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<mods:title id="DD41C2951B05236ABD1B60F969D45D78">Systematic revision of Sarcosuchus hartti (Crocodyliformes) from the Recôncavo Basin (Early Cretaceous) of Bahia, north-eastern Brazil</mods:title>
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<mods:namePart id="448DA5264BEB14EF8AFC911A1F724D2C">Azevedo, Sérgio A. K.</mods:namePart>
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<mods:namePart id="B0DD09FC2D2D3556A79C58B2F253DF05">Riff, Douglas</mods:namePart>
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<mods:namePart id="93F6AEAE90F2E76D2F23FABC8FC6D4DC">Kellner, Alexander W. A.</mods:namePart>
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<subSubSection id="C352FB8AFFF9FFF9EEAAF904447503EE" box="[951,1316,1761,1784]" pageId="18" pageNumber="570" type="nomenclature">
<paragraph id="8BF7A801FFF9FFF9EEAAF904447503EE" blockId="18.[951,1316,1761,1784]" box="[951,1316,1761,1784]" pageId="18" pageNumber="570">
<heading id="D0BF1F6DFFF9FFF9EEAAF904447503EE" box="[951,1316,1761,1784]" centered="true" fontSize="9" level="2" pageId="18" pageNumber="570" reason="2">
<taxonomicName id="4C48D382FFF9FFF9EEAAF904447503EE" authority="PALAEOECOLOGY" authorityName="PALAEOECOLOGY" box="[951,1316,1761,1784]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="18" pageNumber="570" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFF9FFF9EEAAF904450E03EE" box="[951,1119,1761,1784]" italics="true" pageId="18" pageNumber="570">SARCOSUCHUS</emphasis>
PALAEOECOLOGY
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C352FB8AFFF9FFF8EE27F8EC443E07B0" lastPageId="19" lastPageNumber="571" pageId="18" pageNumber="570" type="reference_group">
<paragraph id="8BF7A801FFF9FFF8EE27F8EC402603AF" blockId="18.[826,1442,1801,1885]" lastBlockId="19.[144,761,197,1905]" lastPageId="19" lastPageNumber="571" pageId="18" pageNumber="570">
<taxonomicName id="4C48D382FFF9FFF9EE27F8EC45120209" box="[826,1091,1801,1823]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" kingdom="Animalia" order="Crocodylia" pageId="18" pageNumber="570" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFF9FFF9EE27F8EC428B0208" box="[826,986,1801,1822]" italics="true" pageId="18" pageNumber="570">Sarcosuchus</emphasis>
species
</taxonomicName>
are gigantic semi-aquatic crocodyliforms that inhabited fluvial environments during the Early Cretaceous of what is today known as South America and Africa (
<bibRefCitation id="EFD9D5F0FFF8FFF8ECD6FF2043A005CC" author="Buffetaut E &amp; Taquet P" box="[459,753,197,219]" pageId="19" pageNumber="571" pagination="203 - 208" refId="ref20787" refString="Buffetaut E, Taquet P. 1977. The giant crocodilian Sarcosuchus in the Early Cretaceous of Brazil and Niger. Palaentology 20: 203 - 208." type="journal article" year="1977">Buffetaut &amp; Taquet, 1977</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFF8FFF8ED8DFF01403205EF" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" box="[144,355,228,250]" pageId="19" pageNumber="571" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFF8FFF8EDFBFF00404F05EF" box="[230,286,228,250]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
, 2001
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFF8FFF8EC6DFF0140A105EF" author="Dridi J" box="[368,496,228,250]" pageId="19" pageNumber="571" pagination="268 - 280" refId="ref20959" refString="Dridi J. 2018. New fossils of the giant pholidosaurid genus Sarcosuchus from the Early Cretaceous of Tunisia. Journal of African Earth Sciences 147: 268 - 280." type="journal article" year="2018">Dridi, 2018</bibRefCitation>
). The two
<taxonomicName id="4C48D382FFF8FFF8EF74FF0141B1040E" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFF8FFF8EF74FF0143A905EF" box="[617,760,228,249]" italics="true" pageId="19" pageNumber="571">Sarcosuchus</emphasis>
species
</taxonomicName>
share the same general rostral and mandibular morphology. They have long and wide snouts that are dorsoventrally compressed and, at least in adult specimens, also show well-marked lateral expansions of the anterior ends. The dentition is heterodont on both the upper and lower jaws, with anterior caniniform teeth and small, rounded and robust posterior teeth. A general overbite occlusion pattern is inferred for both species, with interlocking teeth from the premaxilla to at least the level of the seventh alveoli. An interocclusal pattern on the posterior region is suggested by the presence of tooth-marks on the space between the alveoli.
<bibRefCitation id="EFD9D5F0FFF8FFF8EC99FD964338079F" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" box="[388,617,626,649]" pageId="19" pageNumber="571" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFF8FFF8ECC0FD9643490791" box="[477,536,626,648]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
(2001)
</bibRefCitation>
pointed out that despite some morphological adaptations that are traditionally related to primary ichthyophagy feeding in extant
<taxonomicName id="4C48D382FFF8FFF8EDE1FD2B402607F2" box="[252,375,718,740]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="order">Crocodylia</taxonomicName>
,
<taxonomicName id="4C48D382FFF8FFF8EC9CFD2B43D407F2" box="[385,645,718,740]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="imperator">
<emphasis id="B93C7413FFF8FFF8EC9CFD2B43D407F2" box="[385,645,718,740]" italics="true" pageId="19" pageNumber="571">Sarcosuchus imperator</emphasis>
</taxonomicName>
may have had a more generalized diet, including large terrestrial prey, such as dinosaurs. A similar predatory behaviour is observed in
<taxonomicName id="4C48D382FFF8FFF8EC5CFCCF43A20656" authority="Laurenti, 1768" authorityName="Laurenti" authorityYear="1768" box="[321,755,810,832]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="niloticus">
<emphasis id="B93C7413FFF8FFF8EC5CFCCF43640629" box="[321,565,810,831]" italics="true" pageId="19" pageNumber="571">Crocodylus niloticus</emphasis>
Laurenti, 1768
</taxonomicName>
, which feeds on large mammals. Despite the several morphological similarities shared with
<taxonomicName id="4C48D382FFF8FFF8EF79FC8D4189068D" baseAuthorityName="Campos &amp; Kellner" baseAuthorityYear="1991" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="hartti">
<emphasis id="B93C7413FFF8FFF8EF79FC8D4189068D" italics="true" pageId="19" pageNumber="571">Sarcosuchus hartti</emphasis>
</taxonomicName>
, this feeding behaviour was never directly proposed for the Brazilian species. However, some complementary comments are necessary to refine the
<bibRefCitation id="EFD9D5F0FFF8FFF8ED8DFC07402006EE" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" box="[144,369,994,1016]" pageId="19" pageNumber="571" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFF8FFF8EDFAFC06407006E1" box="[231,289,994,1016]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
(2001)
</bibRefCitation>
proposition and construct a more robust palaeoecological hypothesis for the behaviour of
<taxonomicName id="4C48D382FFF8FFF8EDB6FBC540C70120" box="[171,406,1056,1078]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B93C7413FFF8FFF8EDB6FBC540680123" box="[171,313,1056,1077]" italics="true" pageId="19" pageNumber="571">Sarcosuchus</emphasis>
species.
</taxonomicName>
<taxonomicName id="4C48D382FFF8FFF8EC81FBC543D30123" authorityName="Laurenti" authorityYear="1768" box="[412,642,1056,1077]" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="niloticus">
<emphasis id="B93C7413FFF8FFF8EC81FBC543D30123" box="[412,642,1056,1077]" italics="true" pageId="19" pageNumber="571">Crocodylus niloticus</emphasis>
</taxonomicName>
is a largesized extant
<taxonomicName id="4C48D382FFF8FFF8EC37FBDB40F90142" box="[298,424,1086,1108]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="order">Crocodylia</taxonomicName>
that in adult life ambushes big mammals, dragging them underwater to drown and then tears them apart by a death roll movement (
<bibRefCitation id="EFD9D5F0FFF8FFF8ED85FB7F40DD01A6" author="Pooley AC &amp; Gans C" box="[152,396,1178,1200]" pageId="19" pageNumber="571" pagination="114 - 124" refId="ref22320" refString="Pooley AC, Gans C. 1976. The Nile crocodile. Scientific American 234: 114 - 124." type="journal article" year="1976">Pooley &amp; Gans, 1976</bibRefCitation>
).
<bibRefCitation id="EFD9D5F0FFF8FFF8ECBEFB7F43D601A6" author="Blanco RE &amp; Jones WW &amp; Villamal J" box="[419,647,1178,1200]" pageId="19" pageNumber="571" pagination="514 - 524" refId="ref20223" refString="Blanco RE, Jones WW, Villamal J. 2015. The ' death roll' of giant fossil crocodyliforms (Crocodylomorpha: Neosuchia): allometric and skull strength analysis. Historical Biology 27: 514 - 524." type="journal article" year="2015">
Blanco
<emphasis id="B93C7413FFF8FFF8ECE6FB7E436701B9" box="[507,566,1178,1200]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
(2015)
</bibRefCitation>
analysed the allometry and skull strength of several extant
<taxonomicName id="4C48D382FFF8FFF8ED8DFB3D404201F8" box="[144,275,1240,1262]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="order">Crocodylia</taxonomicName>
and other fossil species. Their results suggest that
<taxonomicName id="4C48D382FFF8FFF8EC36FB13436A001A" box="[299,571,1270,1292]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="imperator">
<emphasis id="B93C7413FFF8FFF8EC36FB13436A001A" box="[299,571,1270,1292]" italics="true" pageId="19" pageNumber="571">Sarcosuchus imperator</emphasis>
</taxonomicName>
was not able to perform the death roll movement, contra to what was proposed by
<bibRefCitation id="EFD9D5F0FFF8FFF8EC45FAD1436D005C" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" box="[344,572,1332,1354]" pageId="19" pageNumber="571" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFF8FFF8ECACFAD140BA005F" box="[433,491,1332,1353]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
(2001)
</bibRefCitation>
. So, the feeding strategy of preying on large-sized dinosaurs, larger than
<taxonomicName id="4C48D382FFF8FFF8EDD3FA9440310090" box="[206,352,1393,1414]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFF8FFF8EDD3FA9440310090" box="[206,352,1393,1414]" italics="true" pageId="19" pageNumber="571">Sarcosuchus</emphasis>
</taxonomicName>
itself, is unlikely if the death roll movement is a requirement for that strategy (
<bibRefCitation id="EFD9D5F0FFF8FFF8EFB4FA75404800D2" author="Blanco RE &amp; Jones WW &amp; Villamal J" pageId="19" pageNumber="571" pagination="514 - 524" refId="ref20223" refString="Blanco RE, Jones WW, Villamal J. 2015. The ' death roll' of giant fossil crocodyliforms (Crocodylomorpha: Neosuchia): allometric and skull strength analysis. Historical Biology 27: 514 - 524." type="journal article" year="2015">
Blanco
<emphasis id="B93C7413FFF8FFF8ED8DFA4A419C00D5" box="[144,205,1454,1476]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
, 2015
</bibRefCitation>
; and references therein). Nevertheless, smaller dinosaurs would still be a potential prey for
<taxonomicName id="4C48D382FFF8FFF8ED8DFA0940D50314" box="[144,388,1516,1538]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFF8FFF8ED8DFA0940720317" box="[144,291,1516,1537]" italics="true" pageId="19" pageNumber="571">Sarcosuchus</emphasis>
species.
</taxonomicName>
The feeding behaviour of these extinct animals is probably more like that observed in
<taxonomicName id="4C48D382FFF8FFF8EDADF9CF430B0329" authority="(Muller, 1838)" baseAuthorityName="Muller" baseAuthorityYear="1838" box="[176,602,1577,1599]" class="Reptilia" family="Crocodylidae" genus="Tomistoma" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="schlegelii">
<emphasis id="B93C7413FFF8FFF8EDADF9CF40FB0328" box="[176,426,1577,1599]" italics="true" pageId="19" pageNumber="571">Tomistoma schlegelii</emphasis>
(Müller, 1838)
</taxonomicName>
or
<taxonomicName id="4C48D382FFF8FFF8EF99F9CF408A034B" authority="(Cuvier, 1825)" baseAuthorityName="Cuvier" baseAuthorityYear="1825" class="Reptilia" family="Crocodylidae" genus="Mecistops" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="cataphractus">
<emphasis id="B93C7413FFF8FFF8EF99F9CF407B034B" italics="true" pageId="19" pageNumber="571">Mecistops cataphractus</emphasis>
(Cuvier, 1825)
</taxonomicName>
, which prey on animals smaller than themselves, swallowing them completely without the need of applying a death roll movement (
<bibRefCitation id="EFD9D5F0FFF8FFF8ED85F941403903AF" author="Blanco RE &amp; Jones WW &amp; Villamal J" box="[152,360,1699,1722]" pageId="19" pageNumber="571" pagination="514 - 524" refId="ref20223" refString="Blanco RE, Jones WW, Villamal J. 2015. The ' death roll' of giant fossil crocodyliforms (Crocodylomorpha: Neosuchia): allometric and skull strength analysis. Historical Biology 27: 514 - 524." type="journal article" year="2015">
Blanco
<emphasis id="B93C7413FFF8FFF8EDF1F941407503AF" box="[236,292,1700,1721]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
, 2015
</bibRefCitation>
).
</paragraph>
<paragraph id="8BF7A801FFF8FFF8EDB5F927443E07B0" blockId="19.[144,761,197,1905]" lastBlockId="19.[808,1425,197,955]" pageId="19" pageNumber="571">
There are some features in both
<taxonomicName id="4C48D382FFF8FFF8EF0EF92743A603CE" box="[531,759,1730,1752]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFF8FFF8EF0EF92743F003C1" box="[531,673,1730,1751]" italics="true" pageId="19" pageNumber="571">Sarcosuchus</emphasis>
species
</taxonomicName>
that could support other interesting behaviours. The heterodont dentition suggests a facultative durophagy, as observed among large-sized Alligatoroidea, such as
<taxonomicName id="4C48D382FFF8FFF8EDAFF8D843C70245" authority="(Daudin, 1802)" baseAuthorityName="Daudin" baseAuthorityYear="1802" box="[178,662,1853,1875]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="mississippiensis">
<emphasis id="B93C7413FFF8FFF8EDAFF8D8408E0244" box="[178,479,1853,1874]" italics="true" pageId="19" pageNumber="571">Alligator mississippiensis</emphasis>
(Daudin, 1802)
</taxonomicName>
and the extinct
<taxonomicName id="4C48D382FFF8FFF8EDF6F8BE423805CD" authority="(Colbert &amp; Bird, 1954)" baseAuthorityName="Colbert &amp; Bird" baseAuthorityYear="1954" box="[235,873,197,1905]" class="Reptilia" family="Crocodylidae" genus="Deinosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="species" species="riograndensis">
<emphasis id="B93C7413FFF8FFF8EDF6F8BE437D0266" box="[235,556,1883,1904]" italics="true" pageId="19" pageNumber="571">Deinosuchus riograndensis</emphasis>
(Colbert &amp; Bird, 1954)
</taxonomicName>
(
<bibRefCitation id="EFD9D5F0FFF8FFF8EE66FF20455D05CC" author="Pooley AC" box="[891,1036,197,219]" pageId="19" pageNumber="571" pagination="76 - 91" refId="ref22288" refString="Pooley AC. 1989. Food and feeding habits. In: Ross CA, ed. Crocodile and alligators. New York: Facts on File, 76 - 91." type="book chapter" year="1989">Pooley, 1989</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFF8FFF8E906FF2045BC05CC" author="Schwimmer DR" box="[1051,1261,197,219]" pageId="19" pageNumber="571" refId="ref22621" refString="Schwimmer DR. 2002. King of the crocodylians: the paleobiology of Deinosuchus. Rio de Janeiro: Bloomington: Indiana University Press, 220." type="book" year="2002">Schwimmer, 2002</bibRefCitation>
). In this way,
<taxonomicName id="4C48D382FFF8FFF8EE35FF06429005EE" box="[808,961,227,248]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFF8FFF8EE35FF06429005EE" box="[808,961,227,248]" italics="true" pageId="19" pageNumber="571">Sarcosuchus</emphasis>
</taxonomicName>
could be able to prey on turtles and crush large bones of carcasses. Another interesting feature shared by both species of
<taxonomicName id="4C48D382FFF8FFF8E9A9FEC444170420" box="[1204,1350,289,310]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFF8FFF8E9A9FEC444170420" box="[1204,1350,289,310]" italics="true" pageId="19" pageNumber="571">Sarcosuchus</emphasis>
</taxonomicName>
is the ontogenetic modifications observed on the lateral projections of both the dentary and the premaxilla, which was illustrated by
<bibRefCitation id="EFD9D5F0FFF8FFF8E94FFE9844D80485" author="Buffetaut E &amp; Taquet P" box="[1106,1417,381,403]" pageId="19" pageNumber="571" pagination="203 - 208" refId="ref20787" refString="Buffetaut E, Taquet P. 1977. The giant crocodilian Sarcosuchus in the Early Cretaceous of Brazil and Niger. Palaentology 20: 203 - 208." type="journal article" year="1977">Buffetaut &amp; Taquet (1977)</bibRefCitation>
, and probably imply in a differential dentition pattern observed among juveniles and adults specimens (e.g.
<bibRefCitation id="EFD9D5F0FFF8FFF8EE35FE3C455A04F8" author="Erickson GM &amp; Kristopher Lappin A &amp; Vliet KA" box="[808,1035,473,495]" pageId="19" pageNumber="571" pagination="317 - 327" refId="ref20990" refString="Erickson GM, Kristopher Lappin A, Vliet KA. 2003. The ontogeny of bite-force performance in American alligator (Alligator mississippiensis). Journal of Zoology 260: 317 - 327." type="journal article" year="2003">
Erickson
<emphasis id="B93C7413FFF8FFF8EE8FFE3C429804F8" box="[914,969,473,494]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
, 2003
</bibRefCitation>
). The ontogenetic changes suggest the presence of niche partitioning, a well-known phenomenon observed in extant
<taxonomicName id="4C48D382FFF8FFF8E9DEFDF34418073A" box="[1219,1353,534,556]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="order">Crocodylia</taxonomicName>
, with juveniles preying on insects, crustaceans, mollusks and small fish, while adults prey on large terrestrial and aquatic animals, such as mammals, turtles and big fish, but also carcasses (e.g.
<bibRefCitation id="EFD9D5F0FFF8FFF8E98DFD74443107B1" author="Blanco RE &amp; Jones WW &amp; Villamal J" box="[1168,1376,657,679]" pageId="19" pageNumber="571" pagination="514 - 524" refId="ref20223" refString="Blanco RE, Jones WW, Villamal J. 2015. The ' death roll' of giant fossil crocodyliforms (Crocodylomorpha: Neosuchia): allometric and skull strength analysis. Historical Biology 27: 514 - 524." type="journal article" year="2015">
Blanco
<emphasis id="B93C7413FFF8FFF8E9F8FD74444C07B0" box="[1253,1309,657,678]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
, 2015
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C352FB8AFFF8FFFCEE5DFD4A43F50334" lastPageId="23" lastPageNumber="575" pageId="19" pageNumber="571" type="description">
<paragraph id="8BF7A801FFF8FFF8EE5DFD4A42A806AD" blockId="19.[808,1425,197,955]" pageId="19" pageNumber="571">
The development of different kinds of long snouts in the evolutionary history of Crocodyliformes and other vertebrates remains an undergoing field of research to understand the real natural pressures that positively select for those modifications (see:
<bibRefCitation id="EFD9D5F0FFF8FFF8E9CDFCCF42300648" author="Walmsley CW &amp; Smits PD &amp; Quayle MR &amp; McCurry MR &amp; Richards HS &amp; Oldfield CC &amp; Wroe S &amp; Clausen PD &amp; McHenry CR" pageId="19" pageNumber="571" pagination="53873" refId="ref23024" refString="Walmsley CW, Smits PD, Quayle MR, McCurry MR, Richards HS, Oldfield CC, Wroe S, Clausen PD, McHenry CR. 2013. Why the long face? The mechanics of mandibular symphysis proportions in crocodiles. PLoS ONE 8: e 53873." type="journal article" year="2013">
Walmsley
<emphasis id="B93C7413FFF8FFF8E850FCCE44D80629" box="[1357,1417,810,832]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
, 2013
</bibRefCitation>
). Also, the adult modifications on the rostrum of
<taxonomicName id="4C48D382FFF8FFF8EE5BFC82428B066A" box="[838,986,871,892]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFF8FFF8EE5BFC82428B066A" box="[838,986,871,892]" italics="true" pageId="19" pageNumber="571">Sarcosuchus</emphasis>
</taxonomicName>
are a feature in need of explanation and that requires more specific morphometrics and strength analyses.
</paragraph>
<paragraph id="8BF7A801FFF8FFF8EEDEFC1245A40119" blockId="19.[963,1269,1015,1040]" box="[963,1269,1015,1040]" pageId="19" pageNumber="571">PHYLOGENETIC AFFINITIES</paragraph>
<paragraph id="8BF7A801FFF8FFF8EE35FBC545A50222" blockId="19.[808,1425,1056,1906]" pageId="19" pageNumber="571">
The phylogenetic relationships of
<taxonomicName id="4C48D382FFF8FFF8E9F6FBC5443E0120" authorityName="BENTON &amp; CLARK" authorityYear="1988" box="[1259,1391,1056,1078]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neosuchia" pageId="19" pageNumber="571" phylum="Chordata" rank="order">Neosuchia</taxonomicName>
is one of the most important issues regarding the evolution of crocodylomorphs, yet much of the effort in understanding the morphological variation within the major clades is still in progress (
<bibRefCitation id="EFD9D5F0FFF8FFF8E9C6FB7F44D801A6" author="Pol D &amp; Turner AH &amp; Norell MA" box="[1243,1417,1178,1200]" pageId="19" pageNumber="571" pagination="1 - 103" refId="ref22241" refString="Pol D, Turner AH, Norell MA. 2009. Morphology of the Late Cretaceous crocodylomorph Shamosuchus djadochtaensis and a discussion of neosuchian phylogeny as related to the origin of Eusuchia. Bulletin of the American Museum of Natural History 324: 1 - 103." type="journal article" year="2009">
Pol
<emphasis id="B93C7413FFF8FFF8E815FB7E441201B9" box="[1288,1347,1178,1200]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
, 2009
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFF8FFF8EE35FB5F42E801D9" author="Turner AH &amp; Pritchard AC" box="[808,953,1209,1231]" pageId="19" pageNumber="571" pagination="759" refId="ref22912" refString="Turner AH, Pritchard AC. 2015. The monophyly of Susisuchidae (Crocodyliformes) and its phylogenetic placement in Neosuchia. PeerJ 3: e 759." type="journal article" year="2015">Turner, 2015</bibRefCitation>
). One classic example is the longirostrine problem, i.e. the close affinities of Thalattosuchia with other long-snouted crocodylomorphs, such as the dyrosaurids and pholidosaurids, within the clade of
<taxonomicName id="4C48D382FFF8FFF8EE35FAD145C6005C" authority="(Clark, 1994)" baseAuthorityName="Clark" baseAuthorityYear="1994" box="[808,1175,1332,1354]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Mesoeucrocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="order">Mesoeucrocodylia (Clark, 1994)</taxonomicName>
. The derived position of the group is far from being considered a consensus in the literature and several phylogenetic analyses place them either as basal mesoeucrocodylians or outside Crocodyliformes (e.g.
<bibRefCitation id="EFD9D5F0FFF8FFF8E96CFA4A446E00D2" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" box="[1137,1343,1454,1476]" pageId="19" pageNumber="571" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFF8FFF8E9DBFA4A45AD00D5" box="[1222,1276,1454,1476]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
, 2001
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFF8FFF8E854FA4A42A200F4" author="Young MT &amp; Andrade MB" pageId="19" pageNumber="571" pagination="551 - 585" refId="ref23234" refString="Young MT, Andrade MB. 2009. What is Geosaurus? Redescription of Geosaurus giganteus (Thalattosuchia: Metriorhynchidae) from the Upper Jurassic of Bayern, Germany. Zoological Journal of the Linnean Society 157: 551 - 585." type="journal article" year="2009">Young &amp; Andrade, 2009</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFF8FFF8E91FFA28444F00F4" author="Parrilla-Bel J &amp; Young MT &amp; Moreno-Azanza M &amp; Canudo JI" box="[1026,1310,1485,1507]" pageId="19" pageNumber="571" pagination="54275" refId="ref22143" refString="Parrilla-Bel J, Young MT, Moreno-Azanza M, Canudo JI. 2013. The first metriorhynchid crocodylomorph from the Middle Jurassic of Spain, with implications for evolution of the subclade Rhacheosaurini. PLoS ONE 8: e 54275." type="journal article" year="2013">
Parrilla-Bel
<emphasis id="B93C7413FFF8FFF8E984FA2B458400F4" box="[1177,1237,1485,1507]" italics="true" pageId="19" pageNumber="571">et al.</emphasis>
, 2013
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFF8FFF8E831FA28420E0314" author="Wilberg EW" pageId="19" pageNumber="571" pagination="902846" refId="ref23074" refString="Wilberg EW. 2015. A new metriorhynchoid (Crocodylomorpha, Thalattosuchia) from the Middle Jurassic of Oregon and the evolutionary timing of marine adaptations in thalattosuchian crocodylomorphs. Journal of Vertebrate Paleontology 35: e 902846." type="journal article" year="2015">Wilberg, 2015</bibRefCitation>
). The neosuchian hypothesis for the positioning of thalattosuchians seems to be largely an effect of taxon and character sampling, which also influences the interpretation of character evolution in other species (
<bibRefCitation id="EFD9D5F0FFF8FFF8EE94F9834576036A" author="Wilberg EW" box="[905,1063,1638,1660]" pageId="19" pageNumber="571" pagination="902846" refId="ref23074" refString="Wilberg EW. 2015. A new metriorhynchoid (Crocodylomorpha, Thalattosuchia) from the Middle Jurassic of Oregon and the evolutionary timing of marine adaptations in thalattosuchian crocodylomorphs. Journal of Vertebrate Paleontology 35: e 902846." type="journal article" year="2015">Wilberg, 2015</bibRefCitation>
). For this reason, the alternate non-neosuchian hypothesis is used in the present study with the a priori exclusion of Thalattosuchia from the phylogenetic analysis. This scenario provides new insights in the evolution and biogeography of pholidosaurids and closely related taxa like the dyrosaurids and goniopholidids (
<figureCitation id="1373B484FFF8FFF8E9B8F8FB45B40225" box="[1189,1253,1822,1844]" captionStart="Figure 9" captionStartId="12.[165,245,1501,1523]" captionTargetBox="[326,1277,198,1457]" captionTargetId="figure-180@12.[325,1285,196,1462]" captionTargetPageId="12" captionText="Figure 9. Consensus cladogram resulted from 12 minimum-length trees with 1287 steps (CI: 0.327 and RI: 0.610). The numbered nodes are the following clades: 1, Neosuchia; 2, Susisuchidae; 3, Eusuchia; 4, Allodaposuchidae; 5, Crocodylia; 6, Coelognathosuchia; 7, Goniopholididae; 8, Tethysuchia; 9, Pholidosauridae; 10, Tethysuchoidea; 11, Dyrosauridae." figureDoi="http://doi.org/10.5281/zenodo.5719182" httpUri="https://zenodo.org/record/5719182/files/figure.png" pageId="19" pageNumber="571">Fig. 9</figureCitation>
).
</paragraph>
<paragraph id="8BF7A801FFF8FFFFEE5DF8D843F007F2" blockId="19.[808,1425,1056,1906]" lastBlockId="20.[164,781,197,1905]" lastPageId="20" lastPageNumber="572" pageId="19" pageNumber="571">
There are three main competing hypotheses for the higher relationships of
<taxonomicName id="4C48D382FFF8FFF8E97EF8B944700264" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[1123,1313,1884,1906]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="571" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
. The first one supports a closer affinity with the
<taxonomicName id="4C48D382FFFFFFFFEF70FF20425D05CD" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[621,780,197,219]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
(e.g.
<bibRefCitation id="EFD9D5F0FFFFFFFFEDFFFF01408105EF" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" box="[226,464,228,250]" pageId="20" pageNumber="572" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFFFFFFFEC5FFF0040D205EF" box="[322,387,228,250]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2001
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFFFFFFECFEFF0143F205EF" author="Pol D &amp; Turner AH &amp; Norell MA" box="[483,675,228,250]" pageId="20" pageNumber="572" pagination="1 - 103" refId="ref22241" refString="Pol D, Turner AH, Norell MA. 2009. Morphology of the Late Cretaceous crocodylomorph Shamosuchus djadochtaensis and a discussion of neosuchian phylogeny as related to the origin of Eusuchia. Bulletin of the American Museum of Natural History 324: 1 - 103." type="journal article" year="2009">
Pol
<emphasis id="B93C7413FFFFFFFFEF09FF00430405EF" box="[532,597,228,250]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2009
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFFFFFFEFA8FF014077040E" author="Fortier D &amp; Perea D &amp; Schultz C" pageId="20" pageNumber="572" pagination="272" refId="ref21024" refString="Fortier D, Perea D, Schultz C. 2011. Redescription and phylogenetic relationships of Meridiosaurus valliparadisi, a pholidosaurid from the Late Jurassic of Uruguay. Zoological Journal of the Linnean Society 163: S 257 - S 272." type="journal article" year="2011">
Fortier
<emphasis id="B93C7413FFFFFFFFEDB9FEE6418E0401" box="[164,223,258,280]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2011
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;
<bibRefCitation id="EFD9D5F0FFFFFFFFEC2BFEE7431D040E" author="Montefeltro FC &amp; Larsson HCE &amp; Franca MAG &amp; Langer MC" box="[310,588,258,280]" pageId="20" pageNumber="572" pagination="835 - 841" refId="ref22057" refString="Montefeltro FC, Larsson HCE, Franca MAG, Langer MC. 2013. A new neosuchian with Asian affinities from the Jurassic of northeastern Brazil. Naturwissenschaften 100: 835 - 841." type="journal article" year="2013">
Montefeltro
<emphasis id="B93C7413FFFFFFFFECD4FEE643540401" box="[457,517,258,280]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2013
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFFFFFFEF46FEE7418D0421" author="Halliday TJD &amp; Andrade MB &amp; Benton MJ &amp; Efimov MB" pageId="20" pageNumber="572" pagination="291 - 312" refId="ref21127" refString="Halliday TJD, Andrade MB, Benton MJ, Efimov MB. 2015. A re-evaluation of goniopholidid crocodylomorph material from Central Asia: biogeographic and phylogenetic implications. Acta Palaeontologica Polonica 60: 291 - 312." type="journal article" year="2015">
Halliday
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, 2015
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;
<bibRefCitation id="EFD9D5F0FFFFFFFFEDF7FEC740D30421" author="Turner AH &amp; Pritchard AC" box="[234,386,289,311]" pageId="20" pageNumber="572" pagination="759" refId="ref22912" refString="Turner AH, Pritchard AC. 2015. The monophyly of Susisuchidae (Crocodyliformes) and its phylogenetic placement in Neosuchia. PeerJ 3: e 759." type="journal article" year="2015">Turner, 2015</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFFFFFFEC8DFEC743320420" author="Young MT &amp; Hastings AK &amp; Allan R &amp; Smith TJ" box="[400,611,289,311]" pageId="20" pageNumber="572" pagination="377 - 403" refId="ref23276" refString="Young MT, Hastings AK, Allan R, Smith TJ. 2017. Revision of the enigmatic crocodyliform Elosuchus felixi de Lapparent de Broin, 2002 from the Lower-Upper Cretaceous boundary of Niger: potential evidence for an early origin of the clade Dyrosauridae. Zoological Journal of the Linnean Society 179: 377 - 403." type="journal article" year="2017">
Young
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, 2017
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;
<bibRefCitation id="EFD9D5F0FFFFFFFFEF6CFEC4418F0443" author="Adams TL &amp; Noto CR &amp; Drumheller SK" pageId="20" pageNumber="572" pagination="1349776" refId="ref19990" refString="Adams TL, Noto CR, Drumheller SK. 2017. A Large Neosuchian Crocodyliform From The Upper Cretaceous (Cenomanian) Woodbine Formation Of North Texas. Journal of Vertebrate Paleontology e 1349776." type="book chapter" year="2017">
Adams
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;
<bibRefCitation id="EFD9D5F0FFFFFFFFEDF2FEA540BB0443" author="Schwarz D &amp; Raddatz M &amp; Wings O" box="[239,490,320,342]" pageId="20" pageNumber="572" pagination="0160617" refId="ref22568" refString="Schwarz D, Raddatz M, Wings O. 2017. Knoetschkesuchus langenbergensis gen. nov. sp. nov., a new atoposaurid crocodyliform from the Upper Jurassic Langenberg Quarry (Lower Saxony, northwestern Germany), and its relationships to Theriosuchus. PLoS ONE 12: e 0160617." type="journal article" year="2017">
Schwarz
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).
<bibRefCitation id="EFD9D5F0FFFFFFFFEF19FEA5425A0440" author="Andrade MB &amp; Edmonds R &amp; Benton MJ &amp; Schouten R" box="[516,779,320,342]" pageId="20" pageNumber="572" pagination="108" refId="ref20104" refString="Andrade MB, Edmonds R, Benton MJ, Schouten R. 2011. A new Berriasian species of Goniopholis (Mesoeucrocodylia, Neosuchia) from England, and a review of the genus. Zoological Journal of the Linnean Society 163: S 66 - S 108." type="journal article" year="2011">
Andrade
<emphasis id="B93C7413FFFFFFFFEF6BFEA543E40443" box="[630,693,320,341]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2011)
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proposed a redefinition of the name
<taxonomicName id="4C48D382FFFFFFFFEF4EFEBB43B20462" authorityName="BUFFETAUT" authorityYear="1982" box="[595,739,350,372]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
for this clade, which was composed of
<taxonomicName id="4C48D382FFFFFFFFEF47FE98406904A7" authorityName="Mansel-Pleydell" authorityYear="1888" class="Reptilia" family="Pholidosauridae" genus="Pholidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="purbeckensis">
<emphasis id="B93C7413FFFFFFFFEF47FE98406904A7" italics="true" pageId="20" pageNumber="572">Pholidosaurus purbeckensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C48D382FFFFFFFFEC6EFE7941B404C7" authority="(Thomas, 1839)" baseAuthorityName="Thomas" baseAuthorityYear="1839" class="Reptilia" family="Dyrosauridae" genus="Dyrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="phosphaticus">
<emphasis id="B93C7413FFFFFFFFEC6EFE7943CB04A7" box="[371,666,412,433]" italics="true" pageId="20" pageNumber="572">Dyrosaurus phosphaticus</emphasis>
(Thomas, 1839)
</taxonomicName>
, their common ancestor and all its descendants.
<taxonomicName id="4C48D382FFFFFFFFEDB9FE3C406D04F9" authorityName="BUFFETAUT" authorityYear="1982" box="[164,316,473,495]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
was originally created by Buffetaut (1982) as an infraorder comprising a single family, the
<taxonomicName id="4C48D382FFFFFFFFEDCAFDF340D4073A" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[215,389,534,556]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
. This arrangement recognized the anatomical distinctiveness of dyrosaurids in comparison to other longirostrine taxa, especially teleosaurids and pholidosaurids (Buffetaut, 1982). Goniopholidids are usually recovered as more closely related to Eusuchia in this pholidosaurid-dyrosaurid hypothesis (e.g. Jouve, 2009;
<bibRefCitation id="EFD9D5F0FFFFFFFFECF6FD2B43C007F2" author="Pol D &amp; Turner AH &amp; Norell MA" box="[491,657,718,740]" pageId="20" pageNumber="572" pagination="1 - 103" refId="ref22241" refString="Pol D, Turner AH, Norell MA. 2009. Morphology of the Late Cretaceous crocodylomorph Shamosuchus djadochtaensis and a discussion of neosuchian phylogeny as related to the origin of Eusuchia. Bulletin of the American Museum of Natural History 324: 1 - 103." type="journal article" year="2009">
Pol
<emphasis id="B93C7413FFFFFFFFEF0BFD2A431F07F5" box="[534,590,718,740]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2009
</bibRefCitation>
).
</paragraph>
<paragraph id="8BF7A801FFFFFFFFEDA1FD08405F003C" blockId="20.[164,781,197,1905]" pageId="20" pageNumber="572">
The second hypothesis favours a relationship between
<taxonomicName id="4C48D382FFFFFFFFEC02FCE940A80637" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[287,505,780,802]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
and a paraphyletic arrangement of goniopholidids (Martin &amp; Buffetaut, 2012). This clade was named Coelognathosuchia by Martin
<emphasis id="B93C7413FFFFFFFFEDE2FC8D406B066B" box="[255,314,872,893]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2014), but as observed by Young
<emphasis id="B93C7413FFFFFFFFEFCCFC8D425D066B" box="[721,780,872,893]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2014) the absence of dyrosaurid taxa in the analyses of Martin &amp; Buffetaut (2012) and Martin
<emphasis id="B93C7413FFFFFFFFEF99FC4343ED06AC" box="[644,700,933,955]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2014) means that the monophyly of
<taxonomicName id="4C48D382FFFFFFFFEF0CFC21438406CC" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[529,725,964,986]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
was not properly assessed in the light of all evidence. More than that, the exclusion of dyrosaurids shows that there is no support for a Coelognathosuchia clade as originally proposed by Martin
<emphasis id="B93C7413FFFFFFFFEF05FBDA43050145" box="[536,596,1086,1108]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2014). Rather the phylogenetic analyses performed by
<bibRefCitation id="EFD9D5F0FFFFFFFFEF64FBB841A00184" author="Martin JE &amp; Raslan-Loubatie J &amp; Mazin J-M" pageId="20" pageNumber="572" pagination="43 - 59" refId="ref21983" refString="Martin JE, Raslan-Loubatie J, Mazin J-M. 2016. Cranial anatomy of Pholidosaurus purbeckensis from the Lower Cretaceous of France and its bering on pholidosaurid affinities. Cretaceous Research 66: 43 - 59." type="journal article" year="2016">
Martin
<emphasis id="B93C7413FFFFFFFFEFCEFBBB425D0164" box="[723,780,1117,1139]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2016)
</bibRefCitation>
also supports the
<taxonomicName id="4C48D382FFFFFFFFECFFFB99432B0184" authorityName="BUFFETAUT" authorityYear="1982" box="[482,634,1148,1170]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
hypothesis when dyrosaurids are included in the analysis (
<bibRefCitation id="EFD9D5F0FFFFFFFFEDB1FB5C40D401D9" author="Martin JE &amp; Raslan-Loubatie J &amp; Mazin J-M" box="[172,389,1209,1231]" pageId="20" pageNumber="572" pagination="43 - 59" refId="ref21983" refString="Martin JE, Raslan-Loubatie J, Mazin J-M. 2016. Cranial anatomy of Pholidosaurus purbeckensis from the Lower Cretaceous of France and its bering on pholidosaurid affinities. Cretaceous Research 66: 43 - 59." type="journal article" year="2016">
Martin
<emphasis id="B93C7413FFFFFFFFEC18FB5F406C01D8" box="[261,317,1209,1231]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2016
</bibRefCitation>
: fig. 10A). The Coelognathosuchia hypothesis is only recovered with the exclusion of
<taxonomicName id="4C48D382FFFFFFFFEDB9FB134017001A" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[164,326,1270,1292]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
from the dataset (
<bibRefCitation id="EFD9D5F0FFFFFFFFEF3BFB134256001A" author="Martin JE &amp; Raslan-Loubatie J &amp; Mazin J-M" box="[550,775,1270,1292]" pageId="20" pageNumber="572" pagination="43 - 59" refId="ref21983" refString="Martin JE, Raslan-Loubatie J, Mazin J-M. 2016. Cranial anatomy of Pholidosaurus purbeckensis from the Lower Cretaceous of France and its bering on pholidosaurid affinities. Cretaceous Research 66: 43 - 59." type="journal article" year="2016">
Martin
<emphasis id="B93C7413FFFFFFFFEF9FFB1243EC001D" box="[642,701,1270,1292]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2016
</bibRefCitation>
: fig. 10D).
</paragraph>
<paragraph id="8BF7A801FFFFFFFFEDA1FAD140FF036A" blockId="20.[164,781,197,1905]" pageId="20" pageNumber="572">
A third hypothesis shows a sister-taxon position between
<taxonomicName id="4C48D382FFFFFFFFEC03FAB640A7007E" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[286,502,1362,1384]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
and Thalattosuchia (
<bibRefCitation id="EFD9D5F0FFFFFFFFEDB1FA9440E50090" author="Lauprasert K &amp; Cuny G &amp; Thirakhupt K &amp; Suteethorn V" box="[172,436,1393,1415]" pageId="20" pageNumber="572" pagination="175 - 187" refId="ref21724" refString="Lauprasert K, Cuny G, Thirakhupt K, Suteethorn V. 2009. Khoratosuchus jintasakuli gen. et sp. nov., an advanced neosuchian crocodyliform from the Early Cretaceous (Aptian- Albian) of NE Thailand. Geological Society, London, Special Publications 315: 175 - 187." type="journal article" year="2009">
Lauprasert
<emphasis id="B93C7413FFFFFFFFEC2BFA97403E0090" box="[310,367,1393,1415]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2009
</bibRefCitation>
), but it is important to notice that
<taxonomicName id="4C48D382FFFFFFFFEDC2FA7540DA00B3" authorityName="Meyer" authorityYear="1841" box="[223,395,1424,1445]" class="Reptilia" family="Pholidosauridae" genus="Pholidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFEDC2FA7540DA00B3" box="[223,395,1424,1445]" italics="true" pageId="20" pageNumber="572">Pholidosaurus</emphasis>
</taxonomicName>
is the only pholidosaurid taxon included in this analysis. The
<taxonomicName id="4C48D382FFFFFFFFEF3EFA4B43BD00D2" authorityName="Cope" authorityYear="1875" box="[547,748,1454,1476]" class="Reptilia" family="Goniopholididae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Goniopholididae</taxonomicName>
is monophyletic in this scenario, but the inclusion of a few derived taxa shows that the higher affinities inside
<taxonomicName id="4C48D382FFFFFFFFEDF0F9EF40320336" authorityName="BENTON &amp; CLARK" authorityYear="1988" box="[237,355,1546,1568]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neosuchia" pageId="20" pageNumber="572" phylum="Chordata" rank="order">Neosuchia</taxonomicName>
are poorly resolved (
<bibRefCitation id="EFD9D5F0FFFFFFFFEF56F9EE418C0328" author="Lauprasert K &amp; Cuny G &amp; Thirakhupt K &amp; Suteethorn V" pageId="20" pageNumber="572" pagination="175 - 187" refId="ref21724" refString="Lauprasert K, Cuny G, Thirakhupt K, Suteethorn V. 2009. Khoratosuchus jintasakuli gen. et sp. nov., an advanced neosuchian crocodyliform from the Early Cretaceous (Aptian- Albian) of NE Thailand. Geological Society, London, Special Publications 315: 175 - 187." type="journal article" year="2009">
Lauprasert
<emphasis id="B93C7413FFFFFFFFEFCDF9EE42570309" box="[720,774,1546,1568]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2009
</bibRefCitation>
: fig. 3a, b). In any case, despite this arrangement was new at that time, the authors did not extensively discuss this hypothesis.
</paragraph>
<paragraph id="8BF7A801FFFFFFFFEDA1F96042BB0656" blockId="20.[164,781,197,1905]" lastBlockId="20.[828,1445,197,1905]" pageId="20" pageNumber="572">
The results of the current analysis recovered the
<taxonomicName id="4C48D382FFFFFFFFEDC1F941402E03AF" authorityName="BUFFETAUT" authorityYear="1982" box="[220,383,1700,1722]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
hypothesis (
<figureCitation id="1373B484FFFFFFFFEF25F94143D603AF" box="[568,647,1700,1722]" captionStart="Figure 9" captionStartId="12.[165,245,1501,1523]" captionTargetBox="[326,1277,198,1457]" captionTargetId="figure-180@12.[325,1285,196,1462]" captionTargetPageId="12" captionText="Figure 9. Consensus cladogram resulted from 12 minimum-length trees with 1287 steps (CI: 0.327 and RI: 0.610). The numbered nodes are the following clades: 1, Neosuchia; 2, Susisuchidae; 3, Eusuchia; 4, Allodaposuchidae; 5, Crocodylia; 6, Coelognathosuchia; 7, Goniopholididae; 8, Tethysuchia; 9, Pholidosauridae; 10, Tethysuchoidea; 11, Dyrosauridae." figureDoi="http://doi.org/10.5281/zenodo.5719182" httpUri="https://zenodo.org/record/5719182/files/figure.png" pageId="20" pageNumber="572">Fig. 9</figureCitation>
; node
<figureCitation id="1373B484FFFFFFFFEFF6F94143A803AF" box="[747,761,1700,1721]" captionStart="Figure 8" captionStartId="11.[146,224,490,512]" captionTargetBox="[373,1204,199,449]" captionTargetId="figure-746@11.[368,1204,197,451]" captionTargetPageId="11" captionText="Figure 8. Sarcosuchus sp., osteoderm (BMNH R3224).A, dorsal view. B, ventral view.Abbreviations: art, articular facet; ap, anterior projection; op, ornamentation pit; vcb, ventral compact bone tissue; meb, medial spongiest bone tissue; dcb, dorsal compact bone tissue. Scale bar: 3 cm." figureDoi="http://doi.org/10.5281/zenodo.5719178" httpUri="https://zenodo.org/record/5719178/files/figure.png" pageId="20" pageNumber="572">8</figureCitation>
).
<taxonomicName id="4C48D382FFFFFFFFEDB9F927401F03CE" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[164,334,1730,1752]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
is a less inclusive clade nested in several pholidosaurid lineages, being sister to the
<taxonomicName id="4C48D382FFFFFFFFEDB9F91A40660203" box="[164,311,1791,1813]" class="Reptilia" family="Elosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Elosuchidae</taxonomicName>
.
<taxonomicName id="4C48D382FFFFFFFFEC59F91A43560203" authorityName="Cope" authorityYear="1875" box="[324,519,1791,1813]" class="Reptilia" family="Goniopholididae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Goniopholididae</taxonomicName>
is paraphyletic in its traditional sense. There are several smaller groups that are usually recognized as goniopholidids, and all of them are more closely related to
<taxonomicName id="4C48D382FFFFFFFFEF37F8BE43E40267" authorityName="BUFFETAUT" authorityYear="1982" box="[554,693,1883,1905]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
than to Eusuchia. This large clade, comprising tethysuchians and goniopholidids, was also recovered by
<bibRefCitation id="EFD9D5F0FFFFFFFFE84CFF01429A040E" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" pageId="20" pageNumber="572" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFFFFFFFEE21FEE642290401" box="[828,888,258,280]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2001)
</bibRefCitation>
and
<bibRefCitation id="EFD9D5F0FFFFFFFFE914FEE745A7040E" author="Martin JE &amp; Raslan-Loubatie J &amp; Mazin J-M" box="[1033,1270,258,280]" pageId="20" pageNumber="572" pagination="43 - 59" refId="ref21983" refString="Martin JE, Raslan-Loubatie J, Mazin J-M. 2016. Cranial anatomy of Pholidosaurus purbeckensis from the Lower Cretaceous of France and its bering on pholidosaurid affinities. Cretaceous Research 66: 43 - 59." type="journal article" year="2016">
Martin
<emphasis id="B93C7413FFFFFFFFE97BFEE645F30401" box="[1126,1186,258,280]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2016)
</bibRefCitation>
. These groups of animals have been classified together in the past, but in a gradist scheme. Nopcsa (1928) grouped the subfamilies
<taxonomicName id="4C48D382FFFFFFFFE909FEBB45860462" baseAuthorityName="Dollo" baseAuthorityYear="1914" box="[1044,1239,350,372]" class="Reptilia" family="Goniopholididae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="subFamily" subFamily="Congosaurinae">Congosaurinae</taxonomicName>
,
<taxonomicName id="4C48D382FFFFFFFFE9F4FEBB44CF0462" authorityName="Nopcsa" authorityYear="1928" box="[1257,1438,350,372]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="subFamily" subFamily="Hyposaurinae">Hyposaurinae</taxonomicName>
,
<taxonomicName id="4C48D382FFFFFFFFEE21FE9842B80485" authorityName="Nopcsa" authorityYear="1928" box="[828,1001,381,403]" class="Reptilia" family="Goniopholididae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="subFamily" subFamily="Goniopholinae">Goniopholinae</taxonomicName>
, Pholidosaurinae and Bernissartinae in the family
<taxonomicName id="4C48D382FFFFFFFFEEC8FE7E452A04A7" authorityName="Cope" authorityYear="1875" box="[981,1147,411,433]" class="Reptilia" family="Goniopholidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Goniopholidae</taxonomicName>
. While most analyses find
<taxonomicName id="4C48D382FFFFFFFFEE21FE5E429C04C6" box="[828,973,443,464]" class="Reptilia" family="Bernissartiidae" genus="Bernissartia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFEE21FE5E429C04C6" box="[828,973,443,464]" italics="true" pageId="20" pageNumber="572">Bernissartia</emphasis>
</taxonomicName>
as more closely related to the Eusuchia (e.g.
<bibRefCitation id="EFD9D5F0FFFFFFFFEE6FFE3C451804F8" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" box="[882,1097,473,494]" pageId="20" pageNumber="572" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFFFFFFFEED4FE3C455204F8" box="[969,1027,473,494]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2001
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFFFFFFE94BFE3C441004F9" author="Halliday TJD &amp; Andrade MB &amp; Benton MJ &amp; Efimov MB" box="[1110,1345,473,495]" pageId="20" pageNumber="572" pagination="291 - 312" refId="ref21127" refString="Halliday TJD, Andrade MB, Benton MJ, Efimov MB. 2015. A re-evaluation of goniopholidid crocodylomorph material from Central Asia: biogeographic and phylogenetic implications. Acta Palaeontologica Polonica 60: 291 - 312." type="journal article" year="2015">
Halliday
<emphasis id="B93C7413FFFFFFFFE9DFFE3C45AD04F8" box="[1218,1276,473,494]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2015
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFFFFFFE853FE3C4227071B" author="Turner AH &amp; Pritchard AC" pageId="20" pageNumber="572" pagination="759" refId="ref22912" refString="Turner AH, Pritchard AC. 2015. The monophyly of Susisuchidae (Crocodyliformes) and its phylogenetic placement in Neosuchia. PeerJ 3: e 759." type="journal article" year="2015">Turner, 2015</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFFFFFFEE9BFE12453D071B" author="Adams TL &amp; Noto CR &amp; Drumheller SK" box="[902,1132,503,525]" pageId="20" pageNumber="572" pagination="1349776" refId="ref19990" refString="Adams TL, Noto CR, Drumheller SK. 2017. A Large Neosuchian Crocodyliform From The Upper Cretaceous (Cenomanian) Woodbine Formation Of North Texas. Journal of Vertebrate Paleontology e 1349776." type="book chapter" year="2017">
Adams
<emphasis id="B93C7413FFFFFFFFEEFEFE1D4570071A" box="[995,1057,503,525]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2017
</bibRefCitation>
), that is not the case for the present hypothesis.
<taxonomicName id="4C48D382FFFFFFFFE97DFDF245A4073A" box="[1120,1269,535,556]" class="Reptilia" family="Bernissartiidae" genus="Bernissartia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFE97DFDF245A4073A" box="[1120,1269,535,556]" italics="true" pageId="20" pageNumber="572">Bernissartia</emphasis>
</taxonomicName>
together with
<taxonomicName id="4C48D382FFFFFFFFEE21FDD042B4075C" box="[828,997,565,586]" class="Reptilia" family="Atoposauridae" genus="Theriosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFEE21FDD042B4075C" box="[828,997,565,586]" italics="true" pageId="20" pageNumber="572">Theriosuchus</emphasis>
</taxonomicName>
(i.e.
<taxonomicName id="4C48D382FFFFFFFFE930FDD045B6075D" box="[1069,1255,565,587]" class="Reptilia" family="Atoposauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Atoposauridae</taxonomicName>
) is the sistertaxon of this large unnamed clade of goniopholidids and tethysuchians. Hay (1930) recognizes the Goniopholidiformes as the group including both the
<taxonomicName id="4C48D382FFFFFFFFEE21FD4A42A807D3" authorityName="Cope" authorityYear="1875" box="[828,1017,687,709]" class="Reptilia" family="Goniopholididae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Goniopholididae</taxonomicName>
and the
<taxonomicName id="4C48D382FFFFFFFFE97DFD4A444C07D3" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[1120,1309,687,709]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
, which also encompassed the dyrosaurids. This name would be suitable in case the clade proves to be stable with time, especially after the inclusion of more dyrosaurid taxa in the analysis.
</paragraph>
<paragraph id="8BF7A801FFFFFFFFEE49FCAC4298038C" blockId="20.[828,1445,197,1905]" pageId="20" pageNumber="572">
The phylogenetic hypothesis presented here shows important results regarding the relationships among pholidosaurids, because it includes the higher number of such taxa (nine species), together with that of
<bibRefCitation id="EFD9D5F0FFFFFFFFEE21FC21457106CF" author="Young MT &amp; Hastings AK &amp; Allan R &amp; Smith TJ" box="[828,1056,963,985]" pageId="20" pageNumber="572" pagination="377 - 403" refId="ref23276" refString="Young MT, Hastings AK, Allan R, Smith TJ. 2017. Revision of the enigmatic crocodyliform Elosuchus felixi de Lapparent de Broin, 2002 from the Lower-Upper Cretaceous boundary of Niger: potential evidence for an early origin of the clade Dyrosauridae. Zoological Journal of the Linnean Society 179: 377 - 403." type="journal article" year="2017">
Young
<emphasis id="B93C7413FFFFFFFFEE92FC21429D06CE" box="[911,972,963,985]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2017)
</bibRefCitation>
. Other relevant hypotheses are those of
<bibRefCitation id="EFD9D5F0FFFFFFFFEE82FC0745D206EE" author="Fortier D &amp; Perea D &amp; Schultz C" box="[927,1155,994,1016]" pageId="20" pageNumber="572" pagination="272" refId="ref21024" refString="Fortier D, Perea D, Schultz C. 2011. Redescription and phylogenetic relationships of Meridiosaurus valliparadisi, a pholidosaurid from the Late Jurassic of Uruguay. Zoological Journal of the Linnean Society 163: S 257 - S 272." type="journal article" year="2011">
Fortier
<emphasis id="B93C7413FFFFFFFFEEE5FC06456306E1" box="[1016,1074,994,1016]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2011)
</bibRefCitation>
and
<bibRefCitation id="EFD9D5F0FFFFFFFFE9A0FC07443006EE" author="Turner AH &amp; Pritchard AC" box="[1213,1377,994,1016]" pageId="20" pageNumber="572" pagination="759" refId="ref22912" refString="Turner AH, Pritchard AC. 2015. The monophyly of Susisuchidae (Crocodyliformes) and its phylogenetic placement in Neosuchia. PeerJ 3: e 759." type="journal article" year="2015">Turner (2015)</bibRefCitation>
, each with six species. All those more complete analyses recovered a paraphyletic Pholidosauridae, except for
<bibRefCitation id="EFD9D5F0FFFFFFFFEE78FBDB45010142" author="Fortier D &amp; Perea D &amp; Schultz C" box="[869,1104,1086,1108]" pageId="20" pageNumber="572" pagination="272" refId="ref21024" refString="Fortier D, Perea D, Schultz C. 2011. Redescription and phylogenetic relationships of Meridiosaurus valliparadisi, a pholidosaurid from the Late Jurassic of Uruguay. Zoological Journal of the Linnean Society 163: S 257 - S 272." type="journal article" year="2011">
Fortier
<emphasis id="B93C7413FFFFFFFFEEDCFBDA42AC0145" box="[961,1021,1086,1108]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2011)
</bibRefCitation>
. There are two sister-clades of pholidosaurids in this latter hypothesis: the first unites
<taxonomicName id="4C48D382FFFFFFFFEE95FB9E457C0186" authorityName="Meyer" authorityYear="1841" box="[904,1069,1147,1168]" class="Reptilia" family="Pholidosauridae" genus="Pholidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFEE95FB9E457C0186" box="[904,1069,1147,1168]" italics="true" pageId="20" pageNumber="572">Pholidosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C48D382FFFFFFFFE97FFB9E45A10186" box="[1122,1264,1147,1168]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFE97FFB9E45A10186" box="[1122,1264,1147,1168]" italics="true" pageId="20" pageNumber="572">Sarcosuchus</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFFFFFFE817FB9944F50187" baseAuthorityName="Buffetaut &amp; Wellnhofer" baseAuthorityYear="1980" box="[1290,1444,1148,1169]" class="Reptilia" family="Teleosauridae" genus="Terminonaris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFE817FB9944F50187" box="[1290,1444,1148,1169]" italics="true" pageId="20" pageNumber="572">Terminonaris</emphasis>
</taxonomicName>
and the second
<taxonomicName id="4C48D382FFFFFFFFE917FB7F459301B9" authorityName="Mones" authorityYear="1980" box="[1034,1218,1178,1199]" class="Reptilia" family="Pholidosauridae" genus="Meridiosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFE917FB7F459301B9" box="[1034,1218,1178,1199]" italics="true" pageId="20" pageNumber="572">Meridiosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C48D382FFFFFFFFE815FB7F44D901B9" authorityName="de Lapparent de Broin" authorityYear="2002" box="[1288,1416,1178,1199]" class="Reptilia" family="Pholidosauridae" genus="Elosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFE815FB7F44D901B9" box="[1288,1416,1178,1199]" italics="true" pageId="20" pageNumber="572">Elosuchus</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFFFFFFEE21FB5C42B501D8" box="[828,996,1209,1230]" class="Reptilia" family="Pholidosauridae" genus="Oceanosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFEE21FB5C42B501D8" box="[828,996,1209,1230]" italics="true" pageId="20" pageNumber="572">Oceanosuchus</emphasis>
</taxonomicName>
. Most phylogenetic hypotheses show
<taxonomicName id="4C48D382FFFFFFFFEE21FB32452A01FA" authorityName="Mansel-Pleydell" authorityYear="1888" box="[828,1147,1239,1260]" class="Reptilia" family="Pholidosauridae" genus="Pholidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="purbeckensis">
<emphasis id="B93C7413FFFFFFFFEE21FB32452A01FA" box="[828,1147,1239,1260]" italics="true" pageId="20" pageNumber="572">Pholidosaurus purbeckensis</emphasis>
</taxonomicName>
as more distantly related to the other pholidosaurids, which usually are the sister-taxon of the dyrosaurids (e.g.
<bibRefCitation id="EFD9D5F0FFFFFFFFE81CFAF04222005E" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" pageId="20" pageNumber="572" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFFFFFFFE843FAF044CC003C" box="[1374,1437,1301,1322]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2001
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFFFFFFEE9CFAD64538005F" author="Halliday TJD &amp; Andrade MB &amp; Benton MJ &amp; Efimov MB" box="[897,1129,1331,1353]" pageId="20" pageNumber="572" pagination="291 - 312" refId="ref21127" refString="Halliday TJD, Andrade MB, Benton MJ, Efimov MB. 2015. A re-evaluation of goniopholidid crocodylomorph material from Central Asia: biogeographic and phylogenetic implications. Acta Palaeontologica Polonica 60: 291 - 312." type="journal article" year="2015">
Halliday
<emphasis id="B93C7413FFFFFFFFEEF6FAD14575005E" box="[1003,1060,1331,1353]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2015
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFFFFFFE96BFAD14458005F" author="Turner AH &amp; Pritchard AC" box="[1142,1289,1331,1353]" pageId="20" pageNumber="572" pagination="759" refId="ref22912" refString="Turner AH, Pritchard AC. 2015. The monophyly of Susisuchidae (Crocodyliformes) and its phylogenetic placement in Neosuchia. PeerJ 3: e 759." type="journal article" year="2015">Turner, 2015</bibRefCitation>
; Young
<emphasis id="B93C7413FFFFFFFFE878FAD144CF005E" box="[1381,1438,1331,1353]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2016;
<bibRefCitation id="EFD9D5F0FFFFFFFFEE9CFAB7450A0071" author="Adams TL &amp; Noto CR &amp; Drumheller SK" box="[897,1115,1362,1384]" pageId="20" pageNumber="572" pagination="1349776" refId="ref19990" refString="Adams TL, Noto CR, Drumheller SK. 2017. A Large Neosuchian Crocodyliform From The Upper Cretaceous (Cenomanian) Woodbine Formation Of North Texas. Journal of Vertebrate Paleontology e 1349776." type="book chapter" year="2017">
Adams
<emphasis id="B93C7413FFFFFFFFEEC7FAB645450071" box="[986,1044,1362,1384]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2017
</bibRefCitation>
). In the current hypothesis,
<taxonomicName id="4C48D382FFFFFFFFEE21FA9442BB0090" authorityName="Mansel-Pleydell" authorityYear="1888" box="[828,1002,1393,1414]" class="Reptilia" family="Pholidosauridae" genus="Pholidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="purbeckensis">
<emphasis id="B93C7413FFFFFFFFEE21FA94421D0090" box="[828,844,1393,1414]" italics="true" pageId="20" pageNumber="572">P</emphasis>
.
<emphasis id="B93C7413FFFFFFFFEE44FA9442BB0090" box="[857,1002,1393,1414]" italics="true" pageId="20" pageNumber="572">purbeckensis</emphasis>
</taxonomicName>
comprises a clade with
<taxonomicName id="4C48D382FFFFFFFFE9E7FA94445600B2" authority="Lauprasert et al., 2009" authorityName="Lauprasert" authorityYear="2009" class="Reptilia" genus="Khoratosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Neosuchia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="jintasakuli">
<emphasis id="B93C7413FFFFFFFFE9E7FA94429800B2" italics="true" pageId="20" pageNumber="572">Khoratosuchus jintasakuli</emphasis>
<bibRefCitation id="EFD9D5F0FFFFFFFFEECBFA75445600B2" author="Lauprasert K &amp; Cuny G &amp; Thirakhupt K &amp; Suteethorn V" box="[982,1287,1423,1445]" pageId="20" pageNumber="572" pagination="175 - 187" refId="ref21724" refString="Lauprasert K, Cuny G, Thirakhupt K, Suteethorn V. 2009. Khoratosuchus jintasakuli gen. et sp. nov., an advanced neosuchian crocodyliform from the Early Cretaceous (Aptian- Albian) of NE Thailand. Geological Society, London, Special Publications 315: 175 - 187." type="journal article" year="2009">
Lauprasert
<emphasis id="B93C7413FFFFFFFFE96EFA7545E700B2" box="[1139,1206,1423,1445]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
, 2009
</bibRefCitation>
</taxonomicName>
(a putative advanced neosuchian),
<taxonomicName id="4C48D382FFFFFFFFE969FA4B44F200D5" box="[1140,1443,1454,1475]" class="Reptilia" family="Pholidosauridae" genus="Oceanosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="boecensis">
<emphasis id="B93C7413FFFFFFFFE969FA4B44F200D5" box="[1140,1443,1454,1475]" italics="true" pageId="20" pageNumber="572">Oceanosuchus boecensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C48D382FFFFFFFFEE6DFA28444A00F4" authority="Efimov, 1975" authorityName="Efimov" authorityYear="1975" box="[880,1307,1484,1507]" class="Reptilia" family="Goniopholididae" genus="Kansajsuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="extensus">
<emphasis id="B93C7413FFFFFFFFEE6DFA28452D00F4" box="[880,1148,1485,1506]" italics="true" pageId="20" pageNumber="572">Kansajsuchus extensus</emphasis>
Efimov, 1975
</taxonomicName>
(a putative goniopholidid). This latter clade, which is sister to the other tethysuchians (i.e. the other pholidosaurids and
<taxonomicName id="4C48D382FFFFFFFFEE21F9CC42890329" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[828,984,1577,1599]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
) is considered here to be
<taxonomicName id="4C48D382FFFFFFFFE9F6F9CC44F50329" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[1259,1444,1577,1599]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
<emphasis id="B93C7413FFFFFFFFEE21F9AD429C034B" box="[828,973,1608,1629]" italics="true" pageId="20" pageNumber="572">sensu stricto</emphasis>
(
<figureCitation id="1373B484FFFFFFFFEEC0F9A2454F034A" box="[989,1054,1607,1629]" captionStart="Figure 9" captionStartId="12.[165,245,1501,1523]" captionTargetBox="[326,1277,198,1457]" captionTargetId="figure-180@12.[325,1285,196,1462]" captionTargetPageId="12" captionText="Figure 9. Consensus cladogram resulted from 12 minimum-length trees with 1287 steps (CI: 0.327 and RI: 0.610). The numbered nodes are the following clades: 1, Neosuchia; 2, Susisuchidae; 3, Eusuchia; 4, Allodaposuchidae; 5, Crocodylia; 6, Coelognathosuchia; 7, Goniopholididae; 8, Tethysuchia; 9, Pholidosauridae; 10, Tethysuchoidea; 11, Dyrosauridae." figureDoi="http://doi.org/10.5281/zenodo.5719182" httpUri="https://zenodo.org/record/5719182/files/figure.png" pageId="20" pageNumber="572">Fig. 9</figureCitation>
; node 9). Interestingly, the genus
<taxonomicName id="4C48D382FFFFFFFFEE21F98342B0036D" authorityName="Meyer" authorityYear="1841" box="[828,993,1638,1659]" class="Reptilia" family="Pholidosauridae" genus="Pholidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFEE21F98342B0036D" box="[828,993,1638,1659]" italics="true" pageId="20" pageNumber="572">Pholidosaurus</emphasis>
</taxonomicName>
is paraphyletic in the analysis of
<bibRefCitation id="EFD9D5F0FFFFFFFFE840F9834290038D" author="Young MT &amp; Hastings AK &amp; Allan R &amp; Smith TJ" pageId="20" pageNumber="572" pagination="377 - 403" refId="ref23276" refString="Young MT, Hastings AK, Allan R, Smith TJ. 2017. Revision of the enigmatic crocodyliform Elosuchus felixi de Lapparent de Broin, 2002 from the Lower-Upper Cretaceous boundary of Niger: potential evidence for an early origin of the clade Dyrosauridae. Zoological Journal of the Linnean Society 179: 377 - 403." type="journal article" year="2017">
Young
<emphasis id="B93C7413FFFFFFFFEE21F9604225038C" box="[828,884,1669,1690]" italics="true" pageId="20" pageNumber="572">et al.</emphasis>
(2017)
</bibRefCitation>
.
</paragraph>
<paragraph id="8BF7A801FFFFFFFEEE49F94643AA07D0" blockId="20.[828,1445,197,1905]" lastBlockId="21.[146,763,197,1047]" lastPageId="21" lastPageNumber="573" pageId="20" pageNumber="572">
Our novel analysis shows a second large clade inside
<taxonomicName id="4C48D382FFFFFFFFEE21F927429B03CE" authorityName="BUFFETAUT" authorityYear="1982" box="[828,970,1730,1752]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
with
<taxonomicName id="4C48D382FFFFFFFFE910F927440703C1" box="[1037,1366,1730,1751]" class="Reptilia" family="Pholidosauridae" genus="Meridiosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="valliparadisi">
<emphasis id="B93C7413FFFFFFFFE910F927440703C1" box="[1037,1366,1730,1751]" italics="true" pageId="20" pageNumber="572">Meridiosaurus valliparadisi</emphasis>
</taxonomicName>
as the sister-species of the
<taxonomicName id="4C48D382FFFFFFFFE93AF90445BE03E0" box="[1063,1263,1761,1782]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B93C7413FFFFFFFFE93AF90445E903E0" box="[1063,1208,1761,1782]" italics="true" pageId="20" pageNumber="572">Sarcosuchus</emphasis>
spp.
</taxonomicName>
, plus the group of
<taxonomicName id="4C48D382FFFFFFFFEE4BF8E5451A0202" baseAuthorityName="Mook" baseAuthorityYear="1934" box="[854,1099,1791,1813]" class="Reptilia" family="Teleosauridae" genus="Terminonaris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="robusta">
<emphasis id="B93C7413FFFFFFFFEE4BF8E5451A0202" box="[854,1099,1791,1813]" italics="true" pageId="20" pageNumber="572">Terminonaris robusta</emphasis>
</taxonomicName>
, which is sister to
<taxonomicName id="4C48D382FFFFFFFFE804F91A44F20203" box="[1305,1443,1791,1813]" class="Reptilia" family="Elosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Elosuchidae</taxonomicName>
plus
<taxonomicName id="4C48D382FFFFFFFFEE67F8FB45720222" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[890,1059,1822,1844]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
. This new clade is here named Tethysuchoidea (
<figureCitation id="1373B484FFFFFFFFEEE2F8D9456E0244" box="[1023,1087,1852,1874]" captionStart="Figure 9" captionStartId="12.[165,245,1501,1523]" captionTargetBox="[326,1277,198,1457]" captionTargetId="figure-180@12.[325,1285,196,1462]" captionTargetPageId="12" captionText="Figure 9. Consensus cladogram resulted from 12 minimum-length trees with 1287 steps (CI: 0.327 and RI: 0.610). The numbered nodes are the following clades: 1, Neosuchia; 2, Susisuchidae; 3, Eusuchia; 4, Allodaposuchidae; 5, Crocodylia; 6, Coelognathosuchia; 7, Goniopholididae; 8, Tethysuchia; 9, Pholidosauridae; 10, Tethysuchoidea; 11, Dyrosauridae." figureDoi="http://doi.org/10.5281/zenodo.5719182" httpUri="https://zenodo.org/record/5719182/files/figure.png" pageId="20" pageNumber="572">Fig. 9</figureCitation>
; node 10). In some hypotheses,
<taxonomicName id="4C48D382FFFFFFFFEE21F8B942A50266" box="[828,1012,1883,1905]" class="Reptilia" family="Pholidosauridae" genus="Meridiosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="species" species="valliparadisi">
<emphasis id="B93C7413FFFFFFFFEE21F8B942020267" box="[828,851,1884,1905]" italics="true" pageId="20" pageNumber="572">M</emphasis>
.
<emphasis id="B93C7413FFFFFFFFEE7DF8BE42A50266" box="[864,1012,1883,1904]" italics="true" pageId="20" pageNumber="572">valliparadisi</emphasis>
</taxonomicName>
shows close affinities with
<taxonomicName id="4C48D382FFFFFFFFE832F8BE44F20266" authorityName="de Lapparent de Broin" authorityYear="2002" box="[1327,1443,1883,1904]" class="Reptilia" family="Pholidosauridae" genus="Elosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="20" pageNumber="572" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFFFFFFE832F8BE44F20266" box="[1327,1443,1883,1904]" italics="true" pageId="20" pageNumber="572">Elosuchus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFD9D5F0FFFEFFFEED87FF20403C05CC" author="Fortier D &amp; Perea D &amp; Schultz C" box="[154,365,197,219]" pageId="21" pageNumber="573" pagination="272" refId="ref21024" refString="Fortier D, Perea D, Schultz C. 2011. Redescription and phylogenetic relationships of Meridiosaurus valliparadisi, a pholidosaurid from the Late Jurassic of Uruguay. Zoological Journal of the Linnean Society 163: S 257 - S 272." type="journal article" year="2011">
Fortier
<emphasis id="B93C7413FFFEFFFEEDEDFF23407805CC" box="[240,297,197,219]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2011
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFEFFFEEC64FF23435D05CD" author="Turner AH &amp; Pritchard AC" box="[377,524,197,219]" pageId="21" pageNumber="573" pagination="759" refId="ref22912" refString="Turner AH, Pritchard AC. 2015. The monophyly of Susisuchidae (Crocodyliformes) and its phylogenetic placement in Neosuchia. PeerJ 3: e 759." type="journal article" year="2015">Turner, 2015</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFEFFFEEF05FF2043BA05CC" author="Adams TL &amp; Noto CR &amp; Drumheller SK" box="[536,747,197,219]" pageId="21" pageNumber="573" pagination="1349776" refId="ref19990" refString="Adams TL, Noto CR, Drumheller SK. 2017. A Large Neosuchian Crocodyliform From The Upper Cretaceous (Cenomanian) Woodbine Formation Of North Texas. Journal of Vertebrate Paleontology e 1349776." type="book chapter" year="2017">
Adams
<emphasis id="B93C7413FFFEFFFEEF72FF2343F605CC" box="[623,679,197,219]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2017
</bibRefCitation>
). However, as observed on the current analysis and others (e.g.
<bibRefCitation id="EFD9D5F0FFFEFFFEEC01FEE7435A040E" author="Halliday TJD &amp; Andrade MB &amp; Benton MJ &amp; Efimov MB" box="[284,523,258,280]" pageId="21" pageNumber="573" pagination="291 - 312" refId="ref21127" refString="Halliday TJD, Andrade MB, Benton MJ, Efimov MB. 2015. A re-evaluation of goniopholidid crocodylomorph material from Central Asia: biogeographic and phylogenetic implications. Acta Palaeontologica Polonica 60: 291 - 312." type="journal article" year="2015">
Halliday
<emphasis id="B93C7413FFFEFFFEEC94FEE640950401" box="[393,452,258,280]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2015
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFEFFFEEF04FEE643BA040E" author="Young MT &amp; Hastings AK &amp; Allan R &amp; Smith TJ" box="[537,747,258,280]" pageId="21" pageNumber="573" pagination="377 - 403" refId="ref23276" refString="Young MT, Hastings AK, Allan R, Smith TJ. 2017. Revision of the enigmatic crocodyliform Elosuchus felixi de Lapparent de Broin, 2002 from the Lower-Upper Cretaceous boundary of Niger: potential evidence for an early origin of the clade Dyrosauridae. Zoological Journal of the Linnean Society 179: 377 - 403." type="journal article" year="2017">
Young
<emphasis id="B93C7413FFFEFFFEEF74FEE643F50401" box="[617,676,258,280]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2017
</bibRefCitation>
), this is only correct when
<taxonomicName id="4C48D382FFFEFFFEECA4FEC4438F0420" authorityName="Buffetaut &amp; Hutt" authorityYear="1980" box="[441,734,289,310]" class="Reptilia" family="Goniopholididae" genus="Vectisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="leptognathus">
<emphasis id="B93C7413FFFEFFFEECA4FEC4438F0420" box="[441,734,289,310]" italics="true" pageId="21" pageNumber="573">Vectisuchus leptognathus</emphasis>
</taxonomicName>
is absent. Also the close affinities between
<taxonomicName id="4C48D382FFFEFFFEEF70FEA543AB0440" box="[621,762,320,342]" class="Reptilia" family="Elosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Elosuchidae</taxonomicName>
and
<taxonomicName id="4C48D382FFFEFFFEEDD6FEBB40240462" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[203,373,350,372]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
is shown here. Regarding the relationship of the
<taxonomicName id="4C48D382FFFEFFFEEC73FE9843090485" box="[366,600,381,403]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B93C7413FFFEFFFEEC73FE9840AF0484" box="[366,510,381,402]" italics="true" pageId="21" pageNumber="573">Sarcosuchus</emphasis>
species
</taxonomicName>
, there is little doubt that both species are sister-taxa, even though the inclusion of
<taxonomicName id="4C48D382FFFEFFFEEC4FFE5E40E404D9" authorityName="Marsh" authorityYear="1869" box="[338,437,442,464]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="hartii">
<emphasis id="B93C7413FFFEFFFEEC4FFE5E403304C6" box="[338,354,443,464]" italics="true" pageId="21" pageNumber="573">S</emphasis>
.
<emphasis id="B93C7413FFFEFFFEEC6CFE5F40E404D9" box="[369,437,442,463]" italics="true" pageId="21" pageNumber="573">hartii</emphasis>
</taxonomicName>
in phylogenetic analysis is rare (e.g.,
<bibRefCitation id="EFD9D5F0FFFEFFFEEC14FE3C40A604F8" author="Andrade MB &amp; Edmonds R &amp; Benton MJ &amp; Schouten R" box="[265,503,473,495]" pageId="21" pageNumber="573" pagination="108" refId="ref20104" refString="Andrade MB, Edmonds R, Benton MJ, Schouten R. 2011. A new Berriasian species of Goniopholis (Mesoeucrocodylia, Neosuchia) from England, and a review of the genus. Zoological Journal of the Linnean Society 163: S 66 - S 108." type="journal article" year="2011">
Andrade
<emphasis id="B93C7413FFFEFFFEEC6BFE3F40E004F8" box="[374,433,473,495]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2011
</bibRefCitation>
). Most analyses show
<taxonomicName id="4C48D382FFFEFFFEED8FFE1D4056071B" baseAuthorityName="Mook" baseAuthorityYear="1934" box="[146,263,504,525]" class="Reptilia" family="Teleosauridae" genus="Terminonaris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="robusta">
<emphasis id="B93C7413FFFEFFFEED8FFE1D41F2071B" box="[146,163,504,525]" italics="true" pageId="21" pageNumber="573">T</emphasis>
.
<emphasis id="B93C7413FFFEFFFEEDADFE1D4056071B" box="[176,263,504,525]" italics="true" pageId="21" pageNumber="573">robusta</emphasis>
</taxonomicName>
as the sister-taxon of
<taxonomicName id="4C48D382FFFEFFFEEF16FE1D43CA071B" box="[523,667,504,525]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="imperator">
<emphasis id="B93C7413FFFEFFFEEF16FE1D434A071B" box="[523,539,504,525]" italics="true" pageId="21" pageNumber="573">S</emphasis>
.
<emphasis id="B93C7413FFFEFFFEEF34FE1D43CA071B" box="[553,667,504,525]" italics="true" pageId="21" pageNumber="573">imperator</emphasis>
</taxonomicName>
when it is the sole species of the genus in an analysis (
<bibRefCitation id="EFD9D5F0FFFEFFFEEFB6FDF24040075C" author="Sereno PC &amp; Larsson HCE &amp; Sidor CA &amp; Gado B" pageId="21" pageNumber="573" pagination="1516 - 1519" refId="ref22648" refString="Sereno PC, Larsson HCE, Sidor CA, Gado B. 2001. The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294: 1516 - 1519." type="journal article" year="2001">
Sereno
<emphasis id="B93C7413FFFEFFFEED8FFDD3419D075C" box="[146,204,565,587]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2001
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFEFFFEEC02FDD040A6075C" author="Fortier D &amp; Perea D &amp; Schultz C" box="[287,503,565,587]" pageId="21" pageNumber="573" pagination="272" refId="ref21024" refString="Fortier D, Perea D, Schultz C. 2011. Redescription and phylogenetic relationships of Meridiosaurus valliparadisi, a pholidosaurid from the Late Jurassic of Uruguay. Zoological Journal of the Linnean Society 163: S 257 - S 272." type="journal article" year="2011">
Fortier
<emphasis id="B93C7413FFFEFFFEEC65FDD340E3075C" box="[376,434,565,587]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2011
</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFEFFFEEF19FDD343CB075D" author="Turner AH &amp; Pritchard AC" box="[516,666,565,587]" pageId="21" pageNumber="573" pagination="759" refId="ref22912" refString="Turner AH, Pritchard AC. 2015. The monophyly of Susisuchidae (Crocodyliformes) and its phylogenetic placement in Neosuchia. PeerJ 3: e 759." type="journal article" year="2015">Turner, 2015</bibRefCitation>
;
<bibRefCitation id="EFD9D5F0FFFEFFFEEFB5FDD04045077C" author="Martin JE &amp; Raslan-Loubatie J &amp; Mazin J-M" pageId="21" pageNumber="573" pagination="43 - 59" refId="ref21983" refString="Martin JE, Raslan-Loubatie J, Mazin J-M. 2016. Cranial anatomy of Pholidosaurus purbeckensis from the Lower Cretaceous of France and its bering on pholidosaurid affinities. Cretaceous Research 66: 43 - 59." type="journal article" year="2016">
Martin
<emphasis id="B93C7413FFFEFFFEED8FFDB1419C077F" box="[146,205,596,617]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2016
</bibRefCitation>
,
<bibRefCitation id="EFD9D5F0FFFEFFFEEC3EFDB14351077F" author="Adams TL &amp; Noto CR &amp; Drumheller SK" box="[291,512,596,618]" pageId="21" pageNumber="573" pagination="1349776" refId="ref19990" refString="Adams TL, Noto CR, Drumheller SK. 2017. A Large Neosuchian Crocodyliform From The Upper Cretaceous (Cenomanian) Woodbine Formation Of North Texas. Journal of Vertebrate Paleontology e 1349776." type="book chapter" year="2017">
Adams
<emphasis id="B93C7413FFFEFFFEEC60FDB140E8077F" box="[381,441,596,617]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2017
</bibRefCitation>
). Notable exceptions are close relationships with
<taxonomicName id="4C48D382FFFEFFFEECFEFD97402B07B1" authority="(Young et al., 2017)" baseAuthorityName="Young" baseAuthorityYear="2017" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="thailandicus">
<emphasis id="B93C7413FFFEFFFEECFEFD9743A80791" box="[483,761,626,647]" italics="true" pageId="21" pageNumber="573">Chalawan thailandicus</emphasis>
(
<bibRefCitation id="EFD9D5F0FFFEFFFEED86FD77402307B0" author="Young MT &amp; Hastings AK &amp; Allan R &amp; Smith TJ" box="[155,370,657,679]" pageId="21" pageNumber="573" pagination="377 - 403" refId="ref23276" refString="Young MT, Hastings AK, Allan R, Smith TJ. 2017. Revision of the enigmatic crocodyliform Elosuchus felixi de Lapparent de Broin, 2002 from the Lower-Upper Cretaceous boundary of Niger: potential evidence for an early origin of the clade Dyrosauridae. Zoological Journal of the Linnean Society 179: 377 - 403." type="journal article" year="2017">
Young
<emphasis id="B93C7413FFFEFFFEEDF0FD77407807B0" box="[237,297,657,679]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2017
</bibRefCitation>
)
</taxonomicName>
or
<taxonomicName id="4C48D382FFFEFFFEECB8FD74418207D3" authority="(Halliday et al., 2015)" baseAuthorityName="Halliday" baseAuthorityYear="2015" class="Reptilia" family="Elosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">
Elosuchidae (
<bibRefCitation id="EFD9D5F0FFFEFFFEEF5AFD74419807D0" author="Halliday TJD &amp; Andrade MB &amp; Benton MJ &amp; Efimov MB" pageId="21" pageNumber="573" pagination="291 - 312" refId="ref21127" refString="Halliday TJD, Andrade MB, Benton MJ, Efimov MB. 2015. A re-evaluation of goniopholidid crocodylomorph material from Central Asia: biogeographic and phylogenetic implications. Acta Palaeontologica Polonica 60: 291 - 312." type="journal article" year="2015">
Halliday
<emphasis id="B93C7413FFFEFFFEEFAAFD7743A207B0" box="[695,755,657,679]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2015
</bibRefCitation>
)
</taxonomicName>
. However,
<taxonomicName id="4C48D382FFFEFFFEEC54FD5540F407D3" authorityName="Marsh" authorityYear="1869" box="[329,421,688,709]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="hartii">
<emphasis id="B93C7413FFFEFFFEEC54FD55400807D3" box="[329,345,688,709]" italics="true" pageId="21" pageNumber="573">S</emphasis>
.
<emphasis id="B93C7413FFFEFFFEEC79FD5540F407D3" box="[356,421,688,709]" italics="true" pageId="21" pageNumber="573">hartii</emphasis>
</taxonomicName>
is missing from both analyses.
</paragraph>
<paragraph id="8BF7A801FFFEFFFEEDB7FD2A40950101" blockId="21.[146,763,197,1047]" pageId="21" pageNumber="573">
As discussed above,
<taxonomicName id="4C48D382FFFEFFFEECAFFD2B431B07F2" box="[434,586,718,740]" class="Reptilia" family="Elosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Elosuchidae</taxonomicName>
was found to be the sister-taxon of the
<taxonomicName id="4C48D382FFFEFFFEECD0FD0843230615" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[461,626,749,771]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
.
<taxonomicName id="4C48D382FFFEFFFEEF62FD0840610637" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="thailandicus">
<emphasis id="B93C7413FFFEFFFEEF62FD0840610637" italics="true" pageId="21" pageNumber="573">Chalawan thailandicus</emphasis>
</taxonomicName>
is the only putative pholidosaurid that is recovered outside
<taxonomicName id="4C48D382FFFEFFFEECA0FCCF431D0656" authorityName="BUFFETAUT" authorityYear="1982" box="[445,588,810,832]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
in the current phylogenetic analysis. It is sister to another Thai taxon,
<taxonomicName id="4C48D382FFFEFFFEEDEDFC8D4044068D" authority="Lauprasert et al., 2007" authorityName="Lauprasert" authorityYear="2007" class="Reptilia" family="Goniopholididae" genus="Siamosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="phuphokensis">
<emphasis id="B93C7413FFFEFFFEEDEDFC8D4308066B" box="[240,601,872,893]" italics="true" pageId="21" pageNumber="573">Siamosuchus phuphokensis</emphasis>
Lauprasert
<emphasis id="B93C7413FFFEFFFEED8FFC62419D068D" box="[146,204,902,924]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2007
</taxonomicName>
, and together they form a clade with the Chinese
<taxonomicName id="4C48D382FFFEFFFEEC1CFC4040B606AD" authorityName="Young, 1948 Young" authorityYear="1948" box="[257,487,933,955]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="miaoi">
<emphasis id="B93C7413FFFEFFFEEC1CFC4040B606AD" box="[257,487,933,955]" italics="true" pageId="21" pageNumber="573">Sunosuchus miaoi</emphasis>
</taxonomicName>
. The affinities of the several goniopholidid taxa are complex, but there are certain groups that are more closely related to the tethysuchians than others.
</paragraph>
<paragraph id="8BF7A801FFFEFFFEEC70FBB6434E017D" blockId="21.[365,543,1107,1132]" box="[365,543,1107,1132]" pageId="21" pageNumber="573">BIOGEOGRAPHY</paragraph>
<paragraph id="8BF7A801FFFEFFFEED8FFB99444E04C6" blockId="21.[146,762,1148,1906]" lastBlockId="21.[810,1426,197,1905]" pageId="21" pageNumber="573">
Regarding the biogeography of the genus
<taxonomicName id="4C48D382FFFEFFFEEF76FB9943A80187" box="[619,761,1148,1169]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEEF76FB9943A80187" box="[619,761,1148,1169]" italics="true" pageId="21" pageNumber="573">Sarcosuchus</emphasis>
</taxonomicName>
and the related species, some hypotheses are proposed. The phylogenetic inference made by
<bibRefCitation id="EFD9D5F0FFFEFFFEEF41FB5C418E01F8" author="Fortier D &amp; Perea D &amp; Schultz C" pageId="21" pageNumber="573" pagination="272" refId="ref21024" refString="Fortier D, Perea D, Schultz C. 2011. Redescription and phylogenetic relationships of Meridiosaurus valliparadisi, a pholidosaurid from the Late Jurassic of Uruguay. Zoological Journal of the Linnean Society 163: S 257 - S 272." type="journal article" year="2011">
Fortier
<emphasis id="B93C7413FFFEFFFEEFA6FB5F43AB01D8" box="[699,762,1209,1231]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
(2011)
</bibRefCitation>
resulted in the following topology for the group: (Thalattosuchia (
<taxonomicName id="4C48D382FFFEFFFEECBFFB13436F001A" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[418,574,1270,1292]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
((
<taxonomicName id="4C48D382FFFEFFFEEF48FB1343AB001D" authorityName="Meyer" authorityYear="1841" box="[597,762,1270,1291]" class="Reptilia" family="Pholidosauridae" genus="Pholidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEEF48FB1343AB001D" box="[597,762,1270,1291]" italics="true" pageId="21" pageNumber="573">Pholidosaurus</emphasis>
</taxonomicName>
(
<taxonomicName id="4C48D382FFFEFFFEED81FAF04012003D" box="[156,323,1301,1323]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEED81FAF04012003D" box="[156,323,1301,1323]" italics="true" pageId="21" pageNumber="573">Sarcosuchus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C48D382FFFEFFFEEC46FAF3435E003D" baseAuthorityName="Buffetaut &amp; Wellnhofer" baseAuthorityYear="1980" box="[347,527,1302,1323]" class="Reptilia" family="Teleosauridae" genus="Terminonaris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEEC46FAF3435E003D" box="[347,527,1302,1323]" italics="true" pageId="21" pageNumber="573">Terminonaris</emphasis>
</taxonomicName>
)) (
<taxonomicName id="4C48D382FFFEFFFEEF22FAF043AB003D" box="[575,762,1301,1323]" class="Reptilia" family="Pholidosauridae" genus="Oceanosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEEF22FAF043AB003D" box="[575,762,1301,1323]" italics="true" pageId="21" pageNumber="573">Oceanosuchus</emphasis>
</taxonomicName>
(
<emphasis id="B93C7413FFFEFFFEED87FAD1409B005F" box="[154,458,1332,1353]" italics="true" pageId="21" pageNumber="573">
<taxonomicName id="4C48D382FFFEFFFEED87FAD14017005F" authorityName="Mones" authorityYear="1980" box="[154,326,1332,1353]" class="Reptilia" family="Pholidosauridae" genus="Meridiosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">Meridiosaurus</taxonomicName>
,
<taxonomicName id="4C48D382FFFEFFFEEC4EFAD1409B005F" authorityName="de Lapparent de Broin" authorityYear="2002" box="[339,458,1332,1353]" class="Reptilia" family="Pholidosauridae" genus="Elosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">Elosuchus</taxonomicName>
</emphasis>
))))). Based on this result, together with the occurrence of
<taxonomicName id="4C48D382FFFEFFFEEF18FAB740690091" authority="(Owen, 1884)" baseAuthorityName="Owen" baseAuthorityYear="1884" class="Reptilia" family="Pholidosauridae" genus="Anglosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="geofroyi">
<emphasis id="B93C7413FFFEFFFEEF18FAB743A8007E" box="[517,761,1362,1384]" italics="true" pageId="21" pageNumber="573">Anglosuchus geofroyi</emphasis>
(Owen, 1884)
</taxonomicName>
and
<taxonomicName id="4C48D382FFFEFFFEEC63FA9443E50091" authority="(Owen, 1884)" baseAuthorityName="Owen" baseAuthorityYear="1884" box="[382,692,1393,1415]" class="Reptilia" family="Pholidosauridae" genus="Anglosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="laticeps">
<emphasis id="B93C7413FFFEFFFEEC63FA9443530090" box="[382,514,1393,1414]" italics="true" pageId="21" pageNumber="573">A. laticeps</emphasis>
(Owen, 1884)
</taxonomicName>
from the Bathonian of
<collectingCountry id="F35FE891FFFEFFFEEC68FA75408F00B0" box="[373,478,1424,1446]" name="United Kingdom" pageId="21" pageNumber="573">England</collectingCountry>
(both in Mook, 1942), and
<taxonomicName id="4C48D382FFFEFFFEEDD7FA4B43E400D2" authority="Jourdan, 1857" authorityName="Jourdan" authorityYear="1857" box="[202,693,1454,1476]" class="Reptilia" family="Pholidosauridae" genus="Crocodilaemus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="robustus">
<emphasis id="B93C7413FFFEFFFEEDD7FA4B40A600D5" box="[202,503,1454,1475]" italics="true" pageId="21" pageNumber="573">Crocodilaemus robustus</emphasis>
Jourdan, 1857
</taxonomicName>
from the Kimmeridgian of
<collectingCountry id="F35FE891FFFEFFFEEC82FA2B40A400F5" box="[415,501,1486,1507]" name="France" pageId="21" pageNumber="573">France</collectingCountry>
,
<bibRefCitation id="EFD9D5F0FFFEFFFEEF19FA2843A800F5" author="Fortier D &amp; Perea D &amp; Schultz C" box="[516,761,1485,1507]" pageId="21" pageNumber="573" pagination="272" refId="ref21024" refString="Fortier D, Perea D, Schultz C. 2011. Redescription and phylogenetic relationships of Meridiosaurus valliparadisi, a pholidosaurid from the Late Jurassic of Uruguay. Zoological Journal of the Linnean Society 163: S 257 - S 272." type="journal article" year="2011">
Fortier
<emphasis id="B93C7413FFFEFFFEEF79FA2B43F200F4" box="[612,675,1485,1507]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
(2011)
</bibRefCitation>
defends that the common ancestor of
<taxonomicName id="4C48D382FFFEFFFEEF45FA0943A80314" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[600,761,1516,1538]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
and
<taxonomicName id="4C48D382FFFEFFFEEDD7F9EF40C20336" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[202,403,1546,1568]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
is from the Middle Jurassic of Europe. However, the oldest known fossil record for
<taxonomicName id="4C48D382FFFEFFFEEDDEF9AD40C60348" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[195,407,1608,1630]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
(e.g. Caroll, 1988;
<bibRefCitation id="EFD9D5F0FFFEFFFEEF82F9AD4047036A" author="Fortier D &amp; Perea D &amp; Schultz C" pageId="21" pageNumber="573" pagination="272" refId="ref21024" refString="Fortier D, Perea D, Schultz C. 2011. Redescription and phylogenetic relationships of Meridiosaurus valliparadisi, a pholidosaurid from the Late Jurassic of Uruguay. Zoological Journal of the Linnean Society 163: S 257 - S 272." type="journal article" year="2011">
Fortier
<emphasis id="B93C7413FFFEFFFEED8FF982419C036D" box="[146,205,1638,1660]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2011
</bibRefCitation>
) are species in need of redescriptions to elucidate their relationship with
<taxonomicName id="4C48D382FFFEFFFEEF33F96043A4038D" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[558,757,1669,1691]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
. Therefore, those species were not included in the phylogenetic and biogeographical discussions. Also,
<bibRefCitation id="EFD9D5F0FFFEFFFEED8FF90440D403E1" author="Fortier D &amp; Perea D &amp; Schultz C" box="[146,389,1761,1783]" pageId="21" pageNumber="573" pagination="272" refId="ref21024" refString="Fortier D, Perea D, Schultz C. 2011. Redescription and phylogenetic relationships of Meridiosaurus valliparadisi, a pholidosaurid from the Late Jurassic of Uruguay. Zoological Journal of the Linnean Society 163: S 257 - S 272." type="journal article" year="2011">
Fortier
<emphasis id="B93C7413FFFEFFFEEDECF907407E03E0" box="[241,303,1761,1783]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
(2011)
</bibRefCitation>
defends that
<taxonomicName id="4C48D382FFFEFFFEEF29F90443A803E1" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[564,761,1761,1783]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
remained in Europe until the Late Cenomanian with
<taxonomicName id="4C48D382FFFEFFFEED8FF8FA43930223" authority="(Buffetaut &amp; Wellnhofer, 1980)" baseAuthorityName="Buffetaut &amp; Wellnhofer" baseAuthorityYear="1980" box="[146,706,1822,1845]" class="Reptilia" family="Teleosauridae" genus="Terminonaris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEED8FF8FA40660222" box="[146,311,1823,1844]" italics="true" pageId="21" pageNumber="573">Terminonaris</emphasis>
(Buffetaut &amp; Wellnhofer, 1980)
</taxonomicName>
and points out three dispersion routes for the clade: (1) a dispersion for North Africa and eastern South America during ToarcianKimmeridgian, which results on the occurrence of
<taxonomicName id="4C48D382FFFEFFFEE91CFF0145A405EC" box="[1025,1269,228,250]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B93C7413FFFEFFFEE91CFF0145C705EF" box="[1025,1174,228,249]" italics="true" pageId="21" pageNumber="573">Sarcosuchus</emphasis>
species
</taxonomicName>
, (2) a second dispersion for Africa and South America during KimmeridgianLate Albian explaining, respectively, the species
<taxonomicName id="4C48D382FFFEFFFEEEACFEA545760443" authorityName="de Lapparent de Broin" authorityYear="2002" box="[945,1063,320,341]" class="Reptilia" family="Pholidosauridae" genus="Elosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEEEACFEA545760443" box="[945,1063,320,341]" italics="true" pageId="21" pageNumber="573">Elosuchus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C48D382FFFEFFFEE97FFEA5445A0443" authorityName="Mones" authorityYear="1980" box="[1122,1291,320,341]" class="Reptilia" family="Pholidosauridae" genus="Meridiosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE97FFEA5445A0443" box="[1122,1291,320,341]" italics="true" pageId="21" pageNumber="573">Meridiosaurus</emphasis>
</taxonomicName>
and (3) the last dispersion proposed was between the
<taxonomicName id="4C48D382FFFEFFFEE9E5FEBA422B0485" class="Reptilia" family="Teleosauridae" genus="Terminonaris" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B93C7413FFFEFFFEE9E5FEBA44C30462" box="[1272,1426,351,372]" italics="true" pageId="21" pageNumber="573">Terminonaris</emphasis>
species
</taxonomicName>
from Europe to North America during the Late CenomanianEarly Turonian, being North America the last place inhabited by
<taxonomicName id="4C48D382FFFEFFFEE942FE5F444A04C6" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[1119,1307,442,464]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
.
</paragraph>
<paragraph id="8BF7A801FFFEFFFEEE5FFE3C456D06EE" blockId="21.[810,1426,197,1905]" pageId="21" pageNumber="573">
The phylogenetic inference made by Martin
<emphasis id="B93C7413FFFEFFFEE84BFE3F44C304F8" box="[1366,1426,473,495]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
(2014) resulted in the following relationship of
<taxonomicName id="4C48D382FFFEFFFEEE37FDF342BA073A" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[810,1003,534,556]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
and its sister-group: (
<taxonomicName id="4C48D382FFFEFFFEE9E8FDF3428B075C" class="Reptilia" family="Goniopholididae" genus="Siamosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="phuphokensis">
<emphasis id="B93C7413FFFEFFFEE9E8FDF3428B075C" italics="true" pageId="21" pageNumber="573">Siamosuchus phuphokensis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C48D382FFFEFFFEEEF2FDD045B3075D" box="[1007,1250,565,587]" class="Reptilia" family="Goniopholididae" genus="Goniopholis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="simus">
<emphasis id="B93C7413FFFEFFFEEEF2FDD045B3075D" box="[1007,1250,565,587]" italics="true" pageId="21" pageNumber="573">Goniopholis simus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C48D382FFFEFFFEE9EAFDD042B3077F" authorityName="Schwarz" authorityYear="2002" class="Reptilia" family="Goniopholididae" genus="Goniopholis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="baryglyphaeus">
<emphasis id="B93C7413FFFEFFFEE9EAFDD042B3077F" italics="true" pageId="21" pageNumber="573">Goniopholis baryglyphaeus</emphasis>
</taxonomicName>
(
<taxonomicName id="4C48D382FFFEFFFEEEE5FDB1458F077F" box="[1016,1246,596,617]" class="Reptilia" family="Pholidosauridae" genus="Pholidosaurus" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B93C7413FFFEFFFEEEE5FDB145FE077F" box="[1016,1199,596,617]" italics="true" pageId="21" pageNumber="573">Pholidosaurus</emphasis>
sp.
</taxonomicName>
(
<taxonomicName id="4C48D382FFFEFFFEE9E9FDB142FB079E" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="imperator">
<emphasis id="B93C7413FFFEFFFEE9E9FDB142FB079E" italics="true" pageId="21" pageNumber="573">Sarcosuchus imperator</emphasis>
</taxonomicName>
,
<taxonomicName id="4C48D382FFFEFFFEEEDDFD9745A90791" box="[960,1272,626,647]" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="thailandicus">
<emphasis id="B93C7413FFFEFFFEEEDDFD9745A90791" box="[960,1272,626,647]" italics="true" pageId="21" pageNumber="573">Chalawan thailandicus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C48D382FFFEFFFEE813FD9742E707B0" baseAuthorityName="de Broin" baseAuthorityYear="2002" class="Reptilia" family="Pholidosauridae" genus="Elosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="cherifiensis">
<emphasis id="B93C7413FFFEFFFEE813FD9742E707B0" italics="true" pageId="21" pageNumber="573">Elosuchus cherifiensis</emphasis>
</taxonomicName>
))). Based on this result, Martin
<emphasis id="B93C7413FFFEFFFEE84EFD7744C307B0" box="[1363,1426,657,679]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
(2014) proposed a worldwide distribution for the group in the Late JurassicEarly Cretaceous interval, probably resulting from the conquest of north and south portions of the Tethys Ocean, which connect these regions. Also, they observe the relationship of
<taxonomicName id="4C48D382FFFEFFFEEE37FCAC45190648" box="[810,1096,841,862]" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="thailandicus">
<emphasis id="B93C7413FFFEFFFEEE37FCAC45190648" box="[810,1096,841,862]" italics="true" pageId="21" pageNumber="573">Chalawan thailandicus</emphasis>
</taxonomicName>
from
<collectingCountry id="F35FE891FFFEFFFEE98EFCAC44500649" box="[1171,1281,841,863]" name="Thailand" pageId="21" pageNumber="573">Thailand</collectingCountry>
with South America and Africa as an exception for the region due to dispersal events during the JurassicAptian, because all other strata and crocodilian records from
<collectingCountry id="F35FE891FFFEFFFEEE37FC2142C506CC" box="[810,916,964,986]" name="Thailand" pageId="21" pageNumber="573">Thailand</collectingCountry>
are more closely related to the Asian fauna (Fernandez
<emphasis id="B93C7413FFFEFFFEEEACFC0642B806E1" box="[945,1001,994,1016]" italics="true" pageId="21" pageNumber="573">et al.</emphasis>
, 2010).
</paragraph>
<paragraph id="8BF7A801FFFEFFFEEE5FFBE442C0003D" blockId="21.[810,1426,197,1905]" pageId="21" pageNumber="573">
Both analyses discussed above include species that are not present in the phylogenetic inference (e.g.
<taxonomicName id="4C48D382FFFEFFFEEE37FBDB45540145" baseAuthorityName="Campos &amp; Kellner" baseAuthorityYear="1991" box="[810,1029,1086,1107]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="hartti">
<emphasis id="B93C7413FFFEFFFEEE37FBDB45540145" box="[810,1029,1086,1107]" italics="true" pageId="21" pageNumber="573">Sarcosuchus hartti</emphasis>
</taxonomicName>
), which weakens the methodology of inferring biogeographical hypotheses from topology in a consensus cladogram. Based on the present topology hypotheses (
<figureCitation id="1373B484FFFEFFFEE935FB7F452901B9" box="[1064,1144,1178,1200]" captionStart="Figure 11" captionStartId="22.[142,220,1551,1573]" captionTargetBox="[144,1831,459,1507]" captionTargetId="figure-28@22.[143,1839,457,1511]" captionTargetPageId="22" captionText="Figure 11. Detail of the Tethysuchia clade and their sister-clade from the consensus cladogram. The two phylogenetic hypotheses that originated the polytomy in the consensus cladogram: A, hypothesis two: Sunosuchus shartegensis, Calsoyasuchus, Sunosuchus miaoi, Siamosuchus phuphokensis and Chalawan thailandicus as a clade sister to Tethysuchia. B, hypothesis one: Sunosuchus shartegensis and Calsoyasuchus as sister-group of Tethysuchia." figureDoi="http://doi.org/10.5281/zenodo.5719186" httpUri="https://zenodo.org/record/5719186/files/figure.png" pageId="21" pageNumber="573">Fig. 11</figureCitation>
), and in the calibration of the specimens analysed, a new biogeographical hypothesis with be proposed based on new evidence to better explain the distribution of
<taxonomicName id="4C48D382FFFEFFFEE9D3FB13440E001D" box="[1230,1375,1270,1291]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE9D3FB13440E001D" box="[1230,1375,1270,1291]" italics="true" pageId="21" pageNumber="573">Sarcosuchus</emphasis>
</taxonomicName>
and its allies.
</paragraph>
<paragraph id="8BF7A801FFFEFFFCEE5FFAD6407505EA" blockId="21.[810,1426,197,1905]" lastBlockId="23.[146,762,199,1570]" lastPageId="23" lastPageNumber="575" pageId="21" pageNumber="573">
The present phylogenetic analysis results in two biogeographical hypotheses for the species in the clade ((
<taxonomicName id="4C48D382FFFEFFFEEE99FA9445E50090" authorityName="Efimov" authorityYear="1988" box="[900,1204,1393,1414]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="shartegensis">
<emphasis id="B93C7413FFFEFFFEEE99FA9445E50090" box="[900,1204,1393,1414]" italics="true" pageId="21" pageNumber="573">Sunosuchus shartegensis</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFEFFFEE9CAFA9444D80090" box="[1239,1417,1393,1414]" class="Reptilia" family="Goniopholididae" genus="Calsoyasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE9CAFA9444D80090" box="[1239,1417,1393,1414]" italics="true" pageId="21" pageNumber="573">Calsoyasuchus</emphasis>
</taxonomicName>
) (
<taxonomicName id="4C48D382FFFEFFFEEE2EFA6A454900B3" authorityName="Young, 1948 Young" authorityYear="1948" box="[819,1048,1423,1445]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="miaoi">
<emphasis id="B93C7413FFFEFFFEEE2EFA6A454900B3" box="[819,1048,1423,1445]" italics="true" pageId="21" pageNumber="573">Sunosuchus miaoi</emphasis>
</taxonomicName>
(
<taxonomicName id="4C48D382FFFEFFFEE930FA6A458100B2" box="[1069,1232,1423,1444]" class="Reptilia" family="Goniopholididae" genus="Siamosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE930FA6A458100B2" box="[1069,1232,1423,1444]" italics="true" pageId="21" pageNumber="573">Siamosuchus</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFEFFFEE9EAFA6A442700B2" box="[1271,1398,1423,1444]" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE9EAFA6A442700B2" box="[1271,1398,1423,1444]" italics="true" pageId="21" pageNumber="573">Chalawan</emphasis>
</taxonomicName>
))) (
<taxonomicName id="4C48D382FFFEFFFEEE2FFA4B429800D2" authorityName="BUFFETAUT" authorityYear="1982" box="[818,969,1454,1476]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
)). The first hypothesis is observed in minimum-length trees 1, 3, 5, 6, 7, 9 and 11 (
<figureCitation id="1373B484FFFEFFFEE838FA2844D200F4" box="[1317,1411,1485,1507]" captionStart="Figure 11" captionStartId="22.[142,220,1551,1573]" captionTargetBox="[144,1831,459,1507]" captionTargetId="figure-28@22.[143,1839,457,1511]" captionTargetPageId="22" captionText="Figure 11. Detail of the Tethysuchia clade and their sister-clade from the consensus cladogram. The two phylogenetic hypotheses that originated the polytomy in the consensus cladogram: A, hypothesis two: Sunosuchus shartegensis, Calsoyasuchus, Sunosuchus miaoi, Siamosuchus phuphokensis and Chalawan thailandicus as a clade sister to Tethysuchia. B, hypothesis one: Sunosuchus shartegensis and Calsoyasuchus as sister-group of Tethysuchia." figureDoi="http://doi.org/10.5281/zenodo.5719186" httpUri="https://zenodo.org/record/5719186/files/figure.png" pageId="21" pageNumber="573">Fig. 11A</figureCitation>
), which presents the following topological relationship ((
<taxonomicName id="4C48D382FFFEFFFEEE20F9EF45730336" authorityName="Young, 1948 Young" authorityYear="1948" box="[829,1058,1546,1568]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="miaoi">
<emphasis id="B93C7413FFFEFFFEEE20F9EF45730336" box="[829,1058,1546,1568]" italics="true" pageId="21" pageNumber="573">Sunosuchus miaoi</emphasis>
</taxonomicName>
(
<taxonomicName id="4C48D382FFFEFFFEE92AF9EF458B0309" box="[1079,1242,1546,1567]" class="Reptilia" family="Goniopholididae" genus="Siamosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE92AF9EF458B0309" box="[1079,1242,1546,1567]" italics="true" pageId="21" pageNumber="573">Siamosuchus</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFEFFFEE81DF9EF442E0309" box="[1280,1407,1546,1567]" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE81DF9EF442E0309" box="[1280,1407,1546,1567]" italics="true" pageId="21" pageNumber="573">Chalawan</emphasis>
</taxonomicName>
)) ((
<taxonomicName id="4C48D382FFFEFFFEEE22F9CC45C50328" authorityName="Efimov" authorityYear="1988" box="[831,1172,1577,1598]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="shartegensis">
<emphasis id="B93C7413FFFEFFFEEE22F9CC45C50328" box="[831,1172,1577,1598]" italics="true" pageId="21" pageNumber="573">Sunosuchus shartegensis</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFEFFFEE9DFF9CC44D60328" box="[1218,1415,1577,1598]" class="Reptilia" family="Goniopholididae" genus="Calsoyasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE9DFF9CC44D60328" box="[1218,1415,1577,1598]" italics="true" pageId="21" pageNumber="573">Calsoyasuchus</emphasis>
</taxonomicName>
) (
<taxonomicName id="4C48D382FFFEFFFEEE2FF9A24298034B" authorityName="BUFFETAUT" authorityYear="1982" box="[818,969,1607,1629]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
))). In this phylogenetic scenario, the clade (
<taxonomicName id="4C48D382FFFEFFFEEE6BF983451B036A" authorityName="Young, 1948 Young" authorityYear="1948" box="[886,1098,1638,1660]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="miaoi">
<emphasis id="B93C7413FFFEFFFEEE6BF983451B036A" box="[886,1098,1638,1660]" italics="true" pageId="21" pageNumber="573">Sunosuchus miaoi</emphasis>
</taxonomicName>
(
<taxonomicName id="4C48D382FFFEFFFEE947F98345A2036D" box="[1114,1267,1638,1659]" class="Reptilia" family="Goniopholididae" genus="Siamosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE947F98345A2036D" box="[1114,1267,1638,1659]" italics="true" pageId="21" pageNumber="573">Siamosuchus</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFEFFFEE80FF98344DB036D" box="[1298,1418,1638,1659]" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE80FF98344DB036D" box="[1298,1418,1638,1659]" italics="true" pageId="21" pageNumber="573">Chalawan</emphasis>
</taxonomicName>
) is composed of Asiatic fluvial species from the Late Jurassic to the Early Cretaceous, with the Thai species the sister to the Chinese species. The clade ((
<taxonomicName id="4C48D382FFFEFFFEEE99F90445E503E0" authorityName="Efimov" authorityYear="1988" box="[900,1204,1761,1782]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="species" species="shartegensis">
<emphasis id="B93C7413FFFEFFFEEE99F90445E503E0" box="[900,1204,1761,1782]" italics="true" pageId="21" pageNumber="573">Sunosuchus shartegensis</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFEFFFEE9CAF90444D803E0" box="[1239,1417,1761,1782]" class="Reptilia" family="Goniopholididae" genus="Calsoyasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFEFFFEE9CAF90444D803E0" box="[1239,1417,1761,1782]" italics="true" pageId="21" pageNumber="573">Calsoyasuchus</emphasis>
</taxonomicName>
) (
<taxonomicName id="4C48D382FFFEFFFEEE2FF91A42980203" authorityName="BUFFETAUT" authorityYear="1982" box="[818,969,1791,1813]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="21" pageNumber="573" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
)) has a more complex biogeographic scenario where two fluvial species from the Jurassic of Mongolia and North America are sister species. This phylogenetic context is difficult to explain in a biogeographical scenario for the common ancestor of
<taxonomicName id="4C48D382FFFCFFFCED8FFF03407005EA" authorityName="BUFFETAUT" authorityYear="1982" box="[146,289,230,252]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
.
</paragraph>
<caption id="DF37F889FFFDFFFDED93F9EA44ED0376" ID-DOI="http://doi.org/10.5281/zenodo.5719186" ID-Zenodo-Dep="5719186" httpUri="https://zenodo.org/record/5719186/files/figure.png" pageId="22" pageNumber="574" startId="22.[142,220,1551,1573]" targetBox="[144,1831,459,1507]" targetPageId="22">
<paragraph id="8BF7A801FFFDFFFDED93F9EA44ED0376" blockId="22.[142,1840,1551,1632]" pageId="22" pageNumber="574">
<emphasis id="B93C7413FFFDFFFDED93F9EA40520333" bold="true" box="[142,259,1551,1573]" pageId="22" pageNumber="574">Figure 11.</emphasis>
Detail of the
<taxonomicName id="4C48D382FFFDFFFDEC8BF9EA43470333" authorityName="BUFFETAUT" authorityYear="1982" box="[406,534,1551,1573]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
clade and their sister-clade from the consensus cladogram. The two phylogenetic hypotheses that originated the polytomy in the consensus cladogram: A, hypothesis two:
<taxonomicName id="4C48D382FFFDFFFDEF7EF9C942390354" authorityName="Efimov" authorityYear="1988" box="[611,872,1580,1602]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="species" species="shartegensis">
<emphasis id="B93C7413FFFDFFFDEF7EF9C942390354" box="[611,872,1580,1602]" italics="true" pageId="22" pageNumber="574">Sunosuchus shartegensis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C48D382FFFDFFFDEE69F9C9455F0354" box="[884,1038,1580,1602]" class="Reptilia" family="Goniopholididae" genus="Calsoyasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFDFFFDEE69F9C9455F0354" box="[884,1038,1580,1602]" italics="true" pageId="22" pageNumber="574">Calsoyasuchus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C48D382FFFDFFFDE907F9C9458B0354" authorityName="Young, 1948 Young" authorityYear="1948" box="[1050,1242,1580,1602]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="species" species="miaoi">
<emphasis id="B93C7413FFFDFFFDE907F9C9458B0354" box="[1050,1242,1580,1602]" italics="true" pageId="22" pageNumber="574">Sunosuchus miaoi</emphasis>
</taxonomicName>
,
<taxonomicName id="4C48D382FFFDFFFDE9F8F9C947540354" box="[1253,1541,1580,1602]" class="Reptilia" family="Goniopholididae" genus="Siamosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="species" species="phuphokensis">
<emphasis id="B93C7413FFFDFFFDE9F8F9C947540354" box="[1253,1541,1580,1602]" italics="true" pageId="22" pageNumber="574">Siamosuchus phuphokensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C48D382FFFDFFFDEB24F9C946610354" box="[1593,1840,1580,1602]" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="species" species="thailandicus">
<emphasis id="B93C7413FFFDFFFDEB24F9C946610354" box="[1593,1840,1580,1602]" italics="true" pageId="22" pageNumber="574">Chalawan thailandicus</emphasis>
</taxonomicName>
as a clade sister to
<taxonomicName id="4C48D382FFFDFFFDEC48F9AF40860376" authorityName="BUFFETAUT" authorityYear="1982" box="[341,471,1610,1632]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
. B, hypothesis one:
<taxonomicName id="4C48D382FFFDFFFDEFBEF9AF42F90376" authorityName="Efimov" authorityYear="1988" box="[675,936,1610,1632]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="species" species="shartegensis">
<emphasis id="B93C7413FFFDFFFDEFBEF9AF42F90376" box="[675,936,1610,1632]" italics="true" pageId="22" pageNumber="574">Sunosuchus shartegensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C48D382FFFDFFFDEEC6F9AF45240376" box="[987,1141,1610,1632]" class="Reptilia" family="Goniopholididae" genus="Calsoyasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFDFFFDEEC6F9AF45240376" box="[987,1141,1610,1632]" italics="true" pageId="22" pageNumber="574">Calsoyasuchus</emphasis>
</taxonomicName>
as sister-group of
<taxonomicName id="4C48D382FFFDFFFDE828F9AF44E90376" authorityName="BUFFETAUT" authorityYear="1982" box="[1333,1464,1610,1632]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="22" pageNumber="574" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8BF7A801FFFCFFFCEDB7FEE040BF077A" blockId="23.[146,762,199,1570]" pageId="23" pageNumber="575">
The second hypothesis is based on the topological relationship observed in the minimum-length trees 2, 4, 8, 10 and 12 (
<figureCitation id="1373B484FFFCFFFCEC73FEA740800441" box="[366,465,322,344]" captionStart="Figure 11" captionStartId="22.[142,220,1551,1573]" captionTargetBox="[144,1831,459,1507]" captionTargetId="figure-28@22.[143,1839,457,1511]" captionTargetPageId="22" captionText="Figure 11. Detail of the Tethysuchia clade and their sister-clade from the consensus cladogram. The two phylogenetic hypotheses that originated the polytomy in the consensus cladogram: A, hypothesis two: Sunosuchus shartegensis, Calsoyasuchus, Sunosuchus miaoi, Siamosuchus phuphokensis and Chalawan thailandicus as a clade sister to Tethysuchia. B, hypothesis one: Sunosuchus shartegensis and Calsoyasuchus as sister-group of Tethysuchia." figureDoi="http://doi.org/10.5281/zenodo.5719186" httpUri="https://zenodo.org/record/5719186/files/figure.png" pageId="23" pageNumber="575">Fig. 11B</figureCitation>
), which is (((
<taxonomicName id="4C48D382FFFCFFFCEF70FEA740790460" authorityName="Efimov" authorityYear="1988" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="species" species="shartegensis">
<emphasis id="B93C7413FFFCFFFCEF70FEA740790460" italics="true" pageId="23" pageNumber="575">Sunosuchus shartegensis</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFCFFFCEC56FE8440AF0460" box="[331,510,353,374]" class="Reptilia" family="Goniopholididae" genus="Calsoyasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCEC56FE8440AF0460" box="[331,510,353,374]" italics="true" pageId="23" pageNumber="575">Calsoyasuchus</emphasis>
</taxonomicName>
) (
<taxonomicName id="4C48D382FFFCFFFCEF07FE8443A80460" authorityName="Young, 1948 Young" authorityYear="1948" box="[538,761,353,374]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="species" species="miaoi">
<emphasis id="B93C7413FFFCFFFCEF07FE8443A80460" box="[538,761,353,374]" italics="true" pageId="23" pageNumber="575">Sunosuchus miaoi</emphasis>
</taxonomicName>
(
<taxonomicName id="4C48D382FFFCFFFCED86FE9A406C0482" box="[155,317,383,404]" class="Reptilia" family="Goniopholididae" genus="Siamosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCED86FE9A406C0482" box="[155,317,383,404]" italics="true" pageId="23" pageNumber="575">Siamosuchus</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFCFFFCEC7FFE9A40B10482" box="[354,480,383,404]" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCEC7FFE9A40B10482" box="[354,480,383,404]" italics="true" pageId="23" pageNumber="575">Chalawan</emphasis>
</taxonomicName>
))) (
<taxonomicName id="4C48D382FFFCFFFCEF0DFE9A43FA0483" authorityName="BUFFETAUT" authorityYear="1982" box="[528,683,383,405]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
)). The biogeographical hypothesis for the common ancestor of
<taxonomicName id="4C48D382FFFCFFFCEDA9FE58400104C5" authorityName="BUFFETAUT" authorityYear="1982" box="[180,336,445,467]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
remains doubtful. However, the main difference from the first hypothesis is the clade ((
<taxonomicName id="4C48D382FFFCFFFCEDF1FE1F434D0719" authorityName="Efimov" authorityYear="1988" box="[236,540,506,527]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="species" species="shartegensis">
<emphasis id="B93C7413FFFCFFFCEDF1FE1F434D0719" box="[236,540,506,527]" italics="true" pageId="23" pageNumber="575">Sunosuchus shartegensis</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFCFFFCEF22FE1F43A00719" box="[575,753,506,527]" class="Reptilia" family="Goniopholididae" genus="Calsoyasuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCEF22FE1F43A00719" box="[575,753,506,527]" italics="true" pageId="23" pageNumber="575">Calsoyasuchus</emphasis>
</taxonomicName>
) (
<taxonomicName id="4C48D382FFFCFFFCED86FDFC40D00738" authorityName="Young, 1948 Young" authorityYear="1948" box="[155,385,537,558]" class="Reptilia" family="Goniopholididae" genus="Sunosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="species" species="miaoi">
<emphasis id="B93C7413FFFCFFFCED86FDFC40D00738" box="[155,385,537,558]" italics="true" pageId="23" pageNumber="575">Sunosuchus miaoi</emphasis>
</taxonomicName>
(
<taxonomicName id="4C48D382FFFCFFFCEC88FDFC43690738" box="[405,568,537,558]" class="Reptilia" family="Goniopholididae" genus="Siamosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCEC88FDFC43690738" box="[405,568,537,558]" italics="true" pageId="23" pageNumber="575">Siamosuchus</emphasis>
</taxonomicName>
+
<taxonomicName id="4C48D382FFFCFFFCEF42FDFC438F0738" box="[607,734,537,558]" class="Reptilia" family="Pholidosauridae" genus="Chalawan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCEF42FDFC438F0738" box="[607,734,537,558]" italics="true" pageId="23" pageNumber="575">Chalawan</emphasis>
</taxonomicName>
))) representing an Asian clade with probably a posterior colonization of North America.
</paragraph>
<paragraph id="8BF7A801FFFCFFFCEDB7FD90409C007C" blockId="23.[146,762,199,1570]" pageId="23" pageNumber="575">
Even though it is complex to infer a biogeographical hypothesis for the common ancestor of
<taxonomicName id="4C48D382FFFCFFFCEF7EFD7643A407BF" authorityName="BUFFETAUT" authorityYear="1982" box="[611,757,659,681]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
, its two clades have much less complicated scenarios. The
<taxonomicName id="4C48D382FFFCFFFCEDD6FD3440C407F1" baseAuthorityName="von Zittel &amp; Eastman" baseAuthorityYear="1902" box="[203,405,721,743]" class="Reptilia" family="Pholidosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="family">Pholidosauridae</taxonomicName>
clade contains species from the Early Cretaceous of Asia and Europe, with the European species related to marine environments and the Asian species related to fluvial/terrestrial environments. However, there is no evidences to infer if the common ancestor of this clade lived in Europe or Asia, nor if it was a marine or terrestrial species. On the other hand, the other clade of
<taxonomicName id="4C48D382FFFCFFFCEF7BFC4243A806AB" authorityName="BUFFETAUT" authorityYear="1982" box="[614,761,935,957]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="subOrder" subOrder="Tethysuchia">Tethysuchia</taxonomicName>
includes both marine and fluvial species from the Late Jurassic to the Early Cretaceous of the Americas and Africa. The first species to diverge is the marine
<taxonomicName id="4C48D382FFFCFFFCED8FFBC740120121" authorityName="Mones" authorityYear="1980" box="[146,323,1058,1079]" class="Reptilia" family="Pholidosauridae" genus="Meridiosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCED8FFBC740120121" box="[146,323,1058,1079]" italics="true" pageId="23" pageNumber="575">Meridiosaurus</emphasis>
</taxonomicName>
from the Late Jurassic of
<collectingCountry id="F35FE891FFFCFFFCEF96FBC743A50121" box="[651,756,1058,1079]" name="Uruguay" pageId="23" pageNumber="575">Uruguay</collectingCountry>
, which is the sister-species of a clade that includes the genus
<taxonomicName id="4C48D382FFFCFFFCEC10FBBA40F10162" box="[269,416,1119,1140]" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCEC10FBBA40F10162" box="[269,416,1119,1140]" italics="true" pageId="23" pageNumber="575">Sarcosuchus</emphasis>
</taxonomicName>
and a clade of
<taxonomicName id="4C48D382FFFCFFFCEF49FBBA43B20163" box="[596,739,1119,1141]" class="Reptilia" family="Elosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="family">Elosuchidae</taxonomicName>
+
<taxonomicName id="4C48D382FFFCFFFCED8FFB9B40630182" baseAuthorityName="de Steffano" baseAuthorityYear="1903" box="[146,306,1150,1172]" class="Reptilia" family="Dyrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="family">Dyrosauridae</taxonomicName>
. In this context both fluvial
<taxonomicName id="4C48D382FFFCFFFCEF74FB9B41BA01A4" class="Reptilia" family="Crocodylidae" genus="Sarcosuchus" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B93C7413FFFCFFFCEF74FB9B43AB0185" box="[617,762,1150,1171]" italics="true" pageId="23" pageNumber="575">Sarcosuchus</emphasis>
species
</taxonomicName>
are from the Early Cretaceous of South America and Africa, being its cladogenesis, probably related with the early events of the break-up of Gondwana, as already proposed in earlier literature. The marine North American species of
<taxonomicName id="4C48D382FFFCFFFCEF41FAFD43A8003B" baseAuthorityName="Buffetaut &amp; Wellnhofer" baseAuthorityYear="1980" box="[604,761,1304,1325]" class="Reptilia" family="Teleosauridae" genus="Terminonaris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCEF41FAFD43A8003B" box="[604,761,1304,1325]" italics="true" pageId="23" pageNumber="575">Terminonaris</emphasis>
</taxonomicName>
seems to be related to the North African taxa, such the ancestors of
<taxonomicName id="4C48D382FFFCFFFCEC4DFAB14097007F" authorityName="de Lapparent de Broin" authorityYear="2002" box="[336,454,1364,1385]" class="Reptilia" family="Pholidosauridae" genus="Elosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="23" pageNumber="575" phylum="Chordata" rank="genus">
<emphasis id="B93C7413FFFCFFFCEC4DFAB14097007F" box="[336,454,1364,1385]" italics="true" pageId="23" pageNumber="575">Elosuchus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BF7A801FFFCFFFCEDB7FA9643F50334" blockId="23.[146,762,199,1570]" pageId="23" pageNumber="575">This new phylogenetic hypotheses enables the discussion of some interesting biogeographical scenarios, but further work is needed to provide better supported biogeographical hypotheses and to test these properly. This study is merely a contribution to future, more complete biogeographical studies.</paragraph>
</subSubSection>
</treatment>
</document>