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<mods:title id="99ADF877EAE5B58A5E86E9B9A5D3734D">A new choristodere (Reptilia: Diapsida) from the Lower Cretaceous of western Liaoning Province, China, and phylogenetic relationships of Monjurosuchidae</mods:title>
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<subSubSection id="C344657F0F1A82059A2AF8EDBFD4FF73" box="[853,1415,194,218]" pageId="11" pageNumber="438" type="nomenclature">
<paragraph id="8BE136F40F1A82059A2AF8EDBFD4FF73" blockId="11.[853,1415,194,218]" box="[853,1415,194,218]" pageId="11" pageNumber="438">
<heading id="D0A981980F1A82059A2AF8EDBFD4FF73" box="[853,1415,194,218]" centered="true" fontSize="9" level="2" pageId="11" pageNumber="438" reason="5">
<taxonomicName id="4C5E4D770F1A82059A2AF8EDBE69FF70" authority="AS A MONOPHYLETIC" authorityName="AS A MONOPHYLETIC" box="[853,1082,194,218]" class="Reptilia" family="Monjurosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">MONJUROSUCHIDAE</taxonomicName>
AS A MONOPHYLETIC FAMILY
</heading>
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<paragraph id="8BE136F40F1A82059A45F8C4BE5CFCA0" blockId="11.[826,1442,235,778]" pageId="11" pageNumber="438">
Two genera and species are currently classified in the family
<taxonomicName id="4C5E4D770F1A82059AF4F926BE00FEB5" box="[907,1107,265,287]" class="Reptilia" family="Monjurosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">Monjurosuchidae</taxonomicName>
: the
<typeStatus id="54E588560F1A82059DF3F925BEEEFEB5" box="[1164,1213,266,287]" pageId="11" pageNumber="438">type</typeStatus>
species
<taxonomicName id="4C5E4D770F1A82059C60F925B9B9FE97" authorityName="Endo" authorityYear="1940" class="Reptilia" family="Simoedosauridae" genus="Monjurosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="species" species="splendens">
<emphasis id="B92AEAE60F1A82059C60F925B9B9FE97" italics="true" pageId="11" pageNumber="438">Monjurosuchus splendens</emphasis>
</taxonomicName>
from the Yixian Formation exposed near Lingyuan, and
<taxonomicName id="4C5E4D770F1A82059D55F968BF19FEF6" authority="Gao &amp; Fox, 2005" authorityName="Gao &amp; Fox" authorityYear="2005" box="[1066,1354,327,348]" class="Reptilia" family="Monjurosuchidae" genus="Philydrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="species" species="proseilus" status="gen. et sp. nov.">
<emphasis id="B92AEAE60F1A82059D55F968BF19FEF6" box="[1066,1354,327,348]" italics="true" pageId="11" pageNumber="438">Philydrosaurus proseilus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A219579D0F1A82059C2BF968B9DFFED1" pageId="11" pageNumber="438" rank="species">gen. et sp. nov.</taxonomicNameLabel>
described here from the Chiufotang Formation exposed near Chaoyang. With inclusion of these two taxa in a revised data set (Appendix 2), a parsimony analysis revealed that the monophyly of the family is supported by the following synapomorphies: dorsal process of squamosal absent (13-2; unknown for
<taxonomicName id="4C5E4D770F1A82059C23F9D0B9F9FD98" class="Reptilia" family="Hyphalosauridae" genus="Hyphalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="genus">
<emphasis id="B92AEAE60F1A82059C23F9D0B9F9FD98" italics="true" pageId="11" pageNumber="438">Hyphalosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5E4D770F1A82059A82FA32BE35FD98" box="[1021,1126,541,562]" class="Reptilia" genus="Shokawa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eosuchia" pageId="11" pageNumber="438" phylum="Chordata" rank="genus">
<emphasis id="B92AEAE60F1A82059A82FA32BE35FD98" box="[1021,1126,541,562]" italics="true" pageId="11" pageNumber="438">Shokawa</emphasis>
</taxonomicName>
); supratemporal fenestra smaller than orbit (14-0; character reversal); infratemporal fenestra closed (15-1; homoplastic in
<taxonomicName id="4C5E4D770F1A82059A45FA56B9B6FD24" authorityName="Hecht" authorityYear="1992" box="[826,997,633,654]" class="Reptilia" genus="Lazarussuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eosuchia" pageId="11" pageNumber="438" phylum="Chordata" rank="genus">
<emphasis id="B92AEAE60F1A82059A45FA56B9B6FD24" box="[826,997,633,654]" italics="true" pageId="11" pageNumber="438">Lazarussuchus</emphasis>
</taxonomicName>
, see
<bibRefCitation id="EFCF4B050F1A82059D3FFA55BFC5FD25" author="Evans SE &amp; Klembara J" box="[1088,1430,633,655]" pageId="11" pageNumber="438" pagination="171 - 184" refId="ref10943" refString="Evans SE, Klembara J. 2005. A choristoderan reptile (Reptilia: Diapsida) from the lower Miocene of northwest Bohemia. Journal of Vertebrate Palaeontology 25: 171 - 184." type="journal article" year="2005">Evans &amp; Klembara, 2005</bibRefCitation>
); post-temporal fenestra absent (47-1; also in
<taxonomicName id="4C5E4D770F1A82059C2CFAB7B9E9FD67" authorityName="Lemone" authorityYear="1884" class="Reptilia" family="Simoedosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">Simoedosauridae</taxonomicName>
); posterior process of ischium strongly developed (75-1; uniquely diagnostic for the family
<taxonomicName id="4C5E4D770F1A82059A45FADBBE57FCA0" box="[826,1028,756,778]" class="Reptilia" family="Monjurosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">Monjurosuchidae</taxonomicName>
).
</paragraph>
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<subSubSection id="C344657F0F1A82059AD0FB74BF56FC3B" pageId="11" pageNumber="438" type="multiple">
<paragraph id="8BE136F40F1A82059AD0FB74BF56FC3B" blockId="11.[943,1325,859,915]" pageId="11" pageNumber="438">
A POSSIBLE
<taxonomicName id="4C5E4D770F1A82059D44FB74BF4CFCDB" box="[1083,1311,859,883]" class="Reptilia" family="Monjurosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">MONJUROSUCHIDAE</taxonomicName>
<taxonomicName id="4C5E4D770F1A82059C60FB74BEE7FC3B" authorityName="Gao &amp; Fox" authorityYear="2005" class="Reptilia" family="Hyphalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">HYPHALOSAURIDAE</taxonomicName>
CLADE
</paragraph>
</subSubSection>
<subSubSection id="C344657F0F1A82029A45FB8CB819FF52" lastPageId="12" lastPageNumber="439" pageId="11" pageNumber="438" type="discussion">
<paragraph id="8BE136F40F1A82059A45FB8CB9F0FA2F" blockId="11.[826,1443,931,1904]" pageId="11" pageNumber="438">
<taxonomicName id="4C5E4D770F1A82059A45FB8CB9BBFC12" box="[826,1000,931,952]" class="Reptilia" family="Hyphalosauridae" genus="Hyphalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="genus">
<emphasis id="B92AEAE60F1A82059A45FB8CB9BBFC12" box="[826,1000,931,952]" italics="true" pageId="11" pageNumber="438">Hyphalosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5E4D770F1A82059D5EFB8CBED9FC12" box="[1057,1162,931,952]" class="Reptilia" genus="Shokawa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eosuchia" pageId="11" pageNumber="438" phylum="Chordata" rank="genus">
<emphasis id="B92AEAE60F1A82059D5EFB8CBED9FC12" box="[1057,1162,931,952]" italics="true" pageId="11" pageNumber="438">Shokawa</emphasis>
</taxonomicName>
are two morphologically peculiar, long-necked choristoderes known from the Lower Cretaceous of
<collectingCountry id="F34976640F1A82059D3AFBCEBED9FC5D" box="[1093,1162,993,1015]" name="China" pageId="11" pageNumber="438">China</collectingCountry>
and
<collectingCountry id="F34976640F1A82059DA9FBCEBF4EFC5C" box="[1238,1309,993,1014]" name="Japan" pageId="11" pageNumber="438">Japan</collectingCountry>
(
<bibRefCitation id="EFCF4B050F1A82059C49FBCEB98EFBBE" author="Evans SE &amp; Manabe M" pageId="11" pageNumber="438" pagination="101 - 119" refId="ref10977" refString="Evans SE, Manabe M. 1999. A choristoderan reptile from the Lower Cretaceous of Japan. Special Papers in Palaeontology 60: 101 - 119." type="journal article" year="1999">Evans &amp; Manabe, 1999</bibRefCitation>
;
<bibRefCitation id="EFCF4B050F1A82059A95FBD0BF58FBBF" author="Gao KQ &amp; Tang Z &amp; Wang X" box="[1002,1291,1023,1045]" pageId="11" pageNumber="438" pagination="1 - 8" refId="ref11123" refString="Gao KQ, Tang Z, Wang X. 1999. A long-necked diapsid reptile from the Upper Jurassic / Lower Cretaceous of Liaoning Province, northeastern China. Vertebrata Palasiatica 37: 1 - 8." type="journal article" year="1999">Gao, Tang &amp; Wang, 1999</bibRefCitation>
). Both MPTs (
<figureCitation id="13652A710F1A82059A3DFC31B9D1FB9E" box="[834,898,1054,1076]" captionStart="Figure 6" captionStartId="11.[162,240,1825,1844]" captionTargetBox="[170,768,810,1793]" captionTargetId="graphics-586@11.[201,760,976,1793]" captionTargetPageId="11" captionText="Figure 6. Comparison of the two equally parsimonious trees (TL = 123 steps; CI = 0.85; RI = 0.87) derived from analysis of data presented in Appendices 1 and 2." figureDoi="http://doi.org/10.5281/zenodo.4687375" httpUri="https://zenodo.org/record/4687375/files/figure.png" pageId="11" pageNumber="438">Fig. 6</figureCitation>
) recognize a
<emphasis id="B92AEAE60F1A82059D51FC31BF01FB99" box="[1070,1362,1054,1075]" italics="true" pageId="11" pageNumber="438">
<taxonomicName id="4C5E4D770F1A82059D51FC31BE88FB99" box="[1070,1243,1054,1075]" class="Reptilia" family="Hyphalosauridae" genus="Hyphalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="genus">Hyphalosaurus</taxonomicName>
<taxonomicName id="4C5E4D770F1A82059D95FC31BF01FB99" box="[1258,1362,1054,1075]" class="Reptilia" genus="Shokawa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eosuchia" pageId="11" pageNumber="438" phylum="Chordata" rank="genus">Shokawa</taxonomicName>
</emphasis>
clade, for which the name
<taxonomicName id="4C5E4D770F1A82059D4AFC12BEAAFBF9" authorityName="Gao &amp; Fox" authorityYear="2005" box="[1077,1273,1085,1107]" class="Reptilia" family="Hyphalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">Hyphalosauridae</taxonomicName>
(fam. nov.) is proposed here. The grouping of the two forms in the family
<taxonomicName id="4C5E4D770F1A82059AF3FC55BE03FB3A" authorityName="Gao &amp; Fox" authorityYear="2005" box="[908,1104,1146,1168]" class="Reptilia" family="Hyphalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">Hyphalosauridae</taxonomicName>
is unambiguously supported by two uniquely derived character states of the two genera of the family: cervical vertebral count more than 16 (67-1); and caudal neural spines tall and narrow (72-1). Because the skull morphology is either poorly known or entirely unknown for the two forms, the diagnosis of the family would be enhanced by further study of the known material and discovery of new material.
</paragraph>
<paragraph id="8BE136F40F1A82029A2CFDA1B819FF52" blockId="11.[826,1443,931,1904]" lastBlockId="12.[142,758,195,248]" lastPageId="12" lastPageNumber="439" pageId="11" pageNumber="438">
Within the phylogenetic framework of the
<taxonomicName id="4C5E4D770F1A82059C32FDA1B9D6FA69" authorityName="COPE" authorityYear="1876" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="order">Choristodera</taxonomicName>
, both
<taxonomicName id="4C5E4D770F1A82059AABFD82BECAFA69" box="[980,1177,1453,1475]" class="Reptilia" family="Monjurosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">Monjurosuchidae</taxonomicName>
and
<taxonomicName id="4C5E4D770F1A82059DA2FD82BFF2FA69" authorityName="Gao &amp; Fox" authorityYear="2005" box="[1245,1441,1453,1475]" class="Reptilia" family="Hyphalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">Hyphalosauridae</taxonomicName>
are recognized as robust familial groups outside the neochoristoderan clade; however, the two MPTs differ from one another in placing the two families as either successive sister groups with the
<taxonomicName id="4C5E4D770F1A82059D99FE08BFF2F997" authorityName="Evans &amp; Hecht" authorityYear="1993" box="[1254,1441,1575,1597]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="439" phylum="Chordata" rank="subOrder" subOrder="Neochoristodera">Neochoristodera</taxonomicName>
(
<figureCitation id="13652A710F1A82059A3DFE69B9C5F9F6" box="[834,918,1606,1628]" captionStart="Figure 6" captionStartId="11.[162,240,1825,1844]" captionTargetBox="[170,768,810,1793]" captionTargetId="graphics-586@11.[201,760,976,1793]" captionTargetPageId="11" captionText="Figure 6. Comparison of the two equally parsimonious trees (TL = 123 steps; CI = 0.85; RI = 0.87) derived from analysis of data presented in Appendices 1 and 2." figureDoi="http://doi.org/10.5281/zenodo.4687375" httpUri="https://zenodo.org/record/4687375/files/figure.png" pageId="11" pageNumber="438">Fig. 6A</figureCitation>
), or grouped together as the sister group with the
<taxonomicName id="4C5E4D770F1A82059A19FE4ABE73F9D1" authorityName="Evans &amp; Hecht" authorityYear="1993" box="[870,1056,1637,1659]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="439" phylum="Chordata" rank="subOrder" subOrder="Neochoristodera">Neochoristodera</taxonomicName>
(
<figureCitation id="13652A710F1A82059D50FE4ABED0F9D1" box="[1071,1155,1637,1659]" captionStart="Figure 6" captionStartId="11.[162,240,1825,1844]" captionTargetBox="[170,768,810,1793]" captionTargetId="graphics-586@11.[201,760,976,1793]" captionTargetPageId="11" captionText="Figure 6. Comparison of the two equally parsimonious trees (TL = 123 steps; CI = 0.85; RI = 0.87) derived from analysis of data presented in Appendices 1 and 2." figureDoi="http://doi.org/10.5281/zenodo.4687375" httpUri="https://zenodo.org/record/4687375/files/figure.png" pageId="11" pageNumber="438">Fig. 6B</figureCitation>
). The grouping of
<taxonomicName id="4C5E4D770F1A82059C2FFE4AB9E5F933" class="Reptilia" family="Monjurosuchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">Monjurosuchidae</taxonomicName>
with
<taxonomicName id="4C5E4D770F1A82059A82FEACBE92F933" authorityName="Gao &amp; Fox" authorityYear="2005" box="[1021,1217,1667,1689]" class="Reptilia" family="Hyphalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="11" pageNumber="438" phylum="Chordata" rank="family">Hyphalosauridae</taxonomicName>
as sister groups is weakly supported by the following two character states: lateral borders of frontals strongly constricted (3-1; unknown for
<taxonomicName id="4C5E4D770F1A82059D70FEF0BE2BF95E" box="[1039,1144,1759,1780]" class="Reptilia" genus="Shokawa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eosuchia" pageId="11" pageNumber="438" phylum="Chordata" rank="genus">
<emphasis id="B92AEAE60F1A82059D70FEF0BE2BF95E" box="[1039,1144,1759,1780]" italics="true" pageId="11" pageNumber="438">Shokawa</emphasis>
</taxonomicName>
); and orbit large and dorsally orientated (11-1; unknown for
<taxonomicName id="4C5E4D770F1A82059C55FED1BFC0F8B9" box="[1322,1427,1790,1811]" class="Reptilia" genus="Shokawa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Eosuchia" pageId="11" pageNumber="438" phylum="Chordata" rank="genus">
<emphasis id="B92AEAE60F1A82059C55FED1BFC0F8B9" box="[1322,1427,1790,1811]" italics="true" pageId="11" pageNumber="438">Shokawa</emphasis>
</taxonomicName>
). Because the current data matrix contains a large amount of missing data for the basic taxa of the two families, the robustness of the possible Monjurosuchidae
<taxonomicName id="4C5E4D770F1D8202987EF8ECBB97FF73" authorityName="Gao &amp; Fox" authorityYear="2005" box="[257,452,195,217]" class="Reptilia" family="Hyphalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Choristodera" pageId="12" pageNumber="439" phylum="Chordata" rank="family">Hyphalosauridae</taxonomicName>
clade must be tested further when new data become available.
</paragraph>
</subSubSection>
</treatment>
</document>