefCitation>
</taxonomicName>
;
<taxonomicName authority=", Zweifel &amp; Myers, 1989" authorityName="Zweifel &amp; Myers" authorityYear="1989" box="[809,1245,990,1012]" class="Amphibia" family="Microhylidae" genus="Ctenophryne" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[809,959,990,1011]" italics="true" pageId="10" pageNumber="555">Ctenophryne</emphasis>
,
<bibRefCitation author="Zweifel RG &amp; Myers CW" box="[974,1245,990,1012]" pageId="10" pageNumber="555" pagination="1 - 16" refId="ref20087" refString="Zweifel RG, Myers CW. 1989. A New Frog of the Genus Ctenophryne (Microhylidae) from the Pacific Lowlands of Northwestern South America. American Museum NoVitates 2947: 1 - 16." type="journal article" year="1989">Zweifel &amp; Myers, 1989</bibRefCitation>
</taxonomicName>
;
<taxonomicName authority=", GNF" authorityName="GNF" class="Amphibia" family="Microhylidae" genus="Stereocyclops" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1260,1418,990,1011]" italics="true" pageId="10" pageNumber="555">Stereocyclops</emphasis>
, GNF
</taxonomicName>
, personal observation) and in
<taxonomicName authorityName="Parker" authorityYear="1927" box="[1213,1370,1021,1042]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1213,1370,1021,1042]" italics="true" pageId="10" pageNumber="555">Elachistocleis</emphasis>
</taxonomicName>
(e.g.
<bibRefCitation author="Nelson CE" box="[809,954,1052,1074]" pageId="10" pageNumber="555" pagination="895 - 898" refId="ref17614" refString="Nelson CE. 1972. Distribution and biology of Chiasmocleis panamensis (Amphibia: Microhylidae). Copeia 1972: 895 - 898." type="journal article" year="1972">Nelson, 1972</bibRefCitation>
; Toledo, 2010;
<bibRefCitation author="Toledo LF &amp; Loebmann D &amp; Haddad CFB" box="[1115,1315,1052,1074]" pageId="10" pageNumber="555" pagination="50 - 60" refId="ref19506" refString="Toledo LF, Loebmann D, Haddad CFB. 2010. Revalidation and redescription of Elachistocleis cesarii (Miranda-Ribeiro, 1920) (Anura: Microhylidae). Zootaxa 2418: 50 - 60." type="journal article" year="2010">
Toledo
<emphasis box="[1194,1249,1052,1073]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2010
</bibRefCitation>
;
<bibRefCitation author="Marinho P &amp; Carvalho TR &amp; Bang DL &amp; Teixeira BFDV &amp; Azarak PA &amp; Campos CEC &amp; Giaretta AA" pageId="10" pageNumber="555" pagination="357 - 375" refId="ref17282" refString="Marinho P, Carvalho TR, Bang DL, Teixeira BFDV, Azarak PA, Campos CEC, Giaretta AA. 2018. Advertisement calls, intraspecific variation and species diagnosis of six Brazilian species of Elachistocleis (Anura: Microhylidae: Gastrophryninae). Zootaxa 4521: 357 - 375." type="journal article" year="2018">
Marinho
<emphasis box="[809,867,1082,1104]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2018
</bibRefCitation>
). Notwithstanding, the occurrence of this line and the variation of its extension when present (e.g. from snout to vent or from post-cephalic fold to vent) were used in diagnoses of several
<taxonomicName pageId="10" pageNumber="555">
<emphasis box="[1268,1425,1174,1195]" italics="true" pageId="10" pageNumber="555">Elachistocleis</emphasis>
species
</taxonomicName>
(e.g.
<bibRefCitation author="Caramaschi U &amp; Jim J" box="[960,1259,1205,1227]" pageId="10" pageNumber="555" pagination="390 - 394" refId="ref14682" refString="Caramaschi U, Jim J. 1983. A new microhylid frog, genus Elachistocleis (Amphibia, Anura), from northeastern Brasil. Herpetologica 39: 390 - 394." type="journal article" year="1983">Caramaschi &amp; Jim, 1983</bibRefCitation>
;
<bibRefCitation author="Caramaschi U" pageId="10" pageNumber="555" pagination="1 - 30" refId="ref14635" refString="Caramaschi U. 2010. Notes on the taxonomic status of Elachistocleis oValis (Schneider, 1799) and description of five new species of Elachistocleis Parker, 1927 (Amphibia, Anura, Microhylidae). Boletim do Museu Nacional 527: 1 - 30." type="journal article" year="2010">Caramaschi, 2010</bibRefCitation>
;
<bibRefCitation author="Nunes-de-Almeida CHL &amp; Toledo LF" box="[880,1283,1236,1258]" pageId="10" pageNumber="555" pagination="43 - 50" refId="ref17765" refString="Nunes-de-Almeida CHL, Toledo LF. 2012. A new species of Elachistocleis Parker (Anura, Microhylidae) from the state of Acre, northern Brazil. Zootaxa 3424: 43 - 50." type="journal article" year="2012">Nunes-de-Almeida &amp; Toledo, 2012</bibRefCitation>
;
<bibRefCitation author="Piva A &amp; Caramaschi U &amp; Albuquerque NR" pageId="10" pageNumber="555" pagination="143 - 154" refId="ref18209" refString="Piva A, Caramaschi U, Albuquerque NR. 2017. A new species of Elachistocleis (Anura: Microhylidae) from the Brazilian Pantanal. Phyllomedusa. Journal of Herpetology 16: 143 - 154." type="journal article" year="2017">
Piva
<emphasis box="[1359,1418,1236,1257]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2017
</bibRefCitation>
).
</paragraph>
<paragraph blockId="10.[809,1426,592,1656]" pageId="10" pageNumber="555">
We found a remarkable variation in the occurrence of the mid-dorsal white line in and among species (
<figureCitation captionStart="Figure 1" captionStartId="5.[163,244,1810,1832]" captionTargetBox="[226,1379,195,1770]" captionTargetId="graphics-25@5.[226,1378,239,1770]" captionTargetPageId="5" captionText="Figure 1. Species delimitation in Elachistocleis. The tree shown is from the maximum likelihood (ML) analysis with only 16S + COI from only Elachistocleis samples (tree rooted in E. araios). Numbers on branches are SH-aLRT/UFBoot support values. Asterisks represent support above 80/95. The terminals of the tree are named as: original labelling of" figureDoi="http://doi.org/10.5281/zenodo.7695483" httpUri="https://zenodo.org/record/7695483/files/figure.png" pageId="10" pageNumber="555">Fig. 1</figureCitation>
), similar to the variation reported for
<taxonomicName authority="(Peloso et al., 2014)" baseAuthorityName="Peloso" baseAuthorityYear="2014" class="Amphibia" family="Microhylidae" genus="Chiasmocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1272,1425,1358,1379]" italics="true" pageId="10" pageNumber="555">Chiasmocleis</emphasis>
(
<bibRefCitation author="Peloso PLV &amp; Sturaro MJ &amp; Forlani MC &amp; Gaucher P &amp; Motta AP &amp; Wheeler WC" box="[818,1039,1389,1411]" pageId="10" pageNumber="555" pagination="1 - 112" refId="ref17941" refString="Peloso PLV, Sturaro MJ, Forlani MC, Gaucher P, Motta AP, Wheeler WC. 2014. Phylogeny, taxonomic revision, and character evolution of the genera Chiasmocleis and Syncope (Anura, Microhylidae) in Amazonia, with descriptions of three new species. Bulletin of the American Museum of Natural History 386: 1 - 112." type="journal article" year="2014">
Peloso
<emphasis box="[904,965,1389,1411]" italics="true" pageId="10" pageNumber="555">et al.</emphasis>
, 2014
</bibRefCitation>
)
</taxonomicName>
. This variation challenges the reliability of the trait as a diagnostic feature and weakens the known diagnoses of several currently recognized species (see below). It was impossible to confidently evaluate the extension of the mid-dorsal line, thus we only scored the presence or the absence of the line. Hence, we recommend caution when using the mid-dorsal line in the systematics of
<taxonomicName authorityName="Parker" authorityYear="1927" box="[1264,1419,1604,1625]" class="Amphibia" family="Microhylidae" genus="Elachistocleis" kingdom="Animalia" order="Anura" pageId="10" pageNumber="555" phylum="Chordata" rank="genus">
<emphasis box="[1264,1419,1604,1625]" italics="true" pageId="10" pageNumber="555">Elachistocleis</emphasis>
</taxonomicName>
, especially for diagnostic purposes.
</paragraph>
</subSubSection>
</treatment>
</document>