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<document ID-DOI="10.1038/s41598-020-61973-5" ID-GBIF-Dataset="b42c7378-ff6d-40fc-98da-c1f2ea5649a3" ID-PMC="PMC7181706" ID-PubMed="32327670" ID-Zenodo-Dep="3766727" checkinTime="1587901428986" checkinUser="plazi" docAuthor="Mörs, Thomas, Reguero, Marcelo &amp; Vasilyan, Davit" docDate="2020" docId="03AF87E1EC5EFFFFFE52F92A6171F9E8" docLanguage="en" docName="srep.s41598-020-61973-5.pdf" docOrigin="Scientific Reports 10 (5051)" docStyle="DocumentStyle{}" docTitle="Calyptocephalella Mörs, Reguero &amp; Vasilyan 2020" docType="treatment" docVersion="6" lastPageNumber="7" masterDocId="FF96FF99EC5FFFF9FFEDFFBD641BFFD4" masterDocTitle="First fossil frog from Antarctica: implications for Eocene high latitude climate conditions and Gondwanan cosmopolitanism of Australobatrachia" masterLastPageNumber="11" masterPageNumber="1" pageNumber="2" updateTime="1668144403742" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>First fossil frog from Antarctica: implications for Eocene high latitude climate conditions and Gondwanan cosmopolitanism of Australobatrachia</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Mörs, Thomas</mods:namePart>
<mods:affiliation>Department of Palaeobiology, Swedish Museum of Natural History, P. O. Box 50007, SE- 104 05, Stockholm, Sweden</mods:affiliation>
<mods:nameIdentifier type="email">thomas.moers@nrm.se</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Reguero, Marcelo</mods:namePart>
<mods:affiliation>Instituto Antártico Argentino, Campus Miguelete, 25 de Mayo 1151, 3 ° piso B 1650 HMK, San Martín, Buenos Aires, Argentina</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Vasilyan, Davit</mods:namePart>
<mods:affiliation>Department of Geosciences, University of Fribourg, Chemin du musée 6, 1700, Fribourg, Switzerland</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>Scientific Reports</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2020</mods:date>
<mods:detail type="pubDate">
<mods:number>2020-04-25</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>10</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>5051</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>1</mods:start>
<mods:end>11</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1038/s41598-020-61973-5</mods:identifier>
<mods:identifier type="GBIF-Dataset">b42c7378-ff6d-40fc-98da-c1f2ea5649a3</mods:identifier>
<mods:identifier type="PMC">PMC7181706</mods:identifier>
<mods:identifier type="PubMed">32327670</mods:identifier>
<mods:identifier type="Zenodo-Dep">3766727</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.3803362" ID-GBIF-Taxon="163743997" ID-Zenodo-Dep="3803362" LSID="urn:lsid:plazi:treatment:03AF87E1EC5EFFFFFE52F92A6171F9E8" httpUri="http://treatment.plazi.org/id/03AF87E1EC5EFFFFFE52F92A6171F9E8" lastPageId="6" lastPageNumber="7" pageId="1" pageNumber="2">
<subSubSection box="[447,643,1687,1708]" pageId="1" pageNumber="2" type="nomenclature">
<paragraph blockId="1.[415,1102,1522,1736]" box="[447,643,1687,1708]" pageId="1" pageNumber="2">
<taxonomicName authority="Mörs, Reguero &amp; Vasilyan, 2020" authorityName="Mörs, Reguero &amp; Vasilyan" authorityYear="2020" box="[447,643,1687,1708]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="1" pageNumber="2" phylum="Chordata" rank="species">
<emphasis box="[447,612,1687,1708]" italics="true" pageId="1" pageNumber="2">Calyptocephalella</emphasis>
sp.
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection box="[447,592,1714,1736]" pageId="1" pageNumber="2" type="description">
<paragraph blockId="1.[415,1102,1522,1736]" box="[447,592,1714,1736]" pageId="1" pageNumber="2">
<figureCitation box="[447,535,1714,1736]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="1" pageNumber="2">Figures 2</figureCitation>
and
<figureCitation box="[580,592,1714,1735]" captionStart="Figure 3" captionStartId="3.[415,480,746,768]" captionTargetBox="[418,1372,127,710]" captionTargetId="figure@3.[415,1375,124,711]" captionTargetPageId="3" captionText="Figure 3. Skull bone fragment (NRM-PZ B281) of Calyptocephalella sp. from Seymour Island, Antarctica in dorsal (a), ventral (b) and lateral (c) views." figureDoi="http://doi.org/10.5281/zenodo.3766733" httpUri="https://zenodo.org/record/3766733/files/figure.png" pageId="1" pageNumber="2">3</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="2" type="materials_examined">
<paragraph blockId="1.[415,652,1762,1814]" pageId="1" pageNumber="2">
<heading bold="true" box="[415,652,1762,1786]" centered="true" fontSize="10" level="2" pageId="1" pageNumber="2" reason="0">
<emphasis bold="true" box="[415,652,1762,1786]" pageId="1" pageNumber="2">Referred specimens.</emphasis>
</heading>
<materialsCitation ID-GBIF-Occurrence="2608704112" collectionCode="Swedish Museum of Natural History" latitude="-64.23278" location="IAA 2 / 95, Marsupial site, Seymour Island, Antarctic Peninsula" longLatPrecision="17" longitude="-56.65167" pageId="1" pageNumber="2" specimenCode="NRM-PZ B282, NRM-PZ B281" specimenCount="1">
<collectionCode box="[678,1033,1765,1787]" pageId="1" pageNumber="2">Swedish Museum of Natural History</collectionCode>
<specimenCode box="[1038,1187,1765,1787]" pageId="1" pageNumber="2">NRM-PZ B282</specimenCode>
, right ilium (
<figureCitation box="[1313,1368,1765,1787]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="1" pageNumber="2">Fig. 2</figureCitation>
),
<specimenCode pageId="1" pageNumber="2">
NRM-PZ
<heading box="[415,466,1792,1813]" centered="true" fontSize="9" level="3" pageId="1" pageNumber="2" reason="4">B281</heading>
</specimenCode>
, skull bone (
<figureCitation box="[584,638,1792,1814]" captionStart="Figure 3" captionStartId="3.[415,480,746,768]" captionTargetBox="[418,1372,127,710]" captionTargetId="figure@3.[415,1375,124,711]" captionTargetPageId="3" captionText="Figure 3. Skull bone fragment (NRM-PZ B281) of Calyptocephalella sp. from Seymour Island, Antarctica in dorsal (a), ventral (b) and lateral (c) views." figureDoi="http://doi.org/10.5281/zenodo.3766733" httpUri="https://zenodo.org/record/3766733/files/figure.png" pageId="1" pageNumber="2">Fig. 3</figureCitation>
).
</materialsCitation>
</paragraph>
<paragraph blockId="1.[415,1479,1840,1919]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[415,717,1840,1864]" pageId="1" pageNumber="2">Locality, horizon and age.</emphasis>
<location LSID="urn:lsid:plazi:treatment:03AF87E1EC5EFFFFFE52F92A6171F9E8:8ED9602CEC5EFFF8FD05F8896152F89E" box="[744,1353,1843,1866]" latitude="-64.23278" longLatPrecision="17" longitude="-56.65167" name="IAA 2 / 95, Marsupial site, Seymour Island, Antarctic Peninsula" pageId="1" pageNumber="2">IAA 2/95, Marsupial site, Seymour Island, Antarctic Peninsula</location>
(
<geoCoordinate box="[1365,1475,1841,1866]" degrees="64" direction="south" minutes="13" orientation="latitude" pageId="1" pageNumber="2" precision="15" seconds="58" value="-64.23278">64°1358″S</geoCoordinate>
;
<geoCoordinate box="[415,541,1868,1892]" degrees="56" direction="west" minutes="39" orientation="longitude" pageId="1" pageNumber="2" precision="15" seconds="06" value="-56.65167">56°3906″W</geoCoordinate>
).
<location LSID="urn:lsid:plazi:treatment:03AF87E1EC5EFFFFFE52F92A6171F9E8:8ED9602CEC5EFFF8FDDEF8F36685F8AA" latitude="-64.23278" longLatPrecision="17" longitude="-56.65167" name="Natica horizon' within the Cucullaea I Allomember (Telm 5) of the La Meseta Formation, Bartonian (40 Ma), Eocene" pageId="1" pageNumber="2">
<taxonomicName box="[563,630,1870,1892]" class="Gastropoda" family="Naticidae" genus="Natica" kingdom="Animalia" order="Littorinimorpha" pageId="1" pageNumber="1" phylum="Mollusca" rank="genus">Natica</taxonomicName>
horizon within the
<taxonomicName authority="I Allomember" authorityName="I Allomember" box="[839,1089,1870,1892]" class="Bivalvia" family="Cucullaeidae" genus="Cucullaea" kingdom="Animalia" order="Arcoida" pageId="1" pageNumber="2" phylum="Mollusca" rank="genus">Cucullaea I Allomember</taxonomicName>
(Telm 5) of the La Meseta Formation, Bartonian (40Ma), Eocene
</location>
<superScript attach="left" box="[670,704,1894,1908]" fontSize="6" pageId="1" pageNumber="2">
<bibRefCitation author="Douglas, P. M. J." box="[670,685,1894,1908]" journalOrPublisher="P. Natl Acad. Sci. USA" pageId="1" pageNumber="2" pagination="6582 - 6587" part="111" refId="ref6586" refString="23. Douglas, P. M. J. et al. Pronounced zonal heterogeneity in Eocene southern high-latitude sea surface temperatures. P. Natl Acad. Sci. USA 111, 6582 - 6587 (2014)." title="Pronounced zonal heterogeneity in Eocene southern high-latitude sea surface temperatures" type="journal article" year="2014">23</bibRefCitation>
,
<bibRefCitation author="Amenabar, C. R. &amp; Montes, M. &amp; Nozal, F. &amp; Santillana, S." box="[689,704,1894,1908]" journalOrPublisher="Geological Magazine" pageId="1" pageNumber="2" pagination="1 - 16" refId="ref6627" refString="24. Amenabar, C. R., Montes, M., Nozal, F. &amp; Santillana, S. Dinoflagellate cysts of the La Meseta Formation (middle to late Eocene), Antarctic Peninsula: implications for biostratigraphy, palaeoceanography and palaeoenvironment. Geological Magazine, 1 - 16, https: // doi. org / 10.1017 / S 0016756819000591 (2019)." title="Dinoflagellate cysts of the La Meseta Formation (middle to late Eocene), Antarctic Peninsula: implications for biostratigraphy, palaeoceanography and palaeoenvironment" type="book chapter" year="2019">24</bibRefCitation>
</superScript>
(
<figureCitation box="[716,770,1897,1919]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="1" pageNumber="2">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3766731" ID-Zenodo-Dep="3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="2" pageNumber="3" startId="2.[415,480,1227,1249]" targetBox="[417,1469,126,1190]" targetPageId="2">
<paragraph blockId="2.[415,1470,1227,1435]" pageId="2" pageNumber="3">
<emphasis bold="true" box="[415,503,1227,1249]" pageId="2" pageNumber="3">Figure 2.</emphasis>
Ilium (NRM-PZ B282) of
<taxonomicName box="[763,928,1227,1248]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[763,928,1227,1248]" italics="true" pageId="2" pageNumber="3">Calyptocephalella</emphasis>
</taxonomicName>
sp. from Seymour Island, Antarctica. Ilium in lateral (
<emphasis bold="true" box="[1441,1452,1227,1248]" pageId="2" pageNumber="3">a</emphasis>
), medial (
<emphasis bold="true" box="[494,507,1254,1275]" pageId="2" pageNumber="3">b</emphasis>
), ventral (
<emphasis bold="true" box="[606,616,1254,1275]" pageId="2" pageNumber="3">c</emphasis>
) and dorsal (
<emphasis bold="true" box="[742,755,1254,1275]" pageId="2" pageNumber="3">d</emphasis>
) views. Magnified region of the dorsal protuberance in lateral (
<emphasis bold="true" box="[1354,1382,1254,1275]" pageId="2" pageNumber="3">e </emphasis>
dashed rectangle in orange color) and dorsal (
<emphasis bold="true" box="[778,802,1280,1302]" pageId="2" pageNumber="3">f </emphasis>
dashed rectangle in yellow color) view. The dashed line in black on (
<emphasis bold="true" box="[1451,1462,1280,1301]" pageId="2" pageNumber="3">a</emphasis>
) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (
<emphasis box="[1420,1432,1308,1329]" italics="true" pageId="2" pageNumber="3">*</emphasis>
) on (
<emphasis bold="true" box="[423,434,1334,1355]" pageId="2" pageNumber="3">a</emphasis>
,
<emphasis bold="true" box="[440,450,1334,1355]" pageId="2" pageNumber="3">c</emphasis>
) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (
<emphasis box="[1416,1440,1334,1355]" italics="true" pageId="2" pageNumber="3">**</emphasis>
) indicates the notch caudally from the dorsal protuberance. The dashed red lines on (
<emphasis bold="true" box="[1212,1222,1360,1381]" pageId="2" pageNumber="3">e</emphasis>
,
<emphasis bold="true" box="[1228,1236,1360,1381]" pageId="2" pageNumber="3">f</emphasis>
) outline the intact bone surface. Abbreviations: ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression.
</paragraph>
</caption>
<subSubSection lastPageId="5" lastPageNumber="6" pageId="2" pageNumber="3" type="description">
<paragraph blockId="2.[415,1479,1505,1610]" pageId="2" pageNumber="3">
<emphasis bold="true" box="[415,597,1505,1529]" pageId="2" pageNumber="3">Measurements.</emphasis>
The preserved part of the ilium measures 3.9 mm in length, the distance from the tip of the dorsal acetabular expansion to the (preserved) tip of the ventral acetabular expansion measures 3.3mm, the highest height of the acetabular fossa equals 2.5 mm. The skull bone measures 2.7 mm at its both broadest and longest parts.
</paragraph>
<paragraph blockId="2.[415,1480,1644,1962]" lastBlockId="3.[415,1480,1249,1968]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
<emphasis bold="true" box="[415,482,1644,1668]" pageId="2" pageNumber="3">Ilium.</emphasis>
The fragmentary right ilium (NRM-PZ B282) lacks the caudal portion of the acetabulum and most of the iliac shaft. The dorsal acetabular expansion has a smooth lateral surface and is higher than the preserved part of the ventral acetabular expansion (
<figureCitation box="[815,882,1700,1722]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="2" pageNumber="3">Fig. 2a</figureCitation>
). A large and deep supraacetabular fossa is present at its base (
<figureCitation box="[422,504,1727,1749]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="2" pageNumber="3">Fig. 2a,d</figureCitation>
). The preserved portion of the acetabulum is concave and its shape allows concluding a (semi-)circular outline. The acetabular rim is most prominent at its anterior part (
<figureCitation box="[1033,1112,1754,1776]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="2" pageNumber="3">Fig. 2a,c</figureCitation>
). The barely-developed ventral acetabular expansion projects ventrally. The posterior-most portion of the ventral acetabular expansion is broken off. However, the anterior portion of the ventral acetabular expansion is higher than the preserved posterior portion. In ventral view (
<figureCitation box="[568,647,1834,1856]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="2" pageNumber="3">Fig. 2a,c</figureCitation>
), the lateral surface of the ventral acetabular expansion is convex. The ventral acetabular expansion possesses a shallow and broad depression. In the preacetabular zone, a small and shallow preacetabular fossa is present (
<figureCitation box="[570,634,1887,1909]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="2" pageNumber="3">Fig. 2c</figureCitation>
). The preserved portion of the iliac shaft is damaged and precludes a confident statement whether the dorsal protuberance is present or absent. A narrow and shallow longitudinal groove is observable in the lateral surface of the iliac shaft, which probably corresponds to the posterior extension of the ventral depression (sensu
<bibRefCitation author="Muzzopappa, P. &amp; Baez, A. M." box="[587,602,1249,1263]" journalOrPublisher="Ameghiniana" pageId="3" pageNumber="4" pagination="113 - 125" part="46" refId="ref5842" refString="10. Muzzopappa, P. &amp; Baez, A. M. Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus. Ameghiniana 46, 113 - 125 (2009)." title="Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus" type="journal article" year="2009">
<superScript attach="left" box="[587,602,1249,1263]" fontSize="6" pageId="3" pageNumber="4">10</superScript>
</bibRefCitation>
) (
<figureCitation box="[623,689,1252,1274]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="3" pageNumber="4">Fig. 2a</figureCitation>
). However, intact parts of bone surface are preserved slightly ventral to the dorsal margin on both lateral and medial surfaces (
<figureCitation box="[840,917,1279,1301]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="3" pageNumber="4">Fig. 2e,f</figureCitation>
). The one on the lateral surface is a curved shallow groove and runs posteroventrally (
<figureCitation box="[673,737,1306,1328]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="3" pageNumber="4">Fig. 2e</figureCitation>
). This feature anteroventrally demarcates the slightly elevated roughened scar interpreted above as the dorsal protuberance. The area between the dorsal acetabular expansion and iliac shaft is slightly projected dorsally. This area corresponds to the position of the dorsal protuberance. In fact, no clear evidence of a dorsal protuberance can be found on the ilium, only a slightly roughened area with minimal elevation that corresponds to the dorsal protuberance and the scar for the insertion of the musculus gluteus magnus can be observed. At the caudal side of the dorsal protuberance, a distinct notch is visible (
<figureCitation box="[1198,1264,1439,1461]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="3" pageNumber="4">Fig. 2a</figureCitation>
) which we consider as a further evidence of our interpretation. The area corresponding to the dorsal protuberance is located anteriorly to the anterior margin of the acetabular rim. Medially, the entire surface opposing the acetabulum is lost and the area preserved more anteriorly is slightly convex medially and smooth. Anteriorly and dorsally, just adjacent to the anterior end of the dorsal protuberance a foramen is present (
<figureCitation box="[1032,1098,1546,1568]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="3" pageNumber="4">Fig. 2b</figureCitation>
).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3766733" ID-Zenodo-Dep="3766733" httpUri="https://zenodo.org/record/3766733/files/figure.png" pageId="3" pageNumber="4" startId="3.[415,480,746,768]" targetBox="[418,1372,127,710]" targetPageId="3">
<paragraph blockId="3.[415,1442,746,794]" pageId="3" pageNumber="4">
<emphasis bold="true" box="[415,503,746,768]" pageId="3" pageNumber="4">Figure 3.</emphasis>
Skull bone fragment (NRM-PZ B281) of
<taxonomicName box="[903,1068,746,767]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[903,1068,746,767]" italics="true" pageId="3" pageNumber="4">Calyptocephalella</emphasis>
</taxonomicName>
sp. from Seymour Island, Antarctica in dorsal (
<emphasis bold="true" box="[487,498,773,794]" pageId="3" pageNumber="4">a</emphasis>
), ventral (
<emphasis bold="true" box="[597,610,773,794]" pageId="3" pageNumber="4">b</emphasis>
) and lateral (
<emphasis bold="true" box="[737,747,773,794]" pageId="3" pageNumber="4">c</emphasis>
) views.
</paragraph>
</caption>
<paragraph pageId="3" pageNumber="4">
<table box="[423,1418,862,1034]" gridcols="7" gridrows="6" pageId="3" pageNumber="4">
<tr box="[423,1418,862,879]" gridrow="0" pageId="3" pageNumber="4">
<th box="[423,540,862,879]" gridcol="0" gridrow="0" pageId="3" pageNumber="4">
<emphasis bold="true" box="[423,478,862,879]" pageId="3" pageNumber="4">species</emphasis>
</th>
<th box="[557,737,862,879]" gridcol="1" gridrow="0" pageId="3" pageNumber="4">
<emphasis bold="true" box="[557,615,862,879]" pageId="3" pageNumber="4">locality</emphasis>
</th>
<th box="[757,934,862,879]" gridcol="2" gridrow="0" pageId="3" pageNumber="4">
<emphasis bold="true" box="[757,811,862,878]" pageId="3" pageNumber="4">coll nr.</emphasis>
</th>
<th box="[957,1057,862,879]" gridcol="3" gridrow="0" pageId="3" pageNumber="4">
<emphasis bold="true" box="[957,1057,862,879]" pageId="3" pageNumber="4">SVL (in mm)</emphasis>
</th>
<th box="[1080,1174,862,879]" gridcol="4" gridrow="0" pageId="3" pageNumber="4">
<emphasis bold="true" box="[1080,1174,862,879]" pageId="3" pageNumber="4">HT (in mm)</emphasis>
</th>
<th box="[1200,1286,862,879]" gridcol="5" gridrow="0" pageId="3" pageNumber="4">
<emphasis bold="true" box="[1200,1286,862,879]" pageId="3" pageNumber="4">RHS (in %)</emphasis>
</th>
<th box="[1306,1418,862,879]" gridcol="6" gridrow="0" pageId="3" pageNumber="4">
<emphasis bold="true" box="[1306,1382,862,879]" pageId="3" pageNumber="4">Reference</emphasis>
</th>
</tr>
<tr box="[423,1418,889,910]" gridrow="1" pageId="3" pageNumber="4">
<th box="[423,540,889,910]" gridcol="0" gridrow="1" pageId="3" pageNumber="4">
<taxonomicName box="[423,540,893,910]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="pichileufensis">
<emphasis box="[423,540,893,910]" italics="true" pageId="3" pageNumber="4">C. pichileufensis</emphasis>
</taxonomicName>
</th>
<td box="[557,737,889,910]" gridcol="1" gridrow="1" pageId="3" pageNumber="4">Río Pichileufú locality</td>
<td box="[757,934,889,910]" gridcol="2" gridrow="1" pageId="3" pageNumber="4">BAR 85b</td>
<td box="[957,1057,889,910]" gridcol="3" gridrow="1" pageId="3" pageNumber="4">107</td>
<td box="[1080,1174,889,910]" gridcol="4" gridrow="1" pageId="3" pageNumber="4">6</td>
<td box="[1200,1286,889,910]" gridcol="5" gridrow="1" pageId="3" pageNumber="4">5.6</td>
<td box="[1306,1418,889,910]" gridcol="6" gridrow="1" pageId="3" pageNumber="4">
<bibRefCitation author="Gomez, R. O. &amp; Baez, A. M. &amp; Muzzopappa, P." box="[1306,1318,889,901]" journalOrPublisher="J. Vertebr. Paleontol." pageId="3" pageNumber="4" pagination="50 - 59" part="31" refId="ref7642" refString="48. Gomez, R. O., Baez, A. M. &amp; Muzzopappa, P. A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina. J. Vertebr. Paleontol. 31, 50 - 59 (2011)." title="A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina" type="journal article" year="2011">48</bibRefCitation>
: Figs.2 and 4
</td>
</tr>
<tr box="[423,1418,920,941]" gridrow="2" pageId="3" pageNumber="4">
<th box="[423,540,920,941]" gridcol="0" gridrow="2" pageId="3" pageNumber="4">
<taxonomicName baseAuthorityName="Duméril &amp; Bibron" baseAuthorityYear="1841" box="[423,473,924,940]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="gayi">
<emphasis box="[423,473,924,940]" italics="true" pageId="3" pageNumber="4">C. gayi</emphasis>
</taxonomicName>
</th>
<td box="[557,737,920,941]" gridcol="1" gridrow="2" pageId="3" pageNumber="4">unknown</td>
<td box="[757,934,920,941]" gridcol="2" gridrow="2" pageId="3" pageNumber="4">
M13105/cas:sua:10082
<emphasis box="[925,934,924,940]" italics="true" pageId="3" pageNumber="4">*</emphasis>
</td>
<td box="[957,1057,920,941]" gridcol="3" gridrow="2" pageId="3" pageNumber="4">54.7</td>
<td box="[1080,1174,920,941]" gridcol="4" gridrow="2" pageId="3" pageNumber="4">2.9</td>
<td box="[1200,1286,920,941]" gridcol="5" gridrow="2" pageId="3" pageNumber="4">5.3</td>
<td box="[1306,1418,920,941]" gridcol="6" gridrow="2" pageId="3" pageNumber="4">
<bibRefCitation author="Unknown" box="[1306,1318,920,932]" pageId="3" pageNumber="4" publicationUrl="https://www.morphosource.org/" refId="ref8453" refString="65. Unknown. Morphosource. Available at, https: // www. morphosource. org / (2020)." title="Morphosource. Available at" type="url" year="2020">65</bibRefCitation>
, see Fig.S3
</td>
</tr>
<tr box="[423,1418,955,972]" gridrow="3" pageId="3" pageNumber="4" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-6="1">
<th box="[423,540,955,972]" gridcol="0" gridrow="3" pageId="3" pageNumber="4">range</th>
<td box="[1200,1286,955,972]" gridcol="5" gridrow="3" pageId="3" pageNumber="4">5.35.6</td>
</tr>
<tr box="[423,1418,985,1003]" gridrow="4" pageId="3" pageNumber="4" rowspan-1="1" rowspan-2="1" rowspan-3="1" rowspan-4="1" rowspan-6="1">
<th box="[423,540,985,1003]" gridcol="0" gridrow="4" pageId="3" pageNumber="4">mean ± SD</th>
<td box="[1200,1286,985,1003]" gridcol="5" gridrow="4" pageId="3" pageNumber="4">5.45 ± 0.21</td>
</tr>
<tr box="[423,1418,1016,1034]" gridrow="5" pageId="3" pageNumber="4" rowspan-5="1">
<th box="[423,540,1016,1034]" gridcol="0" gridrow="5" pageId="3" pageNumber="4">
<emphasis box="[423,435,1017,1033]" italics="true" pageId="3" pageNumber="4">C</emphasis>
. sp.
</th>
<td box="[557,737,1016,1034]" gridcol="1" gridrow="5" pageId="3" pageNumber="4">IAA 2/95, Marsupial site</td>
<td box="[757,934,1016,1034]" gridcol="2" gridrow="5" pageId="3" pageNumber="4">NRM-PZ B282</td>
<td box="[957,1057,1016,1034]" gridcol="3" gridrow="5" pageId="3" pageNumber="4">
<emphasis bold="true" box="[957,1032,1016,1034]" pageId="3" pageNumber="4">38.4 ± 3.8</emphasis>
</td>
<td box="[1080,1174,1016,1034]" gridcol="4" gridrow="5" pageId="3" pageNumber="4">
<emphasis bold="true" box="[1080,1103,1017,1034]" pageId="3" pageNumber="4">2.1</emphasis>
</td>
<td box="[1306,1418,1016,1034]" gridcol="6" gridrow="5" pageId="3" pageNumber="4">Present work</td>
</tr>
</table>
</paragraph>
<caption pageId="3" pageNumber="4">
<paragraph blockId="3.[415,1469,1067,1169]" pageId="3" pageNumber="4">
<emphasis bold="true" box="[415,493,1067,1088]" pageId="3" pageNumber="4">Table 1.</emphasis>
Measurements of the snout-vent length (SVL) and height of the transition from the iliac shaft and ilial body (HT) of some
<taxonomicName box="[603,768,1094,1115]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[603,768,1094,1115]" italics="true" pageId="3" pageNumber="4">Calyptocephalella</emphasis>
</taxonomicName>
spp. and Antarctic ilia (Fig. S3), with the value of the ratio between HT and SVL (RHS = HT/SVL
<emphasis box="[662,674,1122,1143]" italics="true" pageId="3" pageNumber="4">*</emphasis>
100%) with the value of the standard deviation (SD). The reconstructed value of the snout-vent length of the studied ilium is highlighted in bold.
</paragraph>
</caption>
<paragraph blockId="3.[415,1480,1249,1968]" lastBlockId="4.[415,1480,1200,1622]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
The fragmentary right ilium can be referred to an anuran based on the following characters
<bibRefCitation author="Gardner, J. D." box="[1328,1343,1569,1583]" journalOrPublisher="J. Vertebr. Paleontol." pageId="3" pageNumber="4" pagination="1684 - 1696" part="30" refId="ref6866" refString="30. Gardner, J. D. et al. Comparative morphology of the ilium of anurans and urodeles (Lissamphibia) and a re-assessment of the anuran affinities of Nezpercius dodsoni Blob et al., 2001. J. Vertebr. Paleontol. 30, 1684 - 1696 (2010)." title="Comparative morphology of the ilium of anurans and urodeles (Lissamphibia) and a re-assessment of the anuran affinities of Nezpercius dodsoni Blob et al" type="journal article" year="2010">
<superScript attach="left" box="[1328,1343,1569,1583]" fontSize="6" pageId="3" pageNumber="4">30</superScript>
</bibRefCitation>
(the numbers before the characters correspond to the feature numbers of Appendix 1 in Gardner
<emphasis box="[1223,1264,1599,1621]" italics="true" pageId="3" pageNumber="4">et al</emphasis>
.
<bibRefCitation author="Gardner, J. D." box="[1269,1284,1596,1610]" journalOrPublisher="J. Vertebr. Paleontol." pageId="3" pageNumber="4" pagination="1684 - 1696" part="30" refId="ref6866" refString="30. Gardner, J. D. et al. Comparative morphology of the ilium of anurans and urodeles (Lissamphibia) and a re-assessment of the anuran affinities of Nezpercius dodsoni Blob et al., 2001. J. Vertebr. Paleontol. 30, 1684 - 1696 (2010)." title="Comparative morphology of the ilium of anurans and urodeles (Lissamphibia) and a re-assessment of the anuran affinities of Nezpercius dodsoni Blob et al" type="journal article" year="2010">
<superScript attach="right" box="[1269,1284,1596,1610]" fontSize="6" pageId="3" pageNumber="4">30</superScript>
</bibRefCitation>
):
<emphasis box="[1304,1315,1600,1621]" italics="true" pageId="3" pageNumber="4">7</emphasis>
. (semi-) circular acetabulum;
<emphasis box="[540,551,1626,1647]" italics="true" pageId="3" pageNumber="4">9</emphasis>
. acetabulum with distinct margins;
<emphasis box="[901,924,1626,1647]" italics="true" pageId="3" pageNumber="4">10</emphasis>
. acetabular surface concave;
<emphasis box="[1206,1229,1626,1647]" italics="true" pageId="3" pageNumber="4">13</emphasis>
. at least dorsal acetabular expansion is strongly divergent;
<emphasis box="[725,748,1653,1674]" italics="true" pageId="3" pageNumber="4">18</emphasis>
. the dorsal protuberance present. Thus, the ilium derives from a small-sized frog (3.8 ± 0.4 cm snout-vent length, see methods,
<tableCitation box="[914,984,1679,1701]" captionStart="Table 1" captionStartId="3.[415,469,1067,1088]" captionText="Table 1. Measurements of the snout-vent length (SVL) and height of the transition from the iliac shaft and ilial body (HT) of some Calyptocephalella spp. and Antarctic ilia (Fig. S3), with the value of the ratio between HT and SVL (RHS=HT/SVL*100%) with the value of the standard deviation (SD). The reconstructed value of the snout-vent length of the studied ilium is highlighted in bold." pageId="3" pageNumber="4">Table 1</tableCitation>
). The specimen is partly eroded and rather poorly preserved; however, it can be compared with all South American and Australian frog families (
<figureCitation box="[1337,1402,1706,1728]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="3" pageNumber="4">Figs. 4</figureCitation>
, S
<figureCitation box="[1428,1437,1706,1727]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="3" pageNumber="4">1</figureCitation>
and S
<figureCitation box="[428,440,1733,1754]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="3" pageNumber="4">2</figureCitation>
,
<tableCitation box="[449,531,1732,1755]" captionStart="Table 1" captionStartId="3.[415,469,1067,1088]" captionText="Table 1. Measurements of the snout-vent length (SVL) and height of the transition from the iliac shaft and ilial body (HT) of some Calyptocephalella spp. and Antarctic ilia (Fig. S3), with the value of the ratio between HT and SVL (RHS=HT/SVL*100%) with the value of the standard deviation (SD). The reconstructed value of the snout-vent length of the studied ilium is highlighted in bold." pageId="3" pageNumber="4">Table S1</tableCitation>
). The families
<taxonomicName box="[672,753,1732,1754]" class="Amphibia" family="Ranidae" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Ranidae</taxonomicName>
,
<taxonomicName box="[762,862,1732,1754]" class="Amphibia" family="Bufonidae" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Bufonidae</taxonomicName>
and
<taxonomicName authorityName="Rafinesque" authorityYear="1815" box="[909,985,1732,1754]" class="Amphibia" family="Hylidae" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Hylidae</taxonomicName>
have not been illustrated in the present work, since their morphology is well known
<bibRefCitation author="Bailon, S." box="[720,735,1756,1770]" journalOrPublisher="Centre de Recherches Archeologiques du CNRS" pageId="3" pageNumber="4" refId="ref6921" refString="31. Bailon, S. Differenciation osteologique des Anoures (Amphibia, Anura) de France 1 - 41 (Centre de Recherches Archeologiques du CNRS, 1999)." title="Differenciation osteologique des Anoures (Amphibia, Anura) de France 1" type="book" year="1999">
<superScript attach="left" box="[720,735,1756,1770]" fontSize="6" pageId="3" pageNumber="4">31</superScript>
</bibRefCitation>
(
<tableCitation box="[746,828,1759,1781]" captionStart="Table 1" captionStartId="3.[415,469,1067,1088]" captionText="Table 1. Measurements of the snout-vent length (SVL) and height of the transition from the iliac shaft and ilial body (HT) of some Calyptocephalella spp. and Antarctic ilia (Fig. S3), with the value of the ratio between HT and SVL (RHS=HT/SVL*100%) with the value of the standard deviation (SD). The reconstructed value of the snout-vent length of the studied ilium is highlighted in bold." pageId="3" pageNumber="4">Table S1</tableCitation>
). The comparison has been done at family level, since the ilia display diagnostic features characteristic for identification of the family (dimensions of the dorsal and ventral acetabular expansions; location of the dorsal protuberance relative to the anterior margin of the acetabular rim etc.
<superScript attach="none" box="[1431,1465,1809,1823]" fontSize="6" pageId="3" pageNumber="4">
<bibRefCitation author="Rage, J. - C." box="[1431,1446,1809,1823]" journalOrPublisher="Foss. Imprint" pageId="3" pageNumber="4" pagination="53 - 66" part="72" refId="ref6952" refString="32. Rage, J. - C. Frogs (Amphibia, Anura) from the Eocene and Oligocene of the phosphorites du Quercy (France). An overview. Foss. Imprint 72, 53 - 66 (2016)." title="Frogs (Amphibia, Anura) from the Eocene and Oligocene of the phosphorites du Quercy (France). An overview" type="journal article" year="2016">32</bibRefCitation>
,
<bibRefCitation author="Folie, A." box="[1450,1465,1809,1823]" journalOrPublisher="Acta Palaeontol. Pol." pageId="3" pageNumber="4" pagination="511 - 524" part="58" refId="ref6996" refString="33. Folie, A. et al. Early Eocene frogs from Vastan Lignite Mine, Gujarat, India. Acta Palaeontol. Pol. 58, 511 - 524 (2013)." title="Early Eocene frogs from Vastan Lignite Mine, Gujarat, India" type="journal article" year="2013">33</bibRefCitation>
</superScript>
). The studied ilium (NRM-PZ B282) differs in: (1) Reduced anterior portion of the dorsal acetabular expansion from nearly all South American and Australian frog families and the genus
<taxonomicName authorityName="Schmidt" authorityYear="1952" box="[1158,1277,1866,1887]" class="Amphibia" family="Calyptocephalellidae" genus="Telmatobufo" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[1158,1277,1866,1887]" italics="true" pageId="3" pageNumber="4">Telmatobufo</emphasis>
</taxonomicName>
, which have moderately or strongly developed anterior portion of the dorsal acetabular expansion. Only the genus
<taxonomicName box="[1314,1479,1893,1914]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[1314,1479,1893,1914]" italics="true" pageId="3" pageNumber="4">Calyptocephalella</emphasis>
</taxonomicName>
(
<figureCitation box="[422,502,1919,1941]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="3" pageNumber="4">Fig. 4c,e</figureCitation>
), the families
<taxonomicName box="[635,714,1919,1941]" class="Amphibia" family="Ranidae" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Ranidae</taxonomicName>
<bibRefCitation author="Bailon, S." box="[714,729,1916,1930]" journalOrPublisher="Centre de Recherches Archeologiques du CNRS" pageId="3" pageNumber="4" refId="ref6921" refString="31. Bailon, S. Differenciation osteologique des Anoures (Amphibia, Anura) de France 1 - 41 (Centre de Recherches Archeologiques du CNRS, 1999)." title="Differenciation osteologique des Anoures (Amphibia, Anura) de France 1" type="book" year="1999">
<superScript attach="left" box="[714,729,1916,1930]" fontSize="6" pageId="3" pageNumber="4">31</superScript>
</bibRefCitation>
,
<taxonomicName box="[739,811,1919,1941]" class="Amphibia" family="Pipidae" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Pipidae</taxonomicName>
(
<figureCitation box="[823,900,1919,1941]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="3" pageNumber="4">Fig. S1a</figureCitation>
),
<taxonomicName authorityName="Bonaparte" authorityYear="1850" box="[917,1086,1919,1941]" class="Amphibia" family="Rhinodermatidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Rhinodermatidae</taxonomicName>
(
<figureCitation box="[1097,1170,1919,1941]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="3" pageNumber="4">Fig. S1j</figureCitation>
), and
<taxonomicName box="[1228,1381,1919,1941]" class="Amphibia" family="Leptodactylidae" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Leptodactylidae</taxonomicName>
(
<figureCitation box="[1393,1472,1919,1941]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="3" pageNumber="4">Fig. S2b</figureCitation>
) have similar state/morphology of this character. (2) Dorsal protuberance located either at the level of or anteriorly from the anterior margin of the acetabular rim from nearly all families, besides
<taxonomicName box="[1200,1373,1200,1222]" class="Amphibia" family="Brachycephalidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Brachycephalidae</taxonomicName>
(
<figureCitation box="[1386,1466,1200,1223]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="4" pageNumber="5">Fig. S1a</figureCitation>
),
<taxonomicName authorityName="Bonaparte" authorityYear="1850" box="[415,588,1227,1249]" class="Amphibia" family="Rhinodermatidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Rhinodermatidae</taxonomicName>
(
<figureCitation box="[601,676,1227,1249]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="4" pageNumber="5">Fig. S1j</figureCitation>
),
<taxonomicName authorityName="Fitzinger" authorityYear="1843" box="[695,834,1227,1249]" class="Amphibia" family="Telmatobiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Telmatobiidae</taxonomicName>
(
<figureCitation box="[847,929,1227,1249]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="4" pageNumber="5">Fig. S1k</figureCitation>
), Hyloididae (
<figureCitation box="[1069,1159,1227,1249]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="4" pageNumber="5">Fig. S1m</figureCitation>
),
<taxonomicName box="[1177,1334,1227,1249]" class="Amphibia" family="Leptodactylidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Leptodactylidae</taxonomicName>
(
<figureCitation box="[1347,1429,1227,1249]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="4" pageNumber="5">Fig. S2b</figureCitation>
) and the genera
<taxonomicName box="[520,685,1254,1275]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[520,685,1254,1275]" italics="true" pageId="4" pageNumber="5">Calyptocephalella</emphasis>
</taxonomicName>
(
<figureCitation box="[697,750,1254,1276]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5">Fig. 4</figureCitation>
be) and
<taxonomicName authorityName="Schmidt" authorityYear="1952" box="[840,956,1254,1275]" class="Amphibia" family="Calyptocephalellidae" genus="Telmatobufo" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[840,956,1254,1275]" italics="true" pageId="4" pageNumber="5">Telmatobufo</emphasis>
</taxonomicName>
(
<figureCitation box="[969,1053,1254,1276]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5">Fig. 4g,h</figureCitation>
). (3) Developed dorsal acetabular expansion from the families
<taxonomicName box="[587,667,1280,1302]" class="Amphibia" family="Ranidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Ranidae</taxonomicName>
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<superScript attach="left" box="[666,681,1277,1291]" fontSize="6" pageId="4" pageNumber="5">31</superScript>
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,
<taxonomicName authorityName="Rafinesque" authorityYear="1815" box="[693,769,1280,1302]" class="Amphibia" family="Hylidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Hylidae</taxonomicName>
<bibRefCitation author="Bailon, S." box="[768,783,1277,1291]" journalOrPublisher="Centre de Recherches Archeologiques du CNRS" pageId="4" pageNumber="5" refId="ref6921" refString="31. Bailon, S. Differenciation osteologique des Anoures (Amphibia, Anura) de France 1 - 41 (Centre de Recherches Archeologiques du CNRS, 1999)." title="Differenciation osteologique des Anoures (Amphibia, Anura) de France 1" type="book" year="1999">
<superScript attach="left" box="[768,783,1277,1291]" fontSize="6" pageId="4" pageNumber="5">31</superScript>
</bibRefCitation>
,
<taxonomicName box="[795,895,1280,1302]" class="Amphibia" family="Bufonidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Bufonidae</taxonomicName>
<bibRefCitation author="Bailon, S." box="[894,909,1277,1291]" journalOrPublisher="Centre de Recherches Archeologiques du CNRS" pageId="4" pageNumber="5" refId="ref6921" refString="31. Bailon, S. Differenciation osteologique des Anoures (Amphibia, Anura) de France 1 - 41 (Centre de Recherches Archeologiques du CNRS, 1999)." title="Differenciation osteologique des Anoures (Amphibia, Anura) de France 1" type="book" year="1999">
<superScript attach="left" box="[894,909,1277,1291]" fontSize="6" pageId="4" pageNumber="5">31</superScript>
</bibRefCitation>
,
<taxonomicName authorityName="Schlegel" authorityYear="1850" box="[921,1079,1280,1302]" class="Amphibia" family="Myobatrachidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Myobatrachidae</taxonomicName>
(
<figureCitation box="[1091,1152,1280,1302]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5">Fig. 4i</figureCitation>
),
<taxonomicName box="[1172,1245,1280,1302]" class="Amphibia" family="Pipidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Pipidae</taxonomicName>
(
<figureCitation box="[1257,1332,1280,1302]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="4" pageNumber="5">Fig. S1i</figureCitation>
),
<taxonomicName box="[1349,1479,1280,1302]" class="Amphibia" family="Microhylidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Microhylidae</taxonomicName>
(
<figureCitation box="[422,508,1307,1329]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="4" pageNumber="5">Fig. S1b</figureCitation>
),
<taxonomicName authorityName="Fitzinger" authorityYear="1843" box="[528,671,1307,1329]" class="Amphibia" family="Telmatobiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Telmatobiidae</taxonomicName>
(
<figureCitation box="[685,771,1307,1329]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="4" pageNumber="5">Fig. S1k</figureCitation>
),
<taxonomicName box="[790,952,1307,1329]" class="Amphibia" family="Leptodactylidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Leptodactylidae</taxonomicName>
(
<figureCitation box="[966,1053,1307,1329]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="4" pageNumber="5">Fig. S2b</figureCitation>
),
<taxonomicName authorityName="Goin, Goin &amp; Zug" authorityYear="1978" box="[1071,1218,1307,1329]" class="Amphibia" family="Allophrynidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Allophrynidae</taxonomicName>
(
<figureCitation box="[1233,1317,1307,1329]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="4" pageNumber="5">Fig. S2c</figureCitation>
),
<taxonomicName authorityName="Taylor" authorityYear="1951" box="[1336,1479,1307,1329]" class="Amphibia" family="Centrolenidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Centrolenidae</taxonomicName>
(
<figureCitation box="[422,503,1334,1356]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="4" pageNumber="5">Fig. S2d</figureCitation>
) and the genus
<taxonomicName authorityName="Schmidt" authorityYear="1952" box="[654,772,1334,1355]" class="Amphibia" family="Calyptocephalellidae" genus="Telmatobufo" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[654,772,1334,1355]" italics="true" pageId="4" pageNumber="5">Telmatobufo</emphasis>
</taxonomicName>
(
<figureCitation box="[783,868,1334,1356]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5">Fig. 4g,h</figureCitation>
). Other families have moderately or well-developed dorsal acetabular expansion, however, due to incomplete preservation of the Antarctic frog remain any further comparison is impossible. (4) Weakly developed dorsal protuberance and lack of dorsal tubercle from nearly all families (e.g.
<taxonomicName authorityName="Lynch" authorityYear="1969" box="[415,583,1414,1436]" class="Amphibia" family="Limnodynastidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Limnodynastidae</taxonomicName>
,
<figureCitation box="[591,649,1414,1436]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5">Fig. 4j</figureCitation>
), besides
<taxonomicName authorityName="Reig" authorityYear="1960" box="[741,940,1414,1436]" class="Amphibia" family="Calyptocephalellidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Calyptocephalellidae</taxonomicName>
(
<figureCitation box="[950,1004,1414,1436]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5">Fig. 4</figureCitation>
be,g,h),
<taxonomicName authorityName="Schlegel" authorityYear="1850" box="[1091,1245,1414,1436]" class="Amphibia" family="Myobatrachidae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Myobatrachidae</taxonomicName>
(
<figureCitation box="[1256,1316,1414,1436]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5">Fig. 4i</figureCitation>
),
<taxonomicName authorityName="Hedges, Duellman &amp; Heinicke" authorityYear="2008" box="[1334,1479,1414,1436]" class="Amphibia" family="Craugastoridae" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Craugastoridae</taxonomicName>
(
<figureCitation box="[422,499,1440,1463]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="4" pageNumber="5">Fig. S1e</figureCitation>
), and Dendrabatidae (
<figureCitation box="[711,790,1440,1463]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="4" pageNumber="5">Fig. S2e</figureCitation>
).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3766735" ID-Zenodo-Dep="3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5" startId="4.[415,480,1050,1072]" targetBox="[418,1468,125,1012]" targetPageId="4">
<paragraph blockId="4.[415,1360,1050,1126]" pageId="4" pageNumber="5">
<emphasis bold="true" box="[415,503,1050,1072]" pageId="4" pageNumber="5">Figure 4.</emphasis>
3D models of some skeletal elements of Australobatrachia. (
<emphasis bold="true" box="[1083,1108,1050,1072]" pageId="4" pageNumber="5">a,f</emphasis>
) skull and (
<emphasis bold="true" box="[1221,1256,1051,1072]" pageId="4" pageNumber="5">be</emphasis>
,
<emphasis bold="true" box="[1262,1293,1050,1072]" pageId="4" pageNumber="5">gj</emphasis>
) ilia of
<taxonomicName baseAuthorityName="Duméril &amp; Bibron" baseAuthorityYear="1841" box="[415,624,1077,1099]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="gayi">
<emphasis box="[415,624,1077,1099]" italics="true" pageId="4" pageNumber="5">Calyptocephalella gayi</emphasis>
</taxonomicName>
(
<emphasis bold="true" box="[637,671,1077,1098]" pageId="4" pageNumber="5">ae</emphasis>
);
<taxonomicName baseAuthorityName="Philippi" baseAuthorityYear="1899" box="[690,892,1077,1099]" class="Amphibia" family="Calyptocephalellidae" genus="Telmatobufo" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="venustus">
<emphasis box="[690,892,1077,1099]" italics="true" pageId="4" pageNumber="5">Telmatobufo venustus</emphasis>
</taxonomicName>
(
<emphasis bold="true" box="[906,939,1077,1099]" pageId="4" pageNumber="5">fh</emphasis>
);
<taxonomicName baseAuthorityName="Gray" baseAuthorityYear="1841" box="[958,1160,1077,1098]" class="Amphibia" family="Myobatrachidae" genus="Myobatrachus" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="gouldii">
<emphasis box="[958,1160,1077,1098]" italics="true" pageId="4" pageNumber="5">Myobatrachus gouldii</emphasis>
</taxonomicName>
(
<emphasis bold="true" box="[1174,1181,1077,1099]" pageId="4" pageNumber="5">i</emphasis>
);
<taxonomicName baseAuthorityName="Macleay" baseAuthorityYear="1878" class="Amphibia" family="Limnodynastidae" genus="Limnodynastes" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="convexiusculus">
<emphasis italics="true" pageId="4" pageNumber="5">Limnodynastes convexiusculus</emphasis>
</taxonomicName>
(
<emphasis bold="true" box="[568,575,1104,1126]" pageId="4" pageNumber="5">j</emphasis>
). Collection numbers of each specimen are listed in Table S1.
</paragraph>
</caption>
<paragraph blockId="4.[415,1480,1200,1622]" pageId="4" pageNumber="5">
Among the compared forms, only the South American endemic genus
<taxonomicName box="[1127,1294,1467,1488]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1127,1294,1467,1488]" italics="true" pageId="4" pageNumber="5">Calyptocephalella</emphasis>
</taxonomicName>
resembles all mentioned four characters. In addition to this, a shallow and broad depression on the anterior portion of the ventral acetabular expansion is a unique character observable on our ilium (NRM-PZ B282) and Recent
<taxonomicName baseAuthorityName="Duméril &amp; Bibron" baseAuthorityYear="1841" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="gayi">
<emphasis italics="true" pageId="4" pageNumber="5">Calyptocephalella gayi</emphasis>
</taxonomicName>
(
<figureCitation box="[466,520,1547,1569]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5">Fig. 4</figureCitation>
). Further, the fossil ilium displays a ventral depression on its lateral surface anteriorly to the acetabulum (
<figureCitation box="[464,528,1574,1596]" captionStart="Figure 2" captionStartId="2.[415,480,1227,1249]" captionTargetBox="[417,1469,126,1190]" captionTargetId="figure@2.[415,1471,124,1192]" captionTargetPageId="2" captionText="Figure 2. Ilium (NRM-PZ B282) of Calyptocephalella sp. from Seymour Island, Antarctica. Ilium in lateral (a), medial (b), ventral (c) and dorsal (d) views. Magnified region of the dorsal protuberance in lateral (e dashed rectangle in orange color) and dorsal (f dashed rectangle in yellow color) view. The dashed line in black on (a) indicates the probable outline of the posterior extension of the ventral acetabular expansion. The asterisk (*) on (a,c) indicates the shallow and broad depression of the ventral acetabular expansion. The double asterisk (**) indicates the notch caudally from the dorsal protuberance.The dashed red lines on (e,f) outline the intact bone surface. Abbreviations:ac, acetabulum; ar, acetabular rim; dae, dorsal acetabular expansion; is, iliac shaft; paf, preacetabular fossa; spf, supraacetabular fossa; vae, ventral acetabular expansion; vd, ventral depression." figureDoi="http://doi.org/10.5281/zenodo.3766731" httpUri="https://zenodo.org/record/3766731/files/figure.png" pageId="4" pageNumber="5">Fig. 2c</figureCitation>
), a comparable structure can be observed also in the fossil species
<taxonomicName authorityName="Schaeffer" authorityYear="1949" box="[1151,1398,1574,1595]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="canqueli">
<emphasis box="[1151,1398,1574,1595]" italics="true" pageId="4" pageNumber="5">Calyptocephalella canqueli</emphasis>
</taxonomicName>
<bibRefCitation author="Agnolin, F." box="[1399,1406,1570,1584]" journalOrPublisher="Stud. Geol. Salamanticensia" pageId="4" pageNumber="5" pagination="129 - 178" part="48" refId="ref5798" refString="9. Agnolin, F. A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position. Stud. Geol. Salamanticensia 48, 129 - 178 (2012)." title="A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position" type="journal article" year="2012">
<superScript attach="left" box="[1399,1406,1570,1584]" fontSize="6" pageId="4" pageNumber="5">9</superScript>
</bibRefCitation>
but not in the Recent species
<taxonomicName baseAuthorityName="Duméril &amp; Bibron" baseAuthorityYear="1841" box="[618,682,1601,1622]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="gayi">
<emphasis box="[618,682,1601,1622]" italics="true" pageId="4" pageNumber="5">C. gayi</emphasis>
</taxonomicName>
(
<figureCitation box="[694,748,1600,1622]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="4" pageNumber="5">Fig. 4</figureCitation>
).
</paragraph>
<paragraph blockId="4.[415,1480,1654,1972]" lastBlockId="5.[415,1480,136,959]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
<emphasis bold="true" box="[415,579,1654,1678]" pageId="4" pageNumber="5">Skull element.</emphasis>
The second bone fragment (NRM-PZ B281) is flat and slightly curved. Both sides of the bone have different structures. One surface is covered by small to large circular or reniform in outline, rather deep pits, which sink in the planar surface of the bone (
<figureCitation box="[886,950,1710,1732]" captionStart="Figure 3" captionStartId="3.[415,480,746,768]" captionTargetBox="[418,1372,127,710]" captionTargetId="figure@3.[415,1375,124,711]" captionTargetPageId="3" captionText="Figure 3. Skull bone fragment (NRM-PZ B281) of Calyptocephalella sp. from Seymour Island, Antarctica in dorsal (a), ventral (b) and lateral (c) views." figureDoi="http://doi.org/10.5281/zenodo.3766733" httpUri="https://zenodo.org/record/3766733/files/figure.png" pageId="4" pageNumber="5">Fig. 3a</figureCitation>
). The diameters of pits vary from 0.10.7 mm and some of them are punctured by foramina. The opposite surface of the bone is in general smooth, slightly deepened and is pierced with some foramina, some of which are preceded by a groove (
<figureCitation box="[1154,1220,1764,1786]" captionStart="Figure 3" captionStartId="3.[415,480,746,768]" captionTargetBox="[418,1372,127,710]" captionTargetId="figure@3.[415,1375,124,711]" captionTargetPageId="3" captionText="Figure 3. Skull bone fragment (NRM-PZ B281) of Calyptocephalella sp. from Seymour Island, Antarctica in dorsal (a), ventral (b) and lateral (c) views." figureDoi="http://doi.org/10.5281/zenodo.3766733" httpUri="https://zenodo.org/record/3766733/files/figure.png" pageId="4" pageNumber="5">Fig. 3b</figureCitation>
). One side of the fragment preserves an unbroken margin of the original bone with a distinct process that is bent and that gives the bone a curved shape (
<figureCitation box="[574,641,1817,1839]" captionStart="Figure 3" captionStartId="3.[415,480,746,768]" captionTargetBox="[418,1372,127,710]" captionTargetId="figure@3.[415,1375,124,711]" captionTargetPageId="3" captionText="Figure 3. Skull bone fragment (NRM-PZ B281) of Calyptocephalella sp. from Seymour Island, Antarctica in dorsal (a), ventral (b) and lateral (c) views." figureDoi="http://doi.org/10.5281/zenodo.3766733" httpUri="https://zenodo.org/record/3766733/files/figure.png" pageId="4" pageNumber="5">Fig. 3c</figureCitation>
). The ornamented surface of the bone projects slightly over this process. Comparable ornamentation, build of pits of different size, is found on the dorsal surfaces of different cranial and postcranial bones of amphibians and reptiles
<superScript attach="right" box="[735,769,1867,1881]" fontSize="6" pageId="4" pageNumber="5">
<bibRefCitation author="Clarac, F. &amp; Buffrenil, V. &amp; Brochu, C. &amp; Cubo, J." box="[735,750,1867,1881]" journalOrPublisher="Biol. J. Linn. Soc." pageId="4" pageNumber="5" pagination="395 - 408" part="121" refId="ref7031" refString="34. Clarac, F., Buffrenil, V., de, Brochu, C. &amp; Cubo, J. The evolution of bone ornamentation in Pseudosuchia: morphological constraints versus ecological adaptation. Biol. J. Linn. Soc. 121, 395 - 408 (2017)." title="The evolution of bone ornamentation in Pseudosuchia: morphological constraints versus ecological adaptation" type="journal article" year="2017">34</bibRefCitation>
,
<bibRefCitation author="Scheyer, T. M. &amp; Sander, P. M. &amp; Joyce, W. G. &amp; Bohme, W. &amp; Witzel, U." box="[754,769,1867,1881]" journalOrPublisher="Org. Divers. Evol." pageId="4" pageNumber="5" pagination="136 - 144" part="7" refId="ref7085" refString="35. Scheyer, T. M., Sander, P. M., Joyce, W. G., Bohme, W. &amp; Witzel, U. A plywood structure in the shell of fossil and living soft-shelled turtles (Trionychidae) and its evolutionary implications. Org. Divers. Evol. 7, 136 - 144 (2007)." title="A plywood structure in the shell of fossil and living soft-shelled turtles (Trionychidae) and its evolutionary implications" type="journal article" year="2007">35</bibRefCitation>
</superScript>
. Among them, the following groups can be excluded from consideration: (1)
<taxonomicName authorityName="Fox &amp; Naylor" authorityYear="1982" box="[448,628,1897,1919]" class="Amphibia" family="Albanerpetontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Gymnophiona" pageId="4" pageNumber="5" phylum="Chordata" rank="family">Albanerpetontidae</taxonomicName>
(
<taxonomicName authorityName="Fox &amp; Naylor" authorityYear="1982" box="[640,761,1897,1919]" class="Amphibia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Allocaudata" pageId="4" pageNumber="5" phylum="Chordata" rank="order">Allocaudata</taxonomicName>
); albanerpetontids are a primary Laurasian lissamphibian group with a single occurrence in Northern Africa. So far no evidence of a Gondwanan radiation of albanerpetontids exists
<bibRefCitation author="Gardner, J. D. &amp; Evans, S. E. &amp; Sigogneau-Russell, D." box="[1458,1473,1920,1934]" journalOrPublisher="Acta Palaeontol. Pol." pageId="4" pageNumber="5" pagination="301 - 319" part="48" refId="ref7152" refString="36. Gardner, J. D., Evans, S. E. &amp; Sigogneau-Russell, D. New albanerpetontid amphibians from the Early Cretaceous of Morocco and Middle Jurassic of England. Acta Palaeontol. Pol. 48, 301 - 319 (2003)." title="New albanerpetontid amphibians from the Early Cretaceous of Morocco and Middle Jurassic of England" type="journal article" year="2003">
<superScript attach="left" box="[1458,1473,1920,1934]" fontSize="6" pageId="4" pageNumber="5">36</superScript>
</bibRefCitation>
. In addition to this, all their ornamented bones (e.g. frontal, premaxillae)
<bibRefCitation author="Gardner, J. D." box="[1114,1129,1947,1961]" journalOrPublisher="Geodiversitas" pageId="4" pageNumber="5" pagination="349 - 388" part="22" refId="ref7201" refString="37. Gardner, J. D. Albanerpetontid amphibians from the Upper Cretaceous (Campanian and Maastrichtian) of North America. Geodiversitas 22, 349 - 388 (2000)." title="Albanerpetontid amphibians from the Upper Cretaceous (Campanian and Maastrichtian) of North America" type="journal article" year="2000">
<superScript attach="left" box="[1114,1129,1947,1961]" fontSize="6" pageId="4" pageNumber="5">37</superScript>
</bibRefCitation>
do not resemble the bone described here. (2)
<taxonomicName box="[500,582,136,158]" class="Amphibia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="5" pageNumber="6" phylum="Chordata" rank="order">Caudata</taxonomicName>
; salamanders are also considered as a Laurasian group, with a number of occurrences in Africa which need critical revision
<bibRefCitation author="Gardner, J. D. &amp; Rage, J. - C." box="[678,693,160,174]" journalOrPublisher="Palaeobio. Palaeoenv." pageId="5" pageNumber="6" pagination="169 - 220" part="96" refId="ref7234" refString="38. Gardner, J. D. &amp; Rage, J. - C. The fossil record of lissamphibians from Africa, Madagascar, and the Arabian Plate. Palaeobio. Palaeoenv. 96, 169 - 220 (2016)." title="The fossil record of lissamphibians from Africa, Madagascar, and the Arabian Plate" type="journal article" year="2016">
<superScript attach="left" box="[678,693,160,174]" fontSize="6" pageId="5" pageNumber="6">38</superScript>
</bibRefCitation>
. In salamanders, ornamented bones are found both among skull bones and on vertebrae (on plates located on the tip of the neural arch)
<bibRefCitation author="Estes, R." box="[924,939,186,200]" journalOrPublisher="Gustav Fischer" pageId="5" pageNumber="6" refId="ref7278" refString="39. Estes, R. Gymnophiona, Caudata 1 - 115 (Gustav Fischer, 1981)." title="Gymnophiona, Caudata 1" type="book" year="1981">
<superScript attach="none" box="[924,939,186,200]" fontSize="6" pageId="5" pageNumber="6">39</superScript>
</bibRefCitation>
. Bone ornamentation here (e.g.
<taxonomicName authority=", Chelotriton, Echinotriton" authorityName="Chelotriton, Echinotriton" class="Amphibia" family="Salamandridae" genus="Tylototriton" kingdom="Animalia" order="Caudata" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="5" pageNumber="6">Tylototriton, Chelotriton, Echinotriton</emphasis>
</taxonomicName>
<superScript attach="left" box="[536,570,213,227]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Estes, R." box="[536,551,213,227]" journalOrPublisher="Gustav Fischer" pageId="5" pageNumber="6" refId="ref7278" refString="39. Estes, R. Gymnophiona, Caudata 1 - 115 (Gustav Fischer, 1981)." title="Gymnophiona, Caudata 1" type="book" year="1981">39</bibRefCitation>
,
<bibRefCitation author="Schoch, R. &amp; Poschmann, M. &amp; Kupfer, A." box="[555,570,213,227]" journalOrPublisher="Palaeobio. Palaeoenv." pageId="5" pageNumber="6" pagination="77 - 86" part="95" refId="ref7297" refString="40. Schoch, R., Poschmann, M. &amp; Kupfer, A. The salamandrid Chelotriton paradoxus from Enspel and Randeck Maars (Oligocene-Miocene, Germany). Palaeobio. Palaeoenv. 95, 77 - 86 (2015)." title="The salamandrid Chelotriton paradoxus from Enspel and Randeck Maars (Oligocene-Miocene, Germany)" type="journal article" year="2015">40</bibRefCitation>
</superScript>
) is represented by a network of pits, ridges and pointy spines that do not resemble the bone described here. (3) Crocodylia; in crocodyliforms, comparable patterns of ornamentation with well-developed pits appear only with growth during later ontogenetic stages
<superScript attach="left" box="[993,1031,266,281]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Vickaryous, M. K. &amp; Hall, B. K." box="[993,1008,266,280]" journalOrPublisher="J. Morphol." pageId="5" pageNumber="6" pagination="398 - 422" part="269" refId="ref7341" refString="41. Vickaryous, M. K. &amp; Hall, B. K. Development of the dermal skeleton in Alligator mississippiensis (Archosauria, Crocodylia) with comments on the homology of osteoderms. J. Morphol. 269, 398 - 422 (2008)." title="Development of the dermal skeleton in Alligator mississippiensis (Archosauria, Crocodylia) with comments on the homology of osteoderms" type="journal article" year="2008">41</bibRefCitation>
<bibRefCitation author="Buffrenil" box="[1016,1031,266,280]" journalOrPublisher="Zoomorphology" pageId="5" pageNumber="6" pagination="155 - 166" part="99" refId="ref7436" refString="43. Buffrenil, Vde Morphogenesis of bone ornamentation in extant and extinct crocodilians. Zoomorphology 99, 155 - 166 (1982)." title="Vde Morphogenesis of bone ornamentation in extant and extinct crocodilians" type="journal article" year="1982">43</bibRefCitation>
</superScript>
. On one hand, the bone dimensions indicate a small-sized animal (corresponding to a juvenile crocodilian without such developed ornamentation). On the other hand, crocodylian osteoderms are flat without any processes, unlike the studied bone. (4)
<taxonomicName box="[1322,1426,323,345]" class="Reptilia" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="6" phylum="Chordata" rank="order">Testudines</taxonomicName>
; shell plates of several turtles, such as
<emphasis box="[719,950,350,372]" italics="true" pageId="5" pageNumber="6">
<taxonomicName authority=", Allaeochelys" authorityName="Allaeochelys" box="[719,919,350,372]" class="Reptilia" family="Trionychidae" genus="Trionyx" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Trionyx, Allaeochelys</taxonomicName>
etc
</emphasis>
.
<bibRefCitation author="Scheyer, T. M. &amp; Sander, P. M. &amp; Joyce, W. G. &amp; Bohme, W. &amp; Witzel, U." box="[956,971,346,360]" journalOrPublisher="Org. Divers. Evol." pageId="5" pageNumber="6" pagination="136 - 144" part="7" refId="ref7085" refString="35. Scheyer, T. M., Sander, P. M., Joyce, W. G., Bohme, W. &amp; Witzel, U. A plywood structure in the shell of fossil and living soft-shelled turtles (Trionychidae) and its evolutionary implications. Org. Divers. Evol. 7, 136 - 144 (2007)." title="A plywood structure in the shell of fossil and living soft-shelled turtles (Trionychidae) and its evolutionary implications" type="journal article" year="2007">
<superScript attach="right" box="[956,971,346,360]" fontSize="6" pageId="5" pageNumber="6">35</superScript>
</bibRefCitation>
, are also covered by ornamentation. The ornamentation is characterized by larger and closely arranged pits, which are not always clearly delimited from each other (see Scheyer
<bibRefCitation author="Scheyer, T. M. &amp; Sander, P. M. &amp; Joyce, W. G. &amp; Bohme, W. &amp; Witzel, U." box="[533,548,400,414]" journalOrPublisher="Org. Divers. Evol." pageId="5" pageNumber="6" pagination="136 - 144" part="7" refId="ref7085" refString="35. Scheyer, T. M., Sander, P. M., Joyce, W. G., Bohme, W. &amp; Witzel, U. A plywood structure in the shell of fossil and living soft-shelled turtles (Trionychidae) and its evolutionary implications. Org. Divers. Evol. 7, 136 - 144 (2007)." title="A plywood structure in the shell of fossil and living soft-shelled turtles (Trionychidae) and its evolutionary implications" type="journal article" year="2007">
<superScript attach="left" box="[533,548,400,414]" fontSize="6" pageId="5" pageNumber="6">35</superScript>
</bibRefCitation>
:
<figureCitation box="[559,622,403,425]" captionStart="Figure 1" captionStartId="1.[415,480,1168,1190]" captionTargetBox="[417,1469,126,1132]" captionTargetId="figure@1.[415,1471,124,1133]" captionTargetPageId="1" captionText="Figure 1. Maps and stratigraphic column of the studied area and succession. (a) Map of Antarctica showing the location of the study area.(b) Map of the northern Antarctic Peninsula showing the location of Seymour Island. (c) Geological sketch map of Seymour Island, showing the position of the fossil locality (asterisk). (d) Stratigraphic column of the La Meseta Formation on Seymour Island (from11) showing the position of the fossil locality (asterisk).Redrawn from13." figureDoi="http://doi.org/10.5281/zenodo.3766729" httpUri="https://zenodo.org/record/3766729/files/figure.png" pageId="5" pageNumber="6">Fig. 1a</figureCitation>
). (5) Lacertilia; lizards also have skull bones and osteoderms covered with ornamentation patterns
<bibRefCitation author="Estes, R." box="[493,508,426,440]" journalOrPublisher="Gustav Fischer" pageId="5" pageNumber="6" refId="ref7461" refString="44. Estes, R. Sauria terrestria, Amphisbaenia 1 - 249 (Gustav Fischer, 1983)." title="Sauria terrestria, Amphisbaenia 1" type="book" year="1983">
<superScript attach="left" box="[493,508,426,440]" fontSize="6" pageId="5" pageNumber="6">44</superScript>
</bibRefCitation>
. They all are characterized by a network of spines, grooves, ridges
<bibRefCitation author="Cernansky, A. &amp; Auge, M. L." box="[1132,1147,426,440]" journalOrPublisher="Palaeontology" pageId="5" pageNumber="6" pagination="79 - 94" part="56" refId="ref7481" refString="45. Cernansky, A. &amp; Auge, M. L. New species of the genus Plesiolacerta (Squamata: Lacertidae) from the upper Oligocene (MP 28) of Southern Germany and a revision of the type species Plesiolacerta lydekkeri. Palaeontology 56, 79 - 94 (2013)." title="New species of the genus Plesiolacerta (Squamata: Lacertidae) from the upper Oligocene (MP 28) of Southern Germany and a revision of the type species Plesiolacerta lydekkeri" type="journal article" year="2013">
<superScript attach="left" box="[1132,1147,426,440]" fontSize="6" pageId="5" pageNumber="6">45</superScript>
</bibRefCitation>
and protuberances
<bibRefCitation author="Cicimurri, D. J. &amp; Knight, J. L. &amp; Self-Trail, J. M. &amp; Ebersole, S. M." box="[1330,1345,426,440]" journalOrPublisher="J. Paleontol." pageId="5" pageNumber="6" pagination="147 - 153" part="90" refId="ref7536" refString="46. Cicimurri, D. J., Knight, J. L., Self-Trail, J. M. &amp; Ebersole, S. M. Late Paleocene glyptosaur (Reptilia: Anguidae) osteoderms from South Carolina, USA. J. Paleontol. 90, 147 - 153 (2016)." title="Late Paleocene glyptosaur (Reptilia: Anguidae) osteoderms from South Carolina, USA" type="journal article" year="2016">
<superScript attach="left" box="[1330,1345,426,440]" fontSize="6" pageId="5" pageNumber="6">46</superScript>
</bibRefCitation>
, which differs from the morphology on NRM-PZ B281.
</paragraph>
<paragraph blockId="5.[415,1480,136,959]" pageId="5" pageNumber="6">
The ornamentation pattern found in NRM-PZ B281 is comparable to that of some frog genera, i.e.
<taxonomicName box="[415,586,510,531]" class="Reptilia" genus="Thaumastosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[415,586,510,531]" italics="true" pageId="5" pageNumber="6">Thaumastosaurus</emphasis>
</taxonomicName>
<bibRefCitation author="Vasilyan, D." box="[587,602,506,520]" journalOrPublisher="PeerJ" pageId="5" pageNumber="6" part="6" refId="ref7593" refString="47. Vasilyan, D. Eocene Western European endemic genus Thaumastosaurus: New insights into the question &quot; Are the Ranidae known prior to the Oligocene? &quot;. PeerJ 6, https: // doi. org / 10.7717 / peerj. 5511 (2018)." title="Eocene Western European endemic genus Thaumastosaurus: New insights into the question &quot; Are the Ranidae known prior to the Oligocene?" type="journal volume" year="2018">
<superScript attach="right" box="[587,602,506,520]" fontSize="6" pageId="5" pageNumber="6">47</superScript>
</bibRefCitation>
,
<taxonomicName box="[614,716,510,531]" class="Amphibia" family="Ceratophryidae" genus="Beelzebufo" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[614,716,510,531]" italics="true" pageId="5" pageNumber="6">Beelzebufo</emphasis>
</taxonomicName>
<bibRefCitation author="Gomez, R. O. &amp; Baez, A. M. &amp; Muzzopappa, P." box="[717,732,506,520]" journalOrPublisher="J. Vertebr. Paleontol." pageId="5" pageNumber="6" pagination="50 - 59" part="31" refId="ref7642" refString="48. Gomez, R. O., Baez, A. M. &amp; Muzzopappa, P. A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina. J. Vertebr. Paleontol. 31, 50 - 59 (2011)." title="A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina" type="journal article" year="2011">
<superScript attach="right" box="[717,732,506,520]" fontSize="6" pageId="5" pageNumber="6">48</superScript>
</bibRefCitation>
,
<taxonomicName box="[744,915,510,531]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[744,915,510,531]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
<bibRefCitation author="Agnolin, F." box="[915,922,506,520]" journalOrPublisher="Stud. Geol. Salamanticensia" pageId="5" pageNumber="6" pagination="129 - 178" part="48" refId="ref5798" refString="9. Agnolin, F. A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position. Stud. Geol. Salamanticensia 48, 129 - 178 (2012)." title="A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position" type="journal article" year="2012">
<superScript attach="left" box="[915,922,506,520]" fontSize="6" pageId="5" pageNumber="6">9</superScript>
</bibRefCitation>
and
<taxonomicName authorityName="Baez &amp; Peri" authorityYear="1989" box="[972,1127,510,531]" class="Amphibia" family="Ceratophryidae" genus="Baurubatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[972,1127,510,531]" italics="true" pageId="5" pageNumber="6">Baurubatrachus</emphasis>
</taxonomicName>
<bibRefCitation author="Baez, A. M. &amp; Gomez, R. O." box="[1127,1142,506,520]" journalOrPublisher="J. Syst. Palaeontol." pageId="5" pageNumber="6" pagination="279 - 308" part="16" refId="ref7697" refString="49. Baez, A. M. &amp; Gomez, R. O. Dealing with homoplasy: osteology and phylogenetic relationships of the bizarre neobatrachian frog Baurubatrachus pricei from the Upper Cretaceous of Brazil. J. Syst. Palaeontol. 16, 279 - 308 (2018)." title="Dealing with homoplasy: osteology and phylogenetic relationships of the bizarre neobatrachian frog Baurubatrachus pricei from the Upper Cretaceous of Brazil" type="journal article" year="2018">
<superScript attach="left" box="[1127,1142,506,520]" fontSize="6" pageId="5" pageNumber="6">49</superScript>
</bibRefCitation>
, but only the last three genera are Gondwanan forms and, thus, considered for comparison herein.
<taxonomicName box="[1023,1121,537,558]" class="Amphibia" family="Ceratophryidae" genus="Beelzebufo" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1023,1121,537,558]" italics="true" pageId="5" pageNumber="6">Beelzebufo</emphasis>
</taxonomicName>
is a very large form and the ornamentation pattern is present both on skull bones and on vertebrae
<bibRefCitation author="Evans, S. E. &amp; Groenke, J. R. &amp; Jones, M. E. H. &amp; Turner, A. H. &amp; Krause, D. W." box="[1011,1026,560,574]" journalOrPublisher="Plos One" pageId="5" pageNumber="6" pagination="87236" part="9" refId="ref7749" refString="50. Evans, S. E., Groenke, J. R., Jones, M. E. H., Turner, A. H. &amp; Krause, D. W. New material of Beelzebufo, a hyperossified frog (Amphibia: Anura) from the Late Cretaceous of Madagascar. Plos One 9, e 87236, https: // doi. org / 10.1371 / journal. pone. 0087236 (2014)." title="New material of Beelzebufo, a hyperossified frog (Amphibia: Anura) from the Late Cretaceous of Madagascar" type="journal article" year="2014">
<superScript attach="left" box="[1011,1026,560,574]" fontSize="6" pageId="5" pageNumber="6">50</superScript>
</bibRefCitation>
.
<taxonomicName box="[1038,1206,563,584]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1038,1206,563,584]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
<bibRefCitation author="Agnolin, F." box="[1206,1213,560,574]" journalOrPublisher="Stud. Geol. Salamanticensia" pageId="5" pageNumber="6" pagination="129 - 178" part="48" refId="ref5798" refString="9. Agnolin, F. A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position. Stud. Geol. Salamanticensia 48, 129 - 178 (2012)." title="A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position" type="journal article" year="2012">
<superScript attach="left" box="[1206,1213,560,574]" fontSize="6" pageId="5" pageNumber="6">9</superScript>
</bibRefCitation>
and
<taxonomicName authorityName="Baez &amp; Peri" authorityYear="1989" box="[1261,1414,563,584]" class="Amphibia" family="Ceratophryidae" genus="Baurubatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1261,1414,563,584]" italics="true" pageId="5" pageNumber="6">Baurubatrachus</emphasis>
</taxonomicName>
<bibRefCitation author="Baez, A. M. &amp; Gomez, R. O." box="[1414,1429,560,574]" journalOrPublisher="J. Syst. Palaeontol." pageId="5" pageNumber="6" pagination="279 - 308" part="16" refId="ref7697" refString="49. Baez, A. M. &amp; Gomez, R. O. Dealing with homoplasy: osteology and phylogenetic relationships of the bizarre neobatrachian frog Baurubatrachus pricei from the Upper Cretaceous of Brazil. J. Syst. Palaeontol. 16, 279 - 308 (2018)." title="Dealing with homoplasy: osteology and phylogenetic relationships of the bizarre neobatrachian frog Baurubatrachus pricei from the Upper Cretaceous of Brazil" type="journal article" year="2018">
<superScript attach="left" box="[1414,1429,560,574]" fontSize="6" pageId="5" pageNumber="6">49</superScript>
</bibRefCitation>
have very similar ornamentation patterns on the surfaces of hyperossified skull bones, comparable to our specimen. A recent phylogenetic analysis
<bibRefCitation author="Baez, A. M. &amp; Gomez, R. O." box="[682,697,613,627]" journalOrPublisher="J. Syst. Palaeontol." pageId="5" pageNumber="6" pagination="279 - 308" part="16" refId="ref7697" refString="49. Baez, A. M. &amp; Gomez, R. O. Dealing with homoplasy: osteology and phylogenetic relationships of the bizarre neobatrachian frog Baurubatrachus pricei from the Upper Cretaceous of Brazil. J. Syst. Palaeontol. 16, 279 - 308 (2018)." title="Dealing with homoplasy: osteology and phylogenetic relationships of the bizarre neobatrachian frog Baurubatrachus pricei from the Upper Cretaceous of Brazil" type="journal article" year="2018">
<superScript attach="left" box="[682,697,613,627]" fontSize="6" pageId="5" pageNumber="6">49</superScript>
</bibRefCitation>
placed the Late Cretaceous
<taxonomicName authorityName="Baez &amp; Peri" authorityYear="1989" box="[962,1112,617,638]" class="Amphibia" family="Ceratophryidae" genus="Baurubatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[962,1112,617,638]" italics="true" pageId="5" pageNumber="6">Baurubatrachus</emphasis>
</taxonomicName>
within both Recent calyptocephalellid genera
<taxonomicName box="[484,649,643,664]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[484,649,643,664]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Schmidt" authorityYear="1952" box="[695,811,643,664]" class="Amphibia" family="Calyptocephalellidae" genus="Telmatobufo" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[695,811,643,664]" italics="true" pageId="5" pageNumber="6">Telmatobufo</emphasis>
</taxonomicName>
. Though Muzzopappa and Báez
<bibRefCitation author="Muzzopappa, P. &amp; Baez, A. M." box="[1116,1131,640,654]" journalOrPublisher="Ameghiniana" pageId="5" pageNumber="6" pagination="113 - 125" part="46" refId="ref5842" refString="10. Muzzopappa, P. &amp; Baez, A. M. Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus. Ameghiniana 46, 113 - 125 (2009)." title="Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus" type="journal article" year="2009">
<superScript attach="left" box="[1116,1131,640,654]" fontSize="6" pageId="5" pageNumber="6">10</superScript>
</bibRefCitation>
mention that both
<taxonomicName box="[1314,1479,643,664]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1314,1479,643,664]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Schmidt" authorityYear="1952" box="[457,575,670,691]" class="Amphibia" family="Calyptocephalellidae" genus="Telmatobufo" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[457,575,670,691]" italics="true" pageId="5" pageNumber="6">Telmatobufo</emphasis>
</taxonomicName>
are characterized by a heavily ossified neurocranium, we can confirm this only for the former genus (
<figureCitation box="[482,558,696,718]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="5" pageNumber="6">Fig. 4a,f</figureCitation>
). Within
<taxonomicName box="[650,814,697,718]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[650,814,697,718]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
the ornamentation pattern on skull bones is variable. In
<taxonomicName box="[1344,1459,697,718]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="conquella">
<emphasis box="[1344,1459,697,718]" italics="true" pageId="5" pageNumber="6">C. conquella</emphasis>
</taxonomicName>
<bibRefCitation author="Muzzopappa, P. &amp; Baez, A. M." box="[1459,1474,693,707]" journalOrPublisher="Ameghiniana" pageId="5" pageNumber="6" pagination="113 - 125" part="46" refId="ref5842" refString="10. Muzzopappa, P. &amp; Baez, A. M. Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus. Ameghiniana 46, 113 - 125 (2009)." title="Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus" type="journal article" year="2009">
<superScript attach="left" box="[1459,1474,693,707]" fontSize="6" pageId="5" pageNumber="6">10</superScript>
</bibRefCitation>
, it is built either by network of pits in small individuals, or tuberculated ornamentation in adults. In
<taxonomicName box="[1354,1431,724,745]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="satan">
<emphasis box="[1354,1431,724,745]" italics="true" pageId="5" pageNumber="6">C. satan</emphasis>
</taxonomicName>
<bibRefCitation author="Agnolin, F." box="[1431,1438,720,734]" journalOrPublisher="Stud. Geol. Salamanticensia" pageId="5" pageNumber="6" pagination="129 - 178" part="48" refId="ref5798" refString="9. Agnolin, F. A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position. Stud. Geol. Salamanticensia 48, 129 - 178 (2012)." title="A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position" type="journal article" year="2012">
<superScript attach="left" box="[1431,1438,720,734]" fontSize="6" pageId="5" pageNumber="6">9</superScript>
</bibRefCitation>
and
<taxonomicName box="[415,587,750,772]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="casamayorensis">
<emphasis box="[415,587,750,772]" italics="true" pageId="5" pageNumber="6">C. casamayorensis</emphasis>
</taxonomicName>
<bibRefCitation author="Schaeffer, B." box="[587,602,746,760]" journalOrPublisher="B. Am. Mus. Nat. Hist." pageId="5" pageNumber="6" pagination="41 - 68" part="93" refId="ref7832" refString="51. Schaeffer, B. Anurans from the early Tertiary of Patagonia. B. Am. Mus. Nat. Hist. 93, 41 - 68 (1949)." title="Anurans from the early Tertiary of Patagonia" type="journal article" year="1949">
<superScript attach="left" box="[587,602,746,760]" fontSize="6" pageId="5" pageNumber="6">51</superScript>
</bibRefCitation>
, ornamented skull bones are slightly larger than NRM-PZ B281 but they have a similar pattern built of pits.
<taxonomicName box="[574,728,777,798]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="pichifleufensis">
<emphasis box="[574,728,777,798]" italics="true" pageId="5" pageNumber="6">C. pichifleufensis</emphasis>
</taxonomicName>
<bibRefCitation author="Gomez, R. O. &amp; Baez, A. M. &amp; Muzzopappa, P." box="[728,743,773,787]" journalOrPublisher="J. Vertebr. Paleontol." pageId="5" pageNumber="6" pagination="50 - 59" part="31" refId="ref7642" refString="48. Gomez, R. O., Baez, A. M. &amp; Muzzopappa, P. A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina. J. Vertebr. Paleontol. 31, 50 - 59 (2011)." title="A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina" type="journal article" year="2011">
<superScript attach="left" box="[728,743,773,787]" fontSize="6" pageId="5" pageNumber="6">48</superScript>
</bibRefCitation>
is known by larger individuals which show similar ornamentation patterns but with larger pits. In comparison to these species, the Antarctic frog displays an ornamentation most similar to that of
<taxonomicName box="[438,514,830,852]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="satan">
<emphasis box="[438,514,830,852]" italics="true" pageId="5" pageNumber="6">C. satan</emphasis>
</taxonomicName>
<bibRefCitation author="Agnolin, F." box="[514,521,826,840]" journalOrPublisher="Stud. Geol. Salamanticensia" pageId="5" pageNumber="6" pagination="129 - 178" part="48" refId="ref5798" refString="9. Agnolin, F. A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position. Stud. Geol. Salamanticensia 48, 129 - 178 (2012)." title="A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position" type="journal article" year="2012">
<superScript attach="left" box="[514,521,826,840]" fontSize="6" pageId="5" pageNumber="6">9</superScript>
</bibRefCitation>
and
<taxonomicName box="[566,736,830,852]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="casamayorensis">
<emphasis box="[566,736,830,852]" italics="true" pageId="5" pageNumber="6">C. casamayorensis</emphasis>
</taxonomicName>
<bibRefCitation author="Gomez, R. O. &amp; Baez, A. M. &amp; Muzzopappa, P." box="[735,750,826,840]" journalOrPublisher="J. Vertebr. Paleontol." pageId="5" pageNumber="6" pagination="50 - 59" part="31" refId="ref7642" refString="48. Gomez, R. O., Baez, A. M. &amp; Muzzopappa, P. A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina. J. Vertebr. Paleontol. 31, 50 - 59 (2011)." title="A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina" type="journal article" year="2011">
<superScript attach="left" box="[735,750,826,840]" fontSize="6" pageId="5" pageNumber="6">48</superScript>
</bibRefCitation>
. Taking into account our comparison, we conclude that the ornamented bone fragment NRM-PZ B281 represents a skull bone (most probably a nasal) of a small-sized
<taxonomicName box="[1284,1453,857,878]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1284,1453,857,878]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
or
<taxonomicName authorityName="Baez &amp; Peri" authorityYear="1989" box="[415,564,883,904]" class="Amphibia" family="Ceratophryidae" genus="Baurubatrachus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[415,564,883,904]" italics="true" pageId="5" pageNumber="6">Baurubatrachus</emphasis>
</taxonomicName>
. Given the presence of a small
<taxonomicName box="[851,1014,883,904]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[851,1014,883,904]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
as indicated by the ilium in the same, only few m
<superScript attach="left" box="[1472,1479,880,894]" fontSize="6" pageId="5" pageNumber="6">2</superScript>
measuring outcrop, it is most likely that specimen NRM-PZ B281 belongs to the same genus. A comparable record of an ilium and ornamented bones referable to the genus
<taxonomicName box="[1022,1187,937,958]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1022,1187,937,958]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
has been mentioned in Báez
<bibRefCitation author="Baez, A. M. The" box="[1459,1474,933,947]" journalOrPublisher="Museo Argentino de Sciencias Naturales Bernadino Rivadavia" pageId="5" pageNumber="6" refId="ref7865" refString="52. Baez, A. M. The Late Cretaceous Fauna of Los Alamitos, Patagonia, Argentina (ed. Bonaparte J. F.) 121 - 130 (Museo Argentino de Sciencias Naturales Bernadino Rivadavia, 1987)." title="Late Cretaceous Fauna of Los Alamitos, Patagonia, Argentina (ed. Bonaparte J. F.) 121" type="book" year="1987">
<superScript attach="right" box="[1459,1474,933,947]" fontSize="6" pageId="5" pageNumber="6">52</superScript>
</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="5" pageNumber="6" type="discussion">
<paragraph blockId="5.[415,1480,985,1972]" box="[415,549,985,1011]" pageId="5" pageNumber="6">
<heading bold="true" box="[415,549,985,1011]" fontSize="11" level="1" pageId="5" pageNumber="6" reason="0">
<emphasis bold="true" box="[415,549,985,1011]" pageId="5" pageNumber="6">Discussion</emphasis>
</heading>
</paragraph>
<paragraph blockId="5.[415,1480,985,1972]" pageId="5" pageNumber="6">
Among Recent amphibians, the frogs (
<taxonomicName authorityName="Fischer von Waldheim" authorityYear="1813" box="[803,870,1017,1039]" class="Magnoliopsida" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Tracheophyta" rank="order">Anura</taxonomicName>
) have the widest distribution, covering all continents except Antarctica, where the conditions have been uninhabitable for over tens of millions of years. Contrary to all other continents, no traces of any extant amphibian group, all of which belong to the lissamphibian clade, have been documented from Antarctica. This paper presents the first record of a lissamphibian in Antarctica, with Eocene fossils referable to the order
<taxonomicName authorityName="Fischer von Waldheim" authorityYear="1813" box="[681,744,1124,1145]" class="Magnoliopsida" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Tracheophyta" rank="order">Anura</taxonomicName>
, and most likely to the australobatrachian genus
<taxonomicName box="[1203,1368,1124,1145]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[1203,1368,1124,1145]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
. The family
<taxonomicName authorityName="Reig" authorityYear="1960" box="[415,616,1150,1172]" class="Amphibia" family="Calyptocephalellidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Calyptocephalellidae</taxonomicName>
belongs to neobatrachian frogs and is exclusively known from South America
<superScript attach="left" box="[1360,1393,1147,1161]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Otero, R. A. &amp; Jimenez-Huidobro, P. &amp; Soto-Acuna, S. &amp; Yury-Yanez, R. E." box="[1360,1375,1147,1161]" journalOrPublisher="J. S. Am. Earth Sci." pageId="5" pageNumber="6" pagination="133 - 140" part="55" refId="ref7908" refString="53. Otero, R. A., Jimenez-Huidobro, P., Soto-Acuna, S. &amp; Yury-Yanez, R. E. Evidence of a giant helmeted frog (Australobatrachia, Calyptocephalellidae) from Eocene levels of the Magallanes Basin, southernmost Chile. J. S. Am. Earth Sci. 55, 133 - 140 (2014)." title="Evidence of a giant helmeted frog (Australobatrachia, Calyptocephalellidae) from Eocene levels of the Magallanes Basin, southernmost Chile" type="journal article" year="2014">53</bibRefCitation>
,
<bibRefCitation author="Vitt, L. J. &amp; Caldwell, J. P." box="[1378,1393,1147,1161]" journalOrPublisher="An introductory biology of amphibians and reptiles 1 - 776 (Elsevier Academic Press, Amsterdam" pageId="5" pageNumber="6" refId="ref7973" refString="54. Vitt, L. J. &amp; Caldwell, J. P. Herpetology. An introductory biology of amphibians and reptiles 1 - 776 (Elsevier Academic Press, Amsterdam, 2013)." title="Herpetology" type="book" year="2013">54</bibRefCitation>
</superScript>
. The five extant species, including the monospecific genus
<taxonomicName box="[884,1049,1177,1198]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[884,1049,1177,1198]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
with hyperossified skull bones, are restricted to the Chilenean Andes
<bibRefCitation author="Vitt, L. J. &amp; Caldwell, J. P." box="[647,662,1200,1214]" journalOrPublisher="An introductory biology of amphibians and reptiles 1 - 776 (Elsevier Academic Press, Amsterdam" pageId="5" pageNumber="6" refId="ref7973" refString="54. Vitt, L. J. &amp; Caldwell, J. P. Herpetology. An introductory biology of amphibians and reptiles 1 - 776 (Elsevier Academic Press, Amsterdam, 2013)." title="Herpetology" type="book" year="2013">
<superScript attach="left" box="[647,662,1200,1214]" fontSize="6" pageId="5" pageNumber="6">54</superScript>
</bibRefCitation>
while most fossil representatives are known from Argentine Patagonia
<superScript attach="left" box="[1358,1403,1200,1214]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Agnolin, F." box="[1358,1365,1200,1214]" journalOrPublisher="Stud. Geol. Salamanticensia" pageId="5" pageNumber="6" pagination="129 - 178" part="48" refId="ref5798" refString="9. Agnolin, F. A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position. Stud. Geol. Salamanticensia 48, 129 - 178 (2012)." title="A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position" type="journal article" year="2012">9</bibRefCitation>
,
<bibRefCitation author="Gomez, R. O. &amp; Baez, A. M. &amp; Muzzopappa, P." box="[1369,1384,1200,1214]" journalOrPublisher="J. Vertebr. Paleontol." pageId="5" pageNumber="6" pagination="50 - 59" part="31" refId="ref7642" refString="48. Gomez, R. O., Baez, A. M. &amp; Muzzopappa, P. A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina. J. Vertebr. Paleontol. 31, 50 - 59 (2011)." title="A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina" type="journal article" year="2011">48</bibRefCitation>
,
<bibRefCitation author="Otero, R. A. &amp; Jimenez-Huidobro, P. &amp; Soto-Acuna, S. &amp; Yury-Yanez, R. E." box="[1388,1403,1200,1214]" journalOrPublisher="J. S. Am. Earth Sci." pageId="5" pageNumber="6" pagination="133 - 140" part="55" refId="ref7908" refString="53. Otero, R. A., Jimenez-Huidobro, P., Soto-Acuna, S. &amp; Yury-Yanez, R. E. Evidence of a giant helmeted frog (Australobatrachia, Calyptocephalellidae) from Eocene levels of the Magallanes Basin, southernmost Chile. J. S. Am. Earth Sci. 55, 133 - 140 (2014)." title="Evidence of a giant helmeted frog (Australobatrachia, Calyptocephalellidae) from Eocene levels of the Magallanes Basin, southernmost Chile" type="journal article" year="2014">53</bibRefCitation>
</superScript>
. Today,
<taxonomicName box="[415,582,1230,1251]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[415,582,1230,1251]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
inhabits lowland areas of central Chile (upper elevation limit 500 m) east of the Andes within temperate and humid climates, between latitudes 3043°S. It has an aquatic or semiaquatic lifestyle and populates standing or slow flowing water bodies (lakes, ponds, streams) in the Valdivian temperate
<taxonomicName box="[1258,1364,1284,1305]" class="Magnoliopsida" family="Nothofagaceae" genus="Nothofagus" kingdom="Plantae" order="Fagales" pageId="5" pageNumber="6" phylum="Tracheophyta" rank="genus">
<emphasis box="[1258,1364,1284,1305]" italics="true" pageId="5" pageNumber="6">Nothofagus</emphasis>
</taxonomicName>
forests
<superScript attach="right" box="[1432,1470,1280,1294]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Vitt, L. J. &amp; Caldwell, J. P." box="[1432,1447,1280,1294]" journalOrPublisher="An introductory biology of amphibians and reptiles 1 - 776 (Elsevier Academic Press, Amsterdam" pageId="5" pageNumber="6" refId="ref7973" refString="54. Vitt, L. J. &amp; Caldwell, J. P. Herpetology. An introductory biology of amphibians and reptiles 1 - 776 (Elsevier Academic Press, Amsterdam, 2013)." title="Herpetology" type="book" year="2013">54</bibRefCitation>
<bibRefCitation author="Cei, J. M." box="[1455,1470,1280,1294]" journalOrPublisher="Universidad de Chile, Santiago" pageId="5" pageNumber="6" refId="ref8042" refString="56. Cei, J. M. Batracios de Chile 1 - 128 (Universidad de Chile, Santiago, 1962)." title="Batracios de Chile 1" type="book" year="1962">56</bibRefCitation>
</superScript>
.
</paragraph>
<paragraph blockId="5.[415,1480,985,1972]" pageId="5" pageNumber="6">
The oldest fossils referable to
<taxonomicName box="[747,920,1310,1331]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[747,920,1310,1331]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
are known from the Upper Cretaceous of Argentina
<superScript attach="left" box="[1446,1474,1307,1321]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Agnolin, F." box="[1446,1453,1307,1321]" journalOrPublisher="Stud. Geol. Salamanticensia" pageId="5" pageNumber="6" pagination="129 - 178" part="48" refId="ref5798" refString="9. Agnolin, F. A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position. Stud. Geol. Salamanticensia 48, 129 - 178 (2012)." title="A new Calyptocephalellidae (Anura, Neobatrachia) from the Upper Cretaceous of Patagonia, Argentina, with comments on its systematic position" type="journal article" year="2012">9</bibRefCitation>
,
<bibRefCitation author="Baez, A. M. The" box="[1458,1474,1307,1321]" journalOrPublisher="Museo Argentino de Sciencias Naturales Bernadino Rivadavia" pageId="5" pageNumber="6" refId="ref7865" refString="52. Baez, A. M. The Late Cretaceous Fauna of Los Alamitos, Patagonia, Argentina (ed. Bonaparte J. F.) 121 - 130 (Museo Argentino de Sciencias Naturales Bernadino Rivadavia, 1987)." title="Late Cretaceous Fauna of Los Alamitos, Patagonia, Argentina (ed. Bonaparte J. F.) 121" type="book" year="1987">52</bibRefCitation>
</superScript>
. During the Paleoceneterminal early Miocene, their geographic range was restricted to Patagonia east of the Andes
<superScript attach="left" box="[477,548,1360,1374]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Gomez, R. O. &amp; Baez, A. M. &amp; Muzzopappa, P." box="[477,492,1360,1374]" journalOrPublisher="J. Vertebr. Paleontol." pageId="5" pageNumber="6" pagination="50 - 59" part="31" refId="ref7642" refString="48. Gomez, R. O., Baez, A. M. &amp; Muzzopappa, P. A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina. J. Vertebr. Paleontol. 31, 50 - 59 (2011)." title="A new helmeted frog (Anura: Calyptocephalellidae) from an Eocene subtropical lake in northwestern Patagonia, Argentina" type="journal article" year="2011">48</bibRefCitation>
,
<bibRefCitation author="Schaeffer, B." box="[496,511,1360,1374]" journalOrPublisher="B. Am. Mus. Nat. Hist." pageId="5" pageNumber="6" pagination="41 - 68" part="93" refId="ref7832" refString="51. Schaeffer, B. Anurans from the early Tertiary of Patagonia. B. Am. Mus. Nat. Hist. 93, 41 - 68 (1949)." title="Anurans from the early Tertiary of Patagonia" type="journal article" year="1949">51</bibRefCitation>
,
<bibRefCitation author="Otero, R. A. &amp; Jimenez-Huidobro, P. &amp; Soto-Acuna, S. &amp; Yury-Yanez, R. E." box="[515,530,1360,1374]" journalOrPublisher="J. S. Am. Earth Sci." pageId="5" pageNumber="6" pagination="133 - 140" part="55" refId="ref7908" refString="53. Otero, R. A., Jimenez-Huidobro, P., Soto-Acuna, S. &amp; Yury-Yanez, R. E. Evidence of a giant helmeted frog (Australobatrachia, Calyptocephalellidae) from Eocene levels of the Magallanes Basin, southernmost Chile. J. S. Am. Earth Sci. 55, 133 - 140 (2014)." title="Evidence of a giant helmeted frog (Australobatrachia, Calyptocephalellidae) from Eocene levels of the Magallanes Basin, southernmost Chile" type="journal article" year="2014">53</bibRefCitation>
,
<bibRefCitation author="Nicoli, L. &amp; Muzzopappa, P. &amp; Faivovich, J." box="[533,548,1360,1374]" journalOrPublisher="Alcheringa" pageId="5" pageNumber="6" pagination="153 - 160" part="40" refId="ref8066" refString="57. Nicoli, L., Muzzopappa, P. &amp; Faivovich, J. The taxonomic placement of the Miocene Patagonian frog Wawelia gerholdi (Amphibia: Anura). Alcheringa 40, 153 - 160, https: // doi. org / 10.1080 / 03115518.2016.1101998 (2016)." title="The taxonomic placement of the Miocene Patagonian frog Wawelia gerholdi (Amphibia: Anura)" type="journal article" year="2016">57</bibRefCitation>
</superScript>
. Not until the late Pleistocene did they appear west of the Andes, where they have their endemic present-day distribution
<superScript attach="left" box="[646,685,1387,1401]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Vitt, L. J. &amp; Caldwell, J. P." box="[646,661,1387,1401]" journalOrPublisher="An introductory biology of amphibians and reptiles 1 - 776 (Elsevier Academic Press, Amsterdam" pageId="5" pageNumber="6" refId="ref7973" refString="54. Vitt, L. J. &amp; Caldwell, J. P. Herpetology. An introductory biology of amphibians and reptiles 1 - 776 (Elsevier Academic Press, Amsterdam, 2013)." title="Herpetology" type="book" year="2013">54</bibRefCitation>
<bibRefCitation author="Nicoli, L. &amp; Muzzopappa, P. &amp; Faivovich, J." box="[670,685,1387,1401]" journalOrPublisher="Alcheringa" pageId="5" pageNumber="6" pagination="153 - 160" part="40" refId="ref8066" refString="57. Nicoli, L., Muzzopappa, P. &amp; Faivovich, J. The taxonomic placement of the Miocene Patagonian frog Wawelia gerholdi (Amphibia: Anura). Alcheringa 40, 153 - 160, https: // doi. org / 10.1080 / 03115518.2016.1101998 (2016)." title="The taxonomic placement of the Miocene Patagonian frog Wawelia gerholdi (Amphibia: Anura)" type="journal article" year="2016">57</bibRefCitation>
</superScript>
.
</paragraph>
<paragraph blockId="5.[415,1480,985,1972]" pageId="5" pageNumber="6">
The clade Australobatrachia comprises Myobatrachoidea (families
<taxonomicName authorityName="Schlegel" authorityYear="1850" box="[1113,1274,1417,1439]" class="Amphibia" family="Myobatrachidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Myobatrachidae</taxonomicName>
+
<taxonomicName authorityName="Lynch" authorityYear="1969" box="[1306,1479,1417,1439]" class="Amphibia" family="Limnodynastidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Limnodynastidae</taxonomicName>
sensu
<bibRefCitation author="Frost, D. R." box="[471,487,1440,1454]" journalOrPublisher="B. Am. Mus. Nat. Hist." pageId="5" pageNumber="6" pagination="1 - 370" part="297" refId="ref6760" refString="27. Frost, D. R. et al. The amphibian tree of life. B. Am. Mus. Nat. Hist. 297, 1 - 370 (2006)." title="The amphibian tree of life" type="journal article" year="2006">
<superScript attach="right" box="[471,487,1440,1454]" fontSize="6" pageId="5" pageNumber="6">27</superScript>
</bibRefCitation>
), nowadays distributed in Australia and south of New Guinea, and the family
<taxonomicName authorityName="Reig" authorityYear="1960" box="[1271,1479,1444,1466]" class="Amphibia" family="Calyptocephalellidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Calyptocephalellidae</taxonomicName>
(
<taxonomicName authorityName="Cope" authorityYear="1875" box="[422,614,1470,1492]" family="Batrachophrynidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anuri" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Batrachophrynidae</taxonomicName>
sensu
<bibRefCitation author="Frost, D. R." box="[675,690,1467,1481]" journalOrPublisher="B. Am. Mus. Nat. Hist." pageId="5" pageNumber="6" pagination="1 - 370" part="297" refId="ref6760" refString="27. Frost, D. R. et al. The amphibian tree of life. B. Am. Mus. Nat. Hist. 297, 1 - 370 (2006)." title="The amphibian tree of life" type="journal article" year="2006">
<superScript attach="left" box="[675,690,1467,1481]" fontSize="6" pageId="5" pageNumber="6">27</superScript>
</bibRefCitation>
). Australobatrachia are considered as a stem group of the Hyloidea. The earliest myobatrachoid from Australia is at least as old as early Eocene, based on fragmentary ilia that were referred to the basal extant
<taxonomicName authorityName="Boulenger" authorityYear="1882" box="[567,669,1524,1545]" class="Amphibia" family="Limnodynastidae" genus="Lechriodus" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[567,669,1524,1545]" italics="true" pageId="5" pageNumber="6">Lechriodus</emphasis>
</taxonomicName>
<bibRefCitation author="Tyler, M. J. &amp; Godthelp, H." box="[670,685,1520,1534]" journalOrPublisher="T. Roy. Soc. South Aust." pageId="5" pageNumber="6" pagination="187 - 189" part="117" refId="ref8119" refString="58. Tyler, M. J. &amp; Godthelp, H. A new species of Lechriodus Boulenger (Anura: Leptodactylidae) from the Early Eocene of Queensland. T. Roy. Soc. South Aust. 117, 187 - 189 (1993)." title="A new species of Lechriodus Boulenger (Anura: Leptodactylidae) from the Early Eocene of Queensland" type="journal article" year="1993">
<superScript attach="right" box="[670,685,1520,1534]" fontSize="6" pageId="5" pageNumber="6">58</superScript>
</bibRefCitation>
. The split between
<taxonomicName authorityName="Reig" authorityYear="1960" box="[865,1065,1524,1546]" class="Amphibia" family="Calyptocephalellidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Calyptocephalellidae</taxonomicName>
and
<taxonomicName authorityName="Schlegel" authorityYear="1850" box="[1110,1266,1524,1546]" class="Amphibia" family="Myobatrachidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Myobatrachidae</taxonomicName>
(
<taxonomicName authorityName="Reig" authorityYear="1960" box="[1277,1479,1524,1546]" class="Amphibia" family="Calyptocephalellidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Calyptocephalellidae</taxonomicName>
+ Myobatrachoidea sensu
<bibRefCitation author="Frost, D. R." box="[663,678,1547,1561]" journalOrPublisher="B. Am. Mus. Nat. Hist." pageId="5" pageNumber="6" pagination="1 - 370" part="297" refId="ref6760" refString="27. Frost, D. R. et al. The amphibian tree of life. B. Am. Mus. Nat. Hist. 297, 1 - 370 (2006)." title="The amphibian tree of life" type="journal article" year="2006">
<superScript attach="left" box="[663,678,1547,1561]" fontSize="6" pageId="5" pageNumber="6">27</superScript>
</bibRefCitation>
) occurred ~100 Ma (~Early-Late Cretaceous boundary)
<bibRefCitation author="Feng, Y. - J." box="[1211,1226,1547,1561]" journalOrPublisher="P. Natl Acad. Sci. USA" pageId="5" pageNumber="6" pagination="5864 - E 5870" part="114" refId="ref8168" refString="59. Feng, Y. - J. et al. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous-Paleogene boundary. P. Natl Acad. Sci. USA 114, E 5864 - E 5870 (2017)." title="Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous-Paleogene boundary" type="journal article" year="2017">
<superScript attach="none" box="[1211,1226,1547,1561]" fontSize="6" pageId="5" pageNumber="6">59</superScript>
</bibRefCitation>
. Considering the distributions of extant Australobatrachia (
<figureCitation box="[738,791,1577,1599]" captionStart="Figure 5" captionStartId="6.[415,480,960,982]" captionTargetBox="[417,1053,126,922]" captionTargetId="figure@6.[415,1055,124,924]" captionTargetPageId="6" captionText="Figure 5. Eocene palaeogeography of the south polar region with a cladogram of australobatrachid frogs showing their occurrences on the southern continents. The grey color indicates the outlines of the continents during the Eocene, the black colored outline the present-day outlines of the continents. Map redrawn from an original generated using ArcGIS 10.17.1 (www.esri.com) software, based on the Satellite base map layer in google Maps (Map data ©2019 Google).Abbreviations: ANT, Antarctica; AUS, Australia and Tasmania; NG, New Guinea; NZ, New Zealand; SA, South America. The red star indicates the fossil locality on Seymour Island." figureDoi="http://doi.org/10.5281/zenodo.3766737" httpUri="https://zenodo.org/record/3766737/files/figure.png" pageId="5" pageNumber="6">Fig. 5</figureCitation>
), the earliest fossil records
<bibRefCitation author="Muzzopappa, P. &amp; Baez, A. M." box="[1042,1057,1574,1588]" journalOrPublisher="Ameghiniana" pageId="5" pageNumber="6" pagination="113 - 125" part="46" refId="ref5842" refString="10. Muzzopappa, P. &amp; Baez, A. M. Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus. Ameghiniana 46, 113 - 125 (2009)." title="Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus" type="journal article" year="2009">
<superScript attach="left" box="[1042,1057,1574,1588]" fontSize="6" pageId="5" pageNumber="6">10</superScript>
</bibRefCitation>
and the divergence age (from genetic data)
<bibRefCitation author="Feng, Y. - J." box="[1464,1479,1573,1587]" journalOrPublisher="P. Natl Acad. Sci. USA" pageId="5" pageNumber="6" pagination="5864 - E 5870" part="114" refId="ref8168" refString="59. Feng, Y. - J. et al. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous-Paleogene boundary. P. Natl Acad. Sci. USA 114, E 5864 - E 5870 (2017)." title="Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous-Paleogene boundary" type="journal article" year="2017">
<superScript attach="none" box="[1464,1479,1573,1587]" fontSize="6" pageId="5" pageNumber="6">59</superScript>
</bibRefCitation>
of both
<taxonomicName authorityName="Reig" authorityYear="1960" box="[490,692,1604,1626]" class="Amphibia" family="Calyptocephalellidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Calyptocephalellidae</taxonomicName>
and Myobatrachoidea lineages, it is clear that Antarctica had played an important palaeobiogeographic role for Australobatrachia and their consequent dispersal. Because the most recent common ancestor of the clade, including Hyloidae and
<taxonomicName authorityName="Schlegel" authorityYear="1850" box="[905,1062,1657,1679]" class="Amphibia" family="Myobatrachidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Myobatrachidae</taxonomicName>
+
<taxonomicName authorityName="Reig" authorityYear="1960" box="[1092,1296,1657,1679]" class="Amphibia" family="Calyptocephalellidae" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Calyptocephalellidae</taxonomicName>
, occurred in South America, their origin in South America and consequent dispersal from South America to Australia via Antarctica has been suggested
<bibRefCitation author="Feng, Y. - J." box="[596,611,1707,1721]" journalOrPublisher="P. Natl Acad. Sci. USA" pageId="5" pageNumber="6" pagination="5864 - E 5870" part="114" refId="ref8168" refString="59. Feng, Y. - J. et al. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous-Paleogene boundary. P. Natl Acad. Sci. USA 114, E 5864 - E 5870 (2017)." title="Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous-Paleogene boundary" type="journal article" year="2017">
<superScript attach="left" box="[596,611,1707,1721]" fontSize="6" pageId="5" pageNumber="6">59</superScript>
</bibRefCitation>
. Additionally, this suggests one more case of strong faunistic affinities of the continent with South America and Australia
<superScript attach="left" box="[691,747,1733,1748]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Woodburne, M. O. &amp; Zinsmeister, W. J." box="[691,698,1734,1748]" journalOrPublisher="Science" pageId="5" pageNumber="6" pagination="284 - 286" part="218" refId="ref5542" refString="4. Woodburne, M. O. &amp; Zinsmeister, W. J. Fossil land mammal from Antarctica. Science 218, 284 - 286 (1982)." title="Fossil land mammal from Antarctica" type="journal article" year="1982">4</bibRefCitation>
,
<bibRefCitation author="Gelfo, J. N. &amp; Mors, T. &amp; Lorente, M. &amp; Lopez, G. M. &amp; Reguero, M." box="[702,709,1733,1747]" journalOrPublisher="Palaeontology" pageId="5" pageNumber="6" pagination="101 - 110" part="58" refId="ref5623" refString="6. Gelfo, J. N., Mors, T., Lorente, M., Lopez, G. M. &amp; Reguero, M. The oldest mammals from Antarctica, early Eocene of the La Meseta Formation, Seymour Island. Palaeontology 58, 101 - 110 (2015)." title="The oldest mammals from Antarctica, early Eocene of the La Meseta Formation, Seymour Island" type="journal article" year="2015">6</bibRefCitation>
,
<bibRefCitation author="Chornogubsky, L. &amp; Goin, F. J. &amp; Reguero, M." box="[713,728,1733,1747]" journalOrPublisher="Antarct. Sci." pageId="5" pageNumber="6" pagination="285 - 297" part="21" refId="ref6178" refString="16. Chornogubsky, L., Goin, F. J. &amp; Reguero, M. A reassessment of Antarctic polydolopid marsupials (Middle Eocene, La Meseta Formation). Antarct. Sci. 21, 285 - 297 (2009)." title="A reassessment of Antarctic polydolopid marsupials (Middle Eocene, La Meseta Formation)" type="journal article" year="2009">16</bibRefCitation>
,
<bibRefCitation author="Vizcaino, S. F. &amp; Kay, R. F. &amp; Bargo, M. S." box="[732,747,1733,1747]" journalOrPublisher="Press" pageId="5" pageNumber="6" refId="ref8217" refString="60. Vizcaino, S. F., Kay, R. F. &amp; Bargo, M. S. (eds.). Early Miocene paleobiology in Patagonia: High-latitude paleocommunities of the Santa Cruz Formation 1 - 378 (Cambridge Univ. Press, 2012)." title="Early Miocene paleobiology in Patagonia: High-latitude paleocommunities of the Santa Cruz Formation 1 - 378 (Cambridge Univ" type="book" year="2012">60</bibRefCitation>
</superScript>
. So far, Antarctica has been considered as a dispersal route, but not as a probable place of origin. The new fossil finds support the hypothesis
<bibRefCitation author="Muzzopappa, P. &amp; Baez, A. M." box="[1028,1043,1760,1774]" journalOrPublisher="Ameghiniana" pageId="5" pageNumber="6" pagination="113 - 125" part="46" refId="ref5842" refString="10. Muzzopappa, P. &amp; Baez, A. M. Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus. Ameghiniana 46, 113 - 125 (2009)." title="Systematic status of the mid-Tertiary neobatrachian frog Calyptocephalella canqueli from Patagonia (Argentina), with comments on the evolution of the genus" type="journal article" year="2009">
<superScript attach="left" box="[1028,1043,1760,1774]" fontSize="6" pageId="5" pageNumber="6">10</superScript>
</bibRefCitation>
that Antarctica may have acted as a center of diversification for australobatrachians.
</paragraph>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6" type="distribution">
<paragraph blockId="5.[415,1480,985,1972]" pageId="5" pageNumber="6">
The Seymour Island frog reported herein is the first vertebrate indicative of freshwater habitats on the Eocene Antarctic Peninsula, following invertebrate and plant evidence
<superScript attach="left" box="[1025,1059,1840,1854]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Friis, E. M. &amp; Iglesias, A. &amp; Reguero, M. A. &amp; Mors, T." box="[1025,1040,1840,1854]" journalOrPublisher="Plant Syst. Evol." pageId="5" pageNumber="6" pagination="969 - 980" part="181" refId="ref5937" refString="12. Friis, E. M., Iglesias, A., Reguero, M. A. &amp; Mors, T. Notonuphar antarctica, an extinct water lily (Nymphaeales) from the Eocene of Antarctica. Plant Syst. Evol. 181, 969 - 980 (2017)." title="Notonuphar antarctica, an extinct water lily (Nymphaeales) from the Eocene of Antarctica" type="journal article" year="2017">12</bibRefCitation>
,
<bibRefCitation author="Bomfleur, B. &amp; Mors, T. &amp; Ferraguti, M. &amp; Reguero, M. A. &amp; McLoughlin, S." box="[1044,1059,1840,1854]" journalOrPublisher="Biol. Letters" pageId="5" pageNumber="6" pagination="20150431" part="11" refId="ref6224" refString="17. Bomfleur, B., Mors, T., Ferraguti, M., Reguero, M. A. &amp; McLoughlin, S. Fossilized spermatozoa preserved in a 50 - Myr-old annelid cocoon from Antarctica. Biol. Letters 11, 20150431, https: // doi. org / 10.1098 / rsbl. 2015.0431 (2015)." title="Fossilized spermatozoa preserved in a 50 - Myr-old annelid cocoon from Antarctica" type="journal article" year="2015">17</bibRefCitation>
</superScript>
(
<figureCitation box="[1072,1128,1844,1866]" captionStart="Figure 6" captionStartId="7.[415,480,932,954]" captionTargetBox="[416,1053,125,894]" captionTargetId="figure@7.[415,1055,124,896]" captionTargetPageId="7" captionText="Figure 6. Reconstruction of an Eocene pond in the Nothofagus forest of the Antarctic Peninsula with Calyptocephalella, sitting on a leaf of Notonuphar antarctica which was described from the same locality12. Artwork by Pollyanna von Knorring, Swedish Museum of Natural History.Photo credits: Simon Pierre Barrette and José Grau de Puerto Montt, Wikimedia Commons (CC BY-SA 3.0), and Mats Wedin, Swedish Museum of Natural History." figureDoi="http://doi.org/10.5281/zenodo.3766739" httpUri="https://zenodo.org/record/3766739/files/figure.png" pageId="5" pageNumber="6">Fig. 6</figureCitation>
). It is interesting to note that nearly all fossil localities where
<taxonomicName box="[653,819,1870,1891]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[653,819,1870,1891]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
occurs (excepting those, for which fossil plant data are not available) contain evidence of the presence of
<taxonomicName box="[763,871,1897,1918]" class="Magnoliopsida" family="Nothofagaceae" genus="Nothofagus" kingdom="Plantae" order="Fagales" pageId="5" pageNumber="6" phylum="Tracheophyta" rank="genus">
<emphasis box="[763,871,1897,1918]" italics="true" pageId="5" pageNumber="6">Nothofagus</emphasis>
</taxonomicName>
, including Seymour Island
<superScript attach="left" box="[1134,1187,1893,1908]" fontSize="6" pageId="5" pageNumber="6">
<bibRefCitation author="Chornogubsky, L. &amp; Goin, F. J. &amp; Reguero, M." box="[1134,1149,1893,1907]" journalOrPublisher="Antarct. Sci." pageId="5" pageNumber="6" pagination="285 - 297" part="21" refId="ref6178" refString="16. Chornogubsky, L., Goin, F. J. &amp; Reguero, M. A reassessment of Antarctic polydolopid marsupials (Middle Eocene, La Meseta Formation). Antarct. Sci. 21, 285 - 297 (2009)." title="A reassessment of Antarctic polydolopid marsupials (Middle Eocene, La Meseta Formation)" type="journal article" year="2009">16</bibRefCitation>
,
<bibRefCitation author="Vizcaino, S. F. &amp; Kay, R. F. &amp; Bargo, M. S." box="[1153,1168,1893,1907]" journalOrPublisher="Press" pageId="5" pageNumber="6" refId="ref8217" refString="60. Vizcaino, S. F., Kay, R. F. &amp; Bargo, M. S. (eds.). Early Miocene paleobiology in Patagonia: High-latitude paleocommunities of the Santa Cruz Formation 1 - 378 (Cambridge Univ. Press, 2012)." title="Early Miocene paleobiology in Patagonia: High-latitude paleocommunities of the Santa Cruz Formation 1 - 378 (Cambridge Univ" type="book" year="2012">60</bibRefCitation>
,
<bibRefCitation author="Francis, J. E." box="[1172,1187,1893,1907]" journalOrPublisher="National Academies Press" pageId="5" pageNumber="6" refId="ref8269" refString="61. Francis, J. E. et al. Antarctica: A keystone in a changing world; proceedings of the 10 th International Symposium on Antarctic Earth Sciences, Santa Barbara, California (ed. Cooper, A. K. &amp; Barrett, P.) 19 - 27 (National Academies Press, 2008)." title="Antarctica: A keystone in a changing world; proceedings of the 10 th International Symposium on Antarctic Earth Sciences, Santa Barbara, California (ed. Cooper, A. K. &amp; Barrett, P.) 19" type="proceedings" year="2008">61</bibRefCitation>
</superScript>
. The southern extant range of
<taxonomicName box="[415,583,1924,1945]" class="Amphibia" family="Calyptocephalellidae" genus="Calyptocephalella" kingdom="Animalia" order="Anura" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[415,583,1924,1945]" italics="true" pageId="5" pageNumber="6">Calyptocephalella</emphasis>
</taxonomicName>
occurs sympatrically with the microbiotherian marsupial
<taxonomicName authorityName="Thomas" authorityYear="1894" box="[1148,1342,1924,1945]" class="Mammalia" family="Microbiotheriidae" genus="Dromiciops" kingdom="Animalia" order="Microbiotheria" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="gliroides">
<emphasis box="[1148,1342,1924,1945]" italics="true" pageId="5" pageNumber="6">Dromiciops gliroides</emphasis>
</taxonomicName>
(
<figureCitation box="[1354,1424,1924,1946]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="5" pageNumber="6">Fig. S4</figureCitation>
), also known as “Monito del Monte” or “Colocolo Opossum”, a small mammal with an arboreal lifestyle and an endemic
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3766737" ID-Zenodo-Dep="3766737" httpUri="https://zenodo.org/record/3766737/files/figure.png" pageId="6" pageNumber="7" startId="6.[415,480,960,982]" subCaptionStartIDs="6.[799,936,1066,1088]" subCaptionStarts="Abbr" targetBox="[417,1053,126,922]" targetPageId="6">
<paragraph blockId="6.[415,1475,960,1115]" pageId="6" pageNumber="7">
<emphasis bold="true" box="[415,503,960,982]" pageId="6" pageNumber="7">Figure 5.</emphasis>
Eocene palaeogeography of the south polar region with a cladogram of australobatrachid frogs showing their occurrences on the southern continents. The grey color indicates the outlines of the continents during the Eocene, the black colored outline the present-day outlines of the continents. Map redrawn from an original generated using ArcGIS 10.17.1 (www.esri.com) software, based on the Satellite base map layer in google Maps (Map data ©2019 Google). Abbreviations: ANT, Antarctica; AUS, Australia and Tasmania; NG, New Guinea; NZ, New Zealand; SA, South America. The red star indicates the fossil locality on Seymour Island.
</paragraph>
</caption>
<paragraph blockId="6.[415,1480,1197,1596]" pageId="6" pageNumber="7">
distribution in the dense Valdivian
<taxonomicName box="[764,873,1201,1222]" class="Magnoliopsida" family="Nothofagaceae" genus="Nothofagus" kingdom="Plantae" order="Fagales" pageId="6" pageNumber="7" phylum="Tracheophyta" rank="genus">
<emphasis box="[764,873,1201,1222]" italics="true" pageId="6" pageNumber="7">Nothofagus</emphasis>
</taxonomicName>
forests of highland Argentina and Chile
<bibRefCitation author="Nowak, R. M. &amp; Dickman, C. R." box="[1270,1285,1197,1211]" journalOrPublisher="Johns Hopkins University Press" pageId="6" pageNumber="7" refId="ref8331" refString="62. Nowak, R. M. &amp; Dickman, C. R. Walker's marsupials of the world 1 - 226 (Johns Hopkins University Press, 2005)." title="Walker's marsupials of the world 1" type="book" year="2005">
<superScript attach="left" box="[1270,1285,1197,1211]" fontSize="6" pageId="6" pageNumber="7">62</superScript>
</bibRefCitation>
. The climate of this
<taxonomicName box="[415,520,1227,1248]" class="Magnoliopsida" family="Nothofagaceae" genus="Nothofagus" kingdom="Plantae" order="Fagales" pageId="6" pageNumber="7" phylum="Tracheophyta" rank="genus">
<emphasis box="[415,520,1227,1248]" italics="true" pageId="6" pageNumber="7">Nothofagus</emphasis>
</taxonomicName>
forest area with the sympatric occurrences of these two endemic animals shows humid and temperate conditions (for the numerical values, see Methods and
<tableCitation box="[948,1018,1254,1276]" captionStart="Table 2" captionStartId="7.[415,469,1367,1388]" captionText="Table 2. Climatic parameters (MAP, MCMT, MWMT) and the elevation at the climatic stations according to the database of the word bank groups67. Abbreviations: MAP=mean annual precipitation, MCMT=mean coldest month temperature, MWMT=mean warmest month temperature. The location of the climatic stations can be found on Fig. S4." pageId="6" pageNumber="7">Table 2</tableCitation>
,
<figureCitation box="[1029,1097,1254,1276]" captionStart="Figure 4" captionStartId="4.[415,480,1050,1072]" captionTargetBox="[418,1468,125,1012]" captionTargetId="figure@4.[415,1471,124,1015]" captionTargetPageId="4" captionText="Figure 4. 3D models of some skeletal elements of Australobatrachia. (a,f) skull and (be,gj) ilia of Calyptocephalella gayi (ae); Telmatobufo venustus (fh); Myobatrachus gouldii (i); Limnodynastes convexiusculus (j). Collection numbers of each specimen are listed in Table S1." figureDoi="http://doi.org/10.5281/zenodo.3766735" httpUri="https://zenodo.org/record/3766735/files/figure.png" pageId="6" pageNumber="7">Fig. S4</figureCitation>
).
<taxonomicName authorityName="Thomas" authorityYear="1894" box="[1113,1307,1254,1276]" class="Mammalia" family="Microbiotheriidae" genus="Dromiciops" kingdom="Animalia" order="Microbiotheria" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="gliroides">
<emphasis box="[1113,1307,1254,1276]" italics="true" pageId="6" pageNumber="7">Dromiciops gliroides</emphasis>
</taxonomicName>
is the only extant species of the order
<taxonomicName box="[605,751,1280,1302]" class="Mammalia" kingdom="Animalia" order="Microbiotheria" pageId="6" pageNumber="7" phylum="Chordata" rank="order">Microbiotheria</taxonomicName>
and is considered as the only South American representative of the superor- der Australidelphia which otherwise comprises Australian marsupials
<bibRefCitation author="Nilsson, M. A." box="[1076,1091,1304,1318]" journalOrPublisher="Plos biology" pageId="6" pageNumber="7" pagination="1000436" part="8" refId="ref8363" refString="63. Nilsson, M. A. et al. Tracking marsupial evolution using archaic genomic retroposon insertions. Plos biology 8, e 1000436, https: // doi. org / 10.1371 / journal. pbio. 1000436 (2010)." title="Tracking marsupial evolution using archaic genomic retroposon insertions" type="journal article" year="2010">
<superScript attach="left" box="[1076,1091,1304,1318]" fontSize="6" pageId="6" pageNumber="7">63</superScript>
</bibRefCitation>
. From the same small shell-rich lens that produced the frog remains reported herein, the fossil microbiotherian
<taxonomicName box="[1078,1274,1334,1356]" class="Mammalia" family="Woodburnodontidae" genus="Woodburnodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Microbiotheria" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="casei">
<emphasis box="[1078,1274,1334,1356]" italics="true" pageId="6" pageNumber="7">Woodburnodon casei</emphasis>
</taxonomicName>
has been described
<bibRefCitation author="Goin, F." box="[1459,1474,1330,1344]" journalOrPublisher="Rev. Asoc. Paleontol. Argentina" pageId="6" pageNumber="7" pagination="597 - 603" part="64" refId="ref8408" refString="64. Goin, F. et al. New marsupial (Mammalia) from the Eocene of Antarctica, and the origins and affinities of the Microbiotheria. Rev. Asoc. Paleontol. Argentina 64, 597 - 603 (2007)." title="New marsupial (Mammalia) from the Eocene of Antarctica, and the origins and affinities of the Microbiotheria" type="journal article" year="2007">
<superScript attach="right" box="[1459,1474,1330,1344]" fontSize="6" pageId="6" pageNumber="7">64</superScript>
</bibRefCitation>
. Hence, we hypothesize that the climatic conditions for the Antarctic Peninsula during the Bartonian (late middle Eocene) should be comparable with the climate found today in the concurrent range of the
<emphasis box="[1267,1438,1387,1408]" italics="true" pageId="6" pageNumber="7">Calyptocephalella-</emphasis>
and
<taxonomicName authorityName="Thomas" authorityYear="1894" box="[415,523,1415,1436]" class="Mammalia" family="Microbiotheriidae" genus="Dromiciops" kingdom="Animalia" order="Microbiotheria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis box="[415,523,1415,1436]" italics="true" pageId="6" pageNumber="7">Dromiciops</emphasis>
</taxonomicName>
-inhabited
<taxonomicName box="[627,733,1414,1435]" class="Magnoliopsida" family="Nothofagaceae" genus="Nothofagus" kingdom="Plantae" order="Fagales" pageId="6" pageNumber="7" phylum="Tracheophyta" rank="genus">
<emphasis box="[627,733,1414,1435]" italics="true" pageId="6" pageNumber="7">Nothofagus</emphasis>
</taxonomicName>
forests of South America.
</paragraph>
<paragraph blockId="6.[415,1480,1197,1596]" pageId="6" pageNumber="7">
The fossil finds of a frog and marsupial from Seymour Island, and their fossil and Recent distributions, represent outstanding examples of the role of global climate change on shifting biogeographic ranges. Despite global cooling and the disappearance of the habitats of these groups over large areas from Antarctica to Patagonia, they maintained their relictual occurrence in the
<taxonomicName box="[835,941,1520,1541]" class="Magnoliopsida" family="Nothofagaceae" genus="Nothofagus" kingdom="Plantae" order="Fagales" pageId="6" pageNumber="7" phylum="Tracheophyta" rank="genus">
<emphasis box="[835,941,1520,1541]" italics="true" pageId="6" pageNumber="7">Nothofagus</emphasis>
</taxonomicName>
forests of the central Chilean Andes. Thus, the Valdivian
<taxonomicName box="[415,521,1547,1568]" class="Magnoliopsida" family="Nothofagaceae" genus="Nothofagus" kingdom="Plantae" order="Fagales" pageId="6" pageNumber="7" phylum="Tracheophyta" rank="genus">
<emphasis box="[415,521,1547,1568]" italics="true" pageId="6" pageNumber="7">Nothofagus</emphasis>
</taxonomicName>
forest is a unique environment offering habitats not only for Eocene Antarctic refugees but also provides a modern analogue of the Antarctic climate just prior to the glaciation of the southern continent.
</paragraph>
</subSubSection>
</treatment>
</document>