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<document id="90C07E9CCCD36495C4AE0B99B9C41BE1" ID-DOI="10.1093/zoolinnean/zlad165" ID-ISSN="0024-4082" ID-Zenodo-Dep="11241243" ID-ZooBank="1EEF0D52-180B-4D3D-AB95-91AF3091E272" IM.bibliography_approvedBy="juliana" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="juliana" IM.metadata_approvedBy="juliana" IM.tables_approvedBy="juliana" IM.taxonomicNames_approvedBy="juliana" IM.treatments_approvedBy="juliana" checkinTime="1716364298778" checkinUser="plazi" docAuthor="Young, Mark T., Wilberg, Eric W., Johnson, Michela M., Herrera, Yanina, Brandalise, Marco de Andrade, Brignon, Arnaud, Sachs, Sven, Abel, Pascal, Foffa, Davide, Fernández, Marta S., Vignaud, Patrick, Cowgill, Thomas &amp; Brusatte, Stephen L." docDate="2024" docId="03D08506FF83730A79FD0CD70DCFFB0F" docLanguage="en" docName="zlad165.pdf" docOrigin="Zoological Journal of the Linnean Society 200 (2)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlad165" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.5:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="5" docTitle="Metriorhynchidae Fitzinger 1843" docType="treatment" docVersion="1" lastPageNumber="600" masterDocId="FFE9FD7EFFB0733F796008100D3AFFFF" masterDocTitle="The history, systematics, and nomenclature of Thalattosuchia (Archosauria: Crocodylomorpha)" masterLastPageNumber="617" masterPageNumber="547" pageNumber="598" updateTime="1716820941663" updateUser="juliana">
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<mods:title id="477D2408122D204778C317FF99E26CAF">The history, systematics, and nomenclature of Thalattosuchia (Archosauria: Crocodylomorpha)</mods:title>
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<mods:affiliation id="5090259AD73D99A9A09165634844DC84">Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), División Paleontología Vertebrados, Museo de La Plata, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, B 1900 La Plata, Buenos Aires, Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) &amp; Departamento de Diversidade e Ecologia, Faculdade de Biociências, Pontifícia Universidade Católica do Rio Grande do Sul, Av. Ipiranga 6681, Prédio 12, Porto Alegre, 90619 - 900, Brazil &amp; Setor de Paleontologia, Museu de Ciências e Tecnologia, Pontifícia Universidade Católica do Rio Grande do Sul, Av. Ipiranga 6681, Prédio 40, Porto Alegre, 90619 - 900, Brazil</mods:affiliation>
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<mods:affiliation id="28D7D55B975C33EC4385AD03755ACB5E">Department of Geosciences, Virginia Tech, 4044 Derring Hall 926 West Campus Drive, Blacksburg 24061, Virginia, USA &amp; School of Geography, Earth and Environmental Sciences, University of Birmingham, Edgbaston, Birmingham B 15 2 TT, United Kingdom</mods:affiliation>
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<mods:namePart id="CE09BC408A00EFFEAF7F9215C41AF586">Fernández, Marta S.</mods:namePart>
<mods:affiliation id="84F5D930453EAC78BF36FBE797A0CACD">Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), División Paleontología Vertebrados, Museo de La Plata, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, B 1900 La Plata, Buenos Aires, Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)</mods:affiliation>
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Metriorhynchidae
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(Zoological Code)
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<emphasis id="B90DE802FF83730C79150D150F64FAC1" bold="true" pageId="51" pageNumber="598">
<taxonomicName id="4C794F93FF83730C79150D150CCAFAE0" authority="Fitzinger 1843: 35" authorityName="Fitzinger" authorityPageNumber="35" authorityYear="1843" box="[117,496,1285,1312]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="family">
<emphasis id="B90DE802FF83730C79150D150C12FADF" bold="true" box="[117,296,1285,1312]" italics="true" pageId="51" pageNumber="598">Metriorhynchidae</emphasis>
<bibRefCitation id="EFE849E1FF83730C784D0D150CCAFAE0" author="Fitzinger L" box="[301,496,1285,1312]" pageId="51" pageNumber="598" refId="ref51847" refString="Fitzinger L. Systema Reptilium: Amblyglossae. Fasciculus primus, Part 1. Vienna: Braumuller et Seidel, 1843, 106." type="book" year="1843">Fitzinger 1843: 35</bibRefCitation>
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(as Metriorhynchi), converted clade name (PhyloCode)
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<paragraph id="8BC63410FF83730C79110D5C0D96FA7C" blockId="51.[113,758,1285,1411]" pageId="51" pageNumber="598">RegNum registration number 1017.</paragraph>
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genus:
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<emphasis id="B90DE802FF83730C79880DB40CBBFA43" box="[232,385,1444,1468]" italics="true" pageId="51" pageNumber="598">Metriorhynchus</emphasis>
<bibRefCitation id="EFE849E1FF83730C78E70DB40F08FA43" author="von Meyer H" box="[391,562,1444,1468]" pageId="51" pageNumber="598" refId="ref56694" refString="von Meyer H. Palaeologica zur Geschichte der Erde und ihrer Geschopfe. Frankfurt am Main: Siegmund Schmerber, 1832, 560." type="book" year="1832">von Meyer, 1832</bibRefCitation>
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.
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<emphasis id="B90DE802FF83730C78E30DCB0CD3FA0A" box="[387,489,1499,1525]" italics="true" pageId="51" pageNumber="598">Etymology</emphasis>
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‘The family of
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<emphasis id="B90DE802FF83730C78680E120C9BF9E5" box="[264,417,1538,1562]" italics="true" pageId="51" pageNumber="598">Metriorhynchus</emphasis>
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’. The stem
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- is based on the genus
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<emphasis id="B90DE802FF83730C799C0E310CAFF9C6" box="[252,405,1569,1593]" italics="true" pageId="51" pageNumber="598">Metriorhynchus</emphasis>
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. -
<emphasis id="B90DE802FF83730C78C80E310CEBF9C6" box="[424,465,1569,1593]" italics="true" pageId="51" pageNumber="598">idae</emphasis>
, is a Latin patronymic suffix, introduced into zoological nomenclature, as -idae, by
<bibRefCitation id="EFE849E1FF83730C7BA00E510DD9F987" author="Kirby W" pageId="51" pageNumber="598" pagination="86 - 123" refId="ref55225" refString="Kirby W. Strepsiptera, a new order of insects proposed; and the characters of the order, with those of its genera, laid down. The Transactions of the Linnean Society of London 1815; 11: 86 - 123." type="journal article" year="1815">Kirby (1815: 88)</bibRefCitation>
. Under Article 29.2 of the Zoological Code, the suffix -
<emphasis id="B90DE802FF83730C79190E6F0D98F968" box="[121,162,1663,1687]" italics="true" pageId="51" pageNumber="598">idae</emphasis>
denotes a family rank within the family-group.
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<subSubSection id="C363679BFF83730C78090EAA0E52FEBB" pageId="51" pageNumber="598" type="description">
<paragraph id="8BC63410FF83730C78090EAA0F39F92B" blockId="51.[361,515,1722,1748]" box="[361,515,1722,1748]" pageId="51" pageNumber="598">
<emphasis id="B90DE802FF83730C78090EAA0F39F92B" box="[361,515,1722,1748]" italics="true" pageId="51" pageNumber="598">Geological range</emphasis>
</paragraph>
<paragraph id="8BC63410FF83730C79110EF10D8CF8C8" blockId="51.[113,763,1761,1847]" pageId="51" pageNumber="598">
Middle Jurassic (early Bathonian) to Early Cretaceous (earliest Aptian) (
<bibRefCitation id="EFE849E1FF83730C79B10F100CA6F8E7" author="Cau A &amp; Fanti F" box="[209,412,1792,1816]" pageId="51" pageNumber="598" pagination="550 - 65" refId="ref49900" refString="Cau A, Fanti F. The oldest known metriorhynchid crocodylian from the Middle Jurassic of North-eastern Italy: Neptunidraco ammoniticus gen et sp nov. Gondwana Research 2011; 19: 550 - 65. https: // doi. org / 10.1016 / j. gr. 2010.07.007" type="journal article" year="2011">Cau and Fanti 2011</bibRefCitation>
,
<bibRefCitation id="EFE849E1FF83730C78C80F100F45F8E7" author="Chiarenza A &amp; Foffa D &amp; Young MT" box="[424,639,1792,1816]" pageId="51" pageNumber="598" pagination="608 - 16" refId="ref50078" refString="Chiarenza A, Foffa D, Young MT et al. The youngest record of metriorhynchid crocodylomorphs, with implications for the extinction of Thalattosuchia. Cretaceous Research 2015; 56: 608 - 16." type="journal article" year="2015">
Chiarenza
<emphasis id="B90DE802FF83730C7B760F110F05F8E7" box="[534,575,1792,1816]" italics="true" pageId="51" pageNumber="598">et al</emphasis>
. 2015
</bibRefCitation>
,
<bibRefCitation id="EFE849E1FF83730C7BEB0F100D9FF8C8" author="Sachs S &amp; Young MT &amp; Hornung J" pageId="51" pageNumber="598" pagination="104479" refId="ref58635" refString="Sachs S, Young MT, Hornung J. The enigma of Enaliosuchus, and a reassessment of the Early Cretaceous fossil record of Metriorhynchidae. Cretaceous Research 2020; 114: 104479." type="journal article" year="2020">
Sachs
<emphasis id="B90DE802FF83730C7BAB0F110FCEF8E7" box="[715,756,1792,1816]" italics="true" pageId="51" pageNumber="598">et al</emphasis>
. 2020
</bibRefCitation>
).
</paragraph>
<paragraph id="8BC63410FF83730C78790F720F69F883" blockId="51.[281,595,1890,1916]" box="[281,595,1890,1916]" pageId="51" pageNumber="598">
<emphasis id="B90DE802FF83730C78790F720F69F883" box="[281,595,1890,1916]" italics="true" pageId="51" pageNumber="598">PhyloCode phylogenetic definition</emphasis>
</paragraph>
<paragraph id="8BC63410FF83730C79110F99081AFF18" blockId="51.[113,763,1929,1984]" lastBlockId="51.[810,1459,144,231]" pageId="51" pageNumber="598">
The smallest clade within
<taxonomicName id="4C794F93FF83730C78DE0F990F4CF85E" authorityName="Fitzinger" authorityYear="1843" box="[446,630,1929,1953]" pageId="51" pageNumber="582" rank="superFamily" superFamily="Metriorhynchoidea">
<emphasis id="B90DE802FF83730C78DE0F990F4CF85E" box="[446,630,1929,1953]" italics="true" pageId="51" pageNumber="598">Metriorhynchoidea</emphasis>
</taxonomicName>
containing
<taxonomicName id="4C794F93FF83730C79110FB80CBAF83F" box="[113,384,1960,1984]" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="51" pageNumber="598" phylum="Chordata" rank="species" species="superciliosus">
<emphasis id="B90DE802FF83730C79110FB80CBAF83F" box="[113,384,1960,1984]" italics="true" pageId="51" pageNumber="598">Thalattosuchus superciliosus</emphasis>
</taxonomicName>
(Blainville in
<bibRefCitation id="EFE849E1FF83730C7B710FB80E64FF57" author="Eudes-Deslongchamps JA" box="[529,862,144,1984]" pageId="51" pageNumber="598" refId="ref51191" refString="Eudes-Deslongchamps JA. de Blainville HMD. Lettres sur les Crocodiles Vivants et Fossiles. Caen: A. Hardel, 1852, 38." type="book" year="1852">Eudes-Deslongchamps 1852</bibRefCitation>
),
<taxonomicName id="4C794F93FF83730C7A1D0880083CFF57" authority="(Andrews 1913)" baseAuthorityName="Andrews" baseAuthorityYear="1913" box="[893,1286,144,168]" class="Reptilia" family="Metriorhynchidae" genus="Gracilineustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="species" species="leedsi">
<emphasis id="B90DE802FF83730C7A1D0880097FFF57" box="[893,1093,144,168]" italics="true" pageId="51" pageNumber="598">Gracilineustes leedsi</emphasis>
(
<bibRefCitation id="EFE849E1FF83730C7D3E088009C1FF57" author="Andrews CW" box="[1118,1275,144,168]" pageId="51" pageNumber="598" refId="ref48442" refString="Andrews CW. A Descriptive Catalogue of the Marine Reptiles of the Oxford Clay, Part Two. London: British Museum (Natural History), 1913." type="book" year="1913">Andrews 1913</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="4C794F93FF83730C7C7A08800922FF38" authority="(Holl 1829)" authorityName="Metriorhynchoidea" baseAuthorityName="Holl" baseAuthorityYear="1829" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="species" species="brevirostris">
<emphasis id="B90DE802FF83730C7C7A08800EAFFF38" italics="true" pageId="51" pageNumber="598">Metriorhynchus brevirostris</emphasis>
(
<bibRefCitation id="EFE849E1FF83730C7AC608BF0937FF38" author="Holl F" box="[934,1037,175,199]" pageId="51" pageNumber="598" refId="ref53683" refString="Holl F. Handbuch der Petrefactenkunde Teil 1. Dresden: Hilscher, 1829, 416." type="book" year="1829">Holl 1829</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="4C794F93FF83730C7D4408BF0897FF38" authority="von Meyer 1831" authorityName="von Meyer" authorityYear="1831" box="[1060,1453,175,200]" class="Reptilia" family="Metriorhynchidae" genus="Rhacheosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B90DE802FF83730C7D4408BF083AFF38" box="[1060,1280,175,199]" italics="true" pageId="51" pageNumber="598">Rhacheosaurus gracilis</emphasis>
<bibRefCitation id="EFE849E1FF83730C7C6508A00897FF38" author="von Meyer H" box="[1285,1453,175,200]" pageId="51" pageNumber="598" pagination="171 - 200" refId="ref56665" refString="von Meyer H. Neue fossile Reptilien aus der Ordnung der Saurier. Nova Acta physico-medica Academiae Caesareae Leopoldino-Carolinae Naturae Curiosum 1831; 15: 171 - 200." type="journal article" year="1831">von Meyer 1831</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C794F93FF83730C7A3608DF0826FF18" authority="(von Sommerring 1816)" baseAuthorityName="von Sommerring" baseAuthorityYear="1816" box="[854,1308,207,231]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="species" species="giganteus">
<emphasis id="B90DE802FF83730C7A3608DF0921FF19" box="[854,1051,207,230]" italics="true" pageId="51" pageNumber="598">Geosaurus giganteus</emphasis>
(
<bibRefCitation id="EFE849E1FF83730C7D4A08DF0835FF18" author="von Sommerring ST" box="[1066,1295,207,231]" pageId="51" pageNumber="598" pagination="37 - 58" refId="ref59308" refString="von Sommerring ST. Ueber die Lacerta gigantea der Vorwelt. Denkschriften der Koniglichen Akademie der Wissenschaften zu Munchen Classe der Mathematik und Naturwissenschaften 1816; 6: 37 - 58." type="journal article" year="1816">von Sömmerring 1816</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="8BC63410FF83730C7D72091609F6FEDF" blockId="51.[1042,1228,262,288]" box="[1042,1228,262,288]" pageId="51" pageNumber="598">
<emphasis id="B90DE802FF83730C7D72091609F6FEDF" box="[1042,1228,262,288]" italics="true" pageId="51" pageNumber="598">Reference phylogeny</emphasis>
</paragraph>
<paragraph id="8BC63410FF83730C7A4A093D0E52FEBB" blockId="51.[810,872,301,325]" box="[810,872,301,325]" pageId="51" pageNumber="598">
<figureCitation id="13422895FF83730C7A4A093D0E58FEBA" box="[810,866,301,325]" captionStart="Figure 3" captionStartId="19.[113,178,1185,1209]" captionTargetBox="[117,1455,147,1154]" captionTargetId="figure-283@19.[114,1458,144,1157]" captionTargetPageId="19" captionText="Figure 3. The equal weights, no constraints strict consensus topology (C0). Length = 4164, CI = 0.303, RI = 0.831, RC = 0.251, HI = 0.697, number of most parsimonious cladograms hit the hold limit.Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig. S1 for the entire topology). A, Crocodylomorpha; B, Crocodyliformes; C, Thalattosuchia; D, Neothalattosuchia; E, Teleosauridae; F, Teleosaurinae; G, Aeolodontini; H, Machimosauridae; I, Machimosaurinae; J, Machimosaurini; K, Euthalattosuchia; L, Metriorhynchidae; M, Metriorhynchinae; N, Rhacheosaurini; O, Geosaurinae; P, Geosaurini. Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians.Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T. Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons.org/licenses/by/3.0/." figureDoi="http://doi.org/10.5281/zenodo.11241251" httpUri="https://zenodo.org/record/11241251/files/figure.png" pageId="51" pageNumber="598">Fig. 3</figureCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C363679BFF83730B7A5B09740DECFA14" lastPageId="52" lastPageNumber="599" pageId="51" pageNumber="598" type="diagnosis">
<paragraph id="8BC63410FF83730C7A5B09740898FE81" blockId="51.[827,1442,356,382]" box="[827,1442,356,382]" pageId="51" pageNumber="598">
<emphasis id="B90DE802FF83730C7A5B09740898FE81" box="[827,1442,356,382]" italics="true" pageId="51" pageNumber="598">Zoological Code diagnosis and PhyloCode diagnostic apomorphies</emphasis>
</paragraph>
<paragraph id="8BC63410FF83730C7A4A099B094DFC87" blockId="51.[810,1459,395,888]" pageId="51" pageNumber="598">
Metriorhynchoid crocodylomorphs with the following unique combination of characters (10): absence of pharyngotympanic tube foramina (some Middle Jurassic metriorhynchid specimens appear to have these foramina, it is currently unclear if the loss of these foramina represent an ontogenetic signal or possibly a phylogenetic signal within
<taxonomicName id="4C794F93FF83730C7DFA0A37086DFDC0" authorityName="Fitzinger" authorityYear="1843" box="[1178,1367,551,575]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="family">Metriorhynchidae</taxonomicName>
) (65.0); in the intertemporal bar, the frontal is wider than the parietal (reversal in
<taxonomicName id="4C794F93FF83730C7AC60A760905FD81" authorityName="von Meyer" authorityYear="1832" box="[934,1087,614,638]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF83730C7AC60A760905FD81" box="[934,1087,614,638]" italics="true" pageId="51" pageNumber="598">Metriorhynchus</emphasis>
</taxonomicName>
) (235.1); presence of a true supraorbital notch (285.1); dermic postorbital bar (309.2); dorsal part of the postorbital bar constricted, distinct from the dorsal part of the postorbital (318.1); humerus deltopectoral crest continuous with the proximal surface (753.1); humerus is flattened and has a plate-like appearance (757.1); humerus diaphysis contributes less than 40% of total humerus length (758.1–2); humerus epiphyses are unossified (760.1); radius/ulna are broadly expanded and plate-like (764.1).
</paragraph>
<paragraph id="8BC63410FF83730C7ABB0B880839FC4D" blockId="51.[810,1460,920,1985]" box="[987,1283,920,946]" pageId="51" pageNumber="598">
<emphasis id="B90DE802FF83730C7ABB0B880839FC4D" box="[987,1283,920,946]" italics="true" pageId="51" pageNumber="598">Potentially diagnostic characters</emphasis>
</paragraph>
<paragraph id="8BC63410FF83730C7A4A0BAF0812FBCA" blockId="51.[810,1460,920,1985]" pageId="51" pageNumber="598">
The following 51 characters are also diagnostic for
<taxonomicName id="4C794F93FF83730C7A4A0BCE0EDEFC09" authorityName="Fitzinger" authorityYear="1843" box="[810,996,990,1014]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="family">Metriorhynchidae</taxonomicName>
but may be more widely distributed within
<taxonomicName id="4C794F93FF83730C7A4A0BED0EC8FBEA" authorityName="Fitzinger" authorityYear="1843" box="[810,1010,1021,1045]" pageId="51" pageNumber="582" rank="superFamily" superFamily="Metriorhynchoidea">Metriorhynchoidea</taxonomicName>
once more complete specimens of non-metriorhynchid metriorhynchoids are discovered.
</paragraph>
<paragraph id="8BC63410FF83730C7A250C2C09BDFB8C" blockId="51.[810,1460,920,1985]" pageId="51" pageNumber="598">Characters unknown in all early diverging metriorhynchoids: lack of scales on the body (874.1).</paragraph>
<paragraph id="8BC63410FF83730C7A250C6B097EF83E" blockId="51.[810,1460,920,1985]" pageId="51" pageNumber="598">
Characters unknown in early diverging metriorhynchoids more derived than
<taxonomicName id="4C794F93FF83730C7A9B0C8A0943FB4D" authorityName="Haeckel" authorityYear="1895" box="[1019,1145,1178,1202]" class="Reptilia" family="Metriorhynchidae" genus="Pelagosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF83730C7A9B0C8A0943FB4D" box="[1019,1145,1178,1202]" italics="true" pageId="51" pageNumber="598">Pelagosaurus</emphasis>
</taxonomicName>
: ‘pelagic morphotype’ of the endosseous labyrinth (38.2); cochlear duct shortened (40.1); basisphenoid diverticula cavities indistinct from parent sinus cavity (62.2); no diverticular inflation within the basioccipital (67.0); absence of ‘otoccipital recesses’ (79.0); seven cervical vertebrae (687.2); mid-to-late thoracic vertebrae lack a shallow fossa on the anterior margin of the diapophysis (703.1); shift in neural spine inclination near distal tail (718.1); ventral deflection of distal caudal vertebrae (719.1–3); tail deflection is abrupt, occurring over 5–10 vertebrae (720.2); distal chevrons mediolaterally compressed, with some having a ‘W’-shape created by an anterior process (732.1); forelimbs (humerus + ulna + metacarpal III) are less than 20% of total trunk (presacral vertebrae minus the atlas-axis) length (shared with
<taxonomicName id="4C794F93FF83730C7A070E400EF8F997" authorityName="von Meyer" authorityYear="1832" box="[871,962,1616,1640]" class="Reptilia" family="Teleosauridae" genus="Aeolodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF83730C7A070E400EF8F997" box="[871,962,1616,1640]" italics="true" pageId="51" pageNumber="598">Aeolodon</emphasis>
</taxonomicName>
) (750.4); metacarpal III makes up 20% or more of combined forelimb length (humerus + ulna + metacarpal III) (751.2–3); radiale/ulnare are broadly expanded and plate-like (769.1); radiale and ulnare subequal in length (772.0); metacarpal I is broadly expanded (776.1); hindlimbs (femur + tibia + metatarsal III) are less than 40% of total trunk (presacral vertebrae minus the atlas-axis) length (807.5); forelimbs (humerus + radius) less than 45% of hindlimb (femur + tibia) length (shared with
<taxonomicName id="4C794F93FF83730C7D8A0F5B087FF89C" authorityName="von Meyer" authorityYear="1832" box="[1258,1349,1867,1891]" class="Reptilia" family="Teleosauridae" genus="Aeolodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="51" pageNumber="598" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF83730C7D8A0F5B087FF89C" box="[1258,1349,1867,1891]" italics="true" pageId="51" pageNumber="598">Aeolodon</emphasis>
</taxonomicName>
) (808.4); calcaneum tuber poorly developed (829.1); metatarsals II–IV shorter than their respective digits (831.2); absence of dorsal caudal osteoderms (862.1).
</paragraph>
<paragraph id="8BC63410FF84730B79FC08800C1DFDFF" blockId="52.[129,778,144,1515]" pageId="52" pageNumber="599">
Characters unknown in early diverging metriorhynchoids more derived than
<taxonomicName id="4C794F93FF84730B782508BF0CF3FF38" authorityName="Haeckel" authorityYear="1895" box="[325,457,175,199]" class="Reptilia" family="Metriorhynchidae" genus="Teleidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B782508BF0CF3FF38" box="[325,457,175,199]" italics="true" pageId="52" pageNumber="599">Teleidosaurus</emphasis>
</taxonomicName>
: highly elliptical external nares (shared with
<taxonomicName id="4C794F93FF84730B786308DF0C46FF18" authorityName="Holland" authorityYear="1909" box="[259,380,207,231]" class="Crocodylia" family="Crocodylidae" genus="Deinosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B786308DF0C46FF18" box="[259,380,207,231]" italics="true" pageId="52" pageNumber="599">Deinosuchus</emphasis>
</taxonomicName>
) (96.3); posterodorsal retraction of the anterior margin of the narial fossa, such that it is at least level to the second premaxillary alveoli (103.2–5); external nares at least partially divided by a midline premaxillary septum (105.1–2); premaxilla contributes more than 25% of total rostrum length (120.1); premaxilla is lanceolate in shape, smoothly fitting the rostrum (133.1); of the anterior processes of the surangular, the ventral process is longer than the dorsal process (549.2); premaxilla contacts the anterior margin of the first maxillary alveoli (589.1–2).
</paragraph>
<paragraph id="8BC63410FF84730B79FC0A180CD5FB8C" blockId="52.[129,778,144,1515]" pageId="52" pageNumber="599">
Characters unknown in early diverging metriorhynchoids more derived than
<taxonomicName id="4C794F93FF84730B782C0A370CD9FDC0" authorityName="Ösi, Young, Galácz &amp; Rabi" authorityYear="2018" box="[332,483,551,575]" genus="Magyarosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B782C0A370CD9FDC0" box="[332,483,551,575]" italics="true" pageId="52" pageNumber="599">Magyarosuchus</emphasis>
</taxonomicName>
and the unnamed Toarcian taxon: coronoid process considerably dorsal to the tooth-row (487.1); absence of external mandibular fenestrae (512.0); thoracic vertebrae parapophyses are oriented anteriorly (704.1); anterior thoracic vertebrae in lateral view, have the parapophysis dorsal to the diapophysis (705.1); scapula blade reduced, as wide as, or narrower than, the glenoid region (744.2); ilium anterior process is short and robust (787.1); absence of ilium posterior process (794.1); dorsal border of the ilium 21% or less of femoral length (797.1); ilium lacks acetabular depressions with the peduncles being contiguous with the ventral margin (800.1); ischium anterior processes reduced, lacking clear articulation facets (802.1–2); tibia less than 50% of femoral length (shared with machimosaurine machimosaurids and aeolodontin teleosaurids) (809.3–5); middle of the femoral diaphysis flattened (817: 1); femur distal condyles incompletely ossified (822.2); distal surface between the lateral and medial condyles is flat (823.0); absence of gastral osteoderms (836.0); absence of dorsal thoracic osteoderms (838.0).
</paragraph>
<paragraph id="8BC63410FF84730B79FC0C6A0DECFA14" blockId="52.[129,778,144,1515]" pageId="52" pageNumber="599">
Characters unknown in early diverging metriorhynchoids more derived than
<taxonomicName id="4C794F93FF84730B781B0C8A0CE3FB4E" authorityName="Young, Brusatte, Ruta &amp; Andrade" authorityYear="2010" box="[379,473,1178,1201]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B781B0C8A0CE3FB4E" box="[379,473,1178,1201]" italics="true" pageId="52" pageNumber="599">Eoneustes</emphasis>
</taxonomicName>
: two major longitudinal neurovascular canals in the rostrum (47.2); dorsal alveolar canal innervates the dental alveoli with a canal leaving the alveolus continuing to the external surface of the rostrum (49.1); nasal lateroposterior processes are well-developed (139.2); nasals reach deep into the preorbital fossa (175.2); jugal reaches deep into the preorbital fossa (reversals in
<taxonomicName id="4C794F93FF84730B7BE70D460E3EFA92" box="[647,772,1366,1389]" class="Reptilia" family="Metriorhynchidae" genus="Torvoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7BE70D460E3EFA92" box="[647,772,1366,1389]" italics="true" pageId="52" pageNumber="599">Torvoneustes</emphasis>
</taxonomicName>
,
<taxonomicName id="4C794F93FF84730B79E10D650C1BFA72" authorityName="Parrilla-Bel, Young, Moreno-Azanza &amp; Canudo" authorityYear="2013" box="[129,289,1397,1421]" family="Metriorhynchidae" genus="Maledictosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B79E10D650C1BFA72" box="[129,289,1397,1421]" italics="true" pageId="52" pageNumber="599">Maledictosuchus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C794F93FF84730B783D0D650F00FA72" baseAuthorityName="Kühn" baseAuthorityYear="1936" box="[349,570,1397,1421]" class="Reptilia" family="Metriorhynchidae" genus="Cricosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="species" species="schroederi">
<emphasis id="B90DE802FF84730B783D0D650F00FA72" box="[349,570,1397,1421]" italics="true" pageId="52" pageNumber="599">Cricosaurus schroederi</emphasis>
</taxonomicName>
) (176.1); posterior maxillary alveoli are ventrally offset relative to other alveoli (due to the ventroposterior curvature of the posterior maxilla) (576.1).
</paragraph>
</subSubSection>
<subSubSection id="C363679BFF84730A78E80E1B0DCFFB0F" lastPageId="53" lastPageNumber="600" pageId="52" pageNumber="599" type="discussion">
<paragraph id="8BC63410FF84730B78E80E1B0F3BF9DA" blockId="52.[129,777,1547,1641]" box="[392,513,1547,1573]" pageId="52" pageNumber="599">
<emphasis id="B90DE802FF84730B78E80E1B0F3BF9DA" box="[392,513,1547,1573]" italics="true" pageId="52" pageNumber="599">Composition</emphasis>
</paragraph>
<paragraph id="8BC63410FF84730B79E10E220CE7F997" blockId="52.[129,777,1547,1641]" pageId="52" pageNumber="599">
<taxonomicName id="4C794F93FF84730B79E10E220C39F9B6" authorityName="Arldt" authorityYear="1907" box="[129,259,1586,1609]" class="Reptilia" family="Metriorhynchidae" genus="Neustosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B79E10E220C39F9B6" box="[129,259,1586,1609]" italics="true" pageId="52" pageNumber="599">Neustosaurus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C794F93FF84730B78750E210CA9F9B6" authorityName="Koken" authorityYear="1883" box="[277,403,1585,1609]" class="Reptilia" family="Metriorhynchidae" genus="Enaliosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B78750E210CA9F9B6" box="[277,403,1585,1609]" italics="true" pageId="52" pageNumber="599">Enaliosuchus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C794F93FF84730B78C40E210F0FF9B6" authorityName="Young, Brignon, Sachs, Hornung, Foffa, Kitson, Johnson &amp; Steel" authorityYear="2020" box="[420,565,1585,1609]" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B78C40E210F0FF9B6" box="[420,565,1585,1609]" italics="true" pageId="52" pageNumber="599">Thalattosuchus</emphasis>
</taxonomicName>
, and the subclades
<taxonomicName id="4C794F93FF84730B79E10E410C17F996" authorityName="Lydekker" authorityYear="1888" box="[129,301,1617,1641]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="subFamily" subFamily="Metriorhynchinae">
<emphasis id="B90DE802FF84730B79E10E410C17F996" box="[129,301,1617,1641]" italics="true" pageId="52" pageNumber="599">Metriorhynchinae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C794F93FF84730B783E0E410CEDF997" authorityName="Bonaparte" authorityYear="1845" box="[350,471,1617,1640]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="subFamily" subFamily="Geosaurinae">
<emphasis id="B90DE802FF84730B783E0E410CEDF997" box="[350,471,1617,1640]" italics="true" pageId="52" pageNumber="599">Geosaurinae</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BC63410FF84730B78F20E990CC2F95C" blockId="52.[129,777,1673,1766]" box="[402,504,1673,1699]" pageId="52" pageNumber="599">
<emphasis id="B90DE802FF84730B78F20E990CC2F95C" box="[402,504,1673,1699]" italics="true" pageId="52" pageNumber="599">Comments</emphasis>
</paragraph>
<paragraph id="8BC63410FF84730B79E10EBF0F29F919" blockId="52.[129,777,1673,1766]" pageId="52" pageNumber="599">
<emphasis id="B90DE802FF84730B79E10EBF0DCEF938" box="[129,244,1711,1735]" italics="true" pageId="52" pageNumber="599">Authorship:</emphasis>
The nomen
<taxonomicName id="4C794F93FF84730B78EF0EBF0F06F938" authorityName="Fitzinger" authorityYear="1843" box="[399,572,1711,1735]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="family">
<emphasis id="B90DE802FF84730B78EF0EBF0F06F938" box="[399,572,1711,1735]" italics="true" pageId="52" pageNumber="599">Metriorhynchidae</emphasis>
</taxonomicName>
was first used by
<bibRefCitation id="EFE849E1FF84730B79E10EDE0C68F919" author="Fitzinger L" box="[129,338,1742,1766]" pageId="52" pageNumber="599" refId="ref51847" refString="Fitzinger L. Systema Reptilium: Amblyglossae. Fasciculus primus, Part 1. Vienna: Braumuller et Seidel, 1843, 106." type="book" year="1843">Fitzinger (1843: 35)</bibRefCitation>
, as Metriorhynchi.
</paragraph>
<paragraph id="8BC63410FF84730B79E10F18088BFE7C" blockId="52.[129,778,1799,1980]" lastBlockId="52.[825,1475,144,387]" pageId="52" pageNumber="599">
<emphasis id="B90DE802FF84730B79E10F180C9EF8E0" box="[129,420,1799,1823]" italics="true" pageId="52" pageNumber="599">Prior phylogenetic definition:</emphasis>
<bibRefCitation id="EFE849E1FF84730B78D00F180FE7F8E0" author="Young MT &amp; Andrade MB" box="[432,733,1799,1824]" pageId="52" pageNumber="599" pagination="551 - 85" refId="ref60527" refString="Young MT, Andrade MB. What is Geosaurus? Redescription of Geosaurus giganteus (Thalattosuchia, Metriorhynchidae) from the Upper Jurassic of Bayern, Germany. Zoological Journal of the Linnean Society 2009; 157: 551 - 85." type="journal article" year="2009">Young and Andrade (2009)</bibRefCitation>
defined
<taxonomicName id="4C794F93FF84730B79DC0F370C53F8C0" authorityName="Fitzinger" authorityYear="1843" box="[188,361,1831,1855]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="family">
<emphasis id="B90DE802FF84730B79DC0F370C53F8C0" box="[188,361,1831,1855]" italics="true" pageId="52" pageNumber="599">Metriorhynchidae</emphasis>
</taxonomicName>
as the least inclusive clade consisting of
<emphasis id="B90DE802FF84730B79E10F560C4FF8A1" box="[129,373,1862,1886]" italics="true" pageId="52" pageNumber="599">Metriorhynchusgeoffroyii</emphasis>
vonMeyer 1832 and
<emphasis id="B90DE802FF84730B7B270F560E30F8A2" box="[583,778,1862,1885]" italics="true" pageId="52" pageNumber="599">Geosaurusgiganteus</emphasis>
(
<bibRefCitation id="EFE849E1FF84730B79EB0F760C4CF882" author="von Sommerring ST" box="[139,374,1893,1917]" pageId="52" pageNumber="599" pagination="37 - 58" refId="ref59308" refString="von Sommerring ST. Ueber die Lacerta gigantea der Vorwelt. Denkschriften der Koniglichen Akademie der Wissenschaften zu Munchen Classe der Mathematik und Naturwissenschaften 1816; 6: 37 - 58." type="journal article" year="1816">von Sömmerring 1816</bibRefCitation>
). Here we have expanded the number of internal specifiers, so that
<taxonomicName id="4C794F93FF84730B78B70F950F52F862" authorityName="Young, Brignon, Sachs, Hornung, Foffa, Kitson, Johnson &amp; Steel" authorityYear="2020" box="[471,616,1925,1949]" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B78B70F950F52F862" box="[471,616,1925,1949]" italics="true" pageId="52" pageNumber="599">Thalattosuchus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C794F93FF84730B7B1D0F950E3EF862" box="[637,772,1925,1949]" class="Reptilia" family="Metriorhynchidae" genus="Gracilineustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7B1D0F950E3EF862" box="[637,772,1925,1949]" italics="true" pageId="52" pageNumber="599">Gracilineustes</emphasis>
</taxonomicName>
,
<taxonomicName id="4C794F93FF84730B79E10FB40C20F843" authorityName="von Meyer" authorityYear="1832" box="[129,282,1956,1980]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B79E10FB40C20F843" box="[129,282,1956,1980]" italics="true" pageId="52" pageNumber="599">Metriorhynchus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C794F93FF84730B78210FB40CE9F843" box="[321,467,1956,1980]" class="Reptilia" family="Metriorhynchidae" genus="Rhacheosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B78210FB40CE9F843" box="[321,467,1956,1980]" italics="true" pageId="52" pageNumber="599">Rhacheosaurus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C794F93FF84730B7B220FB40F92F844" authorityName="Bonaparte" authorityYear="1850" box="[578,680,1956,1979]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7B220FB40F92F844" box="[578,680,1956,1979]" italics="true" pageId="52" pageNumber="599">Geosaurus</emphasis>
</taxonomicName>
define
<taxonomicName id="4C794F93FF84730B7A5908800EDCFF57" authorityName="Fitzinger" authorityYear="1843" box="[825,998,144,168]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="family">
<emphasis id="B90DE802FF84730B7A5908800EDCFF57" box="[825,998,144,168]" italics="true" pageId="52" pageNumber="599">Metriorhynchidae</emphasis>
</taxonomicName>
. The new definition solves the
<taxonomicName id="4C794F93FF84730B7C5108800E8AFF38" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="52" pageNumber="599" phylum="Chordata" rank="species" species="superciliosus">
<emphasis id="B90DE802FF84730B7C5108800E8AFF38" italics="true" pageId="52" pageNumber="599">Thalattosuchus superciliosus</emphasis>
</taxonomicName>
issue, as in
<bibRefCitation id="EFE849E1FF84730B7D4C08BF09E1FF38" author="Wilberg EW" box="[1068,1243,175,199]" pageId="52" pageNumber="599" pagination="621 - 37" refId="ref59841" refString="Wilberg EW. What's in an outgroup? The impact of outgroup choice on the phylogenetic position of Thalattosuchia (Crocodylomorpha) and the origin of Crocodyliformes. Systematic Biology 2015 b; 64: 621 - 37. https: // doi. org / 10.1093 / sysbio / syv 020" type="journal article" year="2015">Wilberg (2015b)</bibRefCitation>
<taxonomicName id="4C794F93FF84730B7D8208BF0849FF38" authorityName="Young, Brignon, Sachs, Hornung, Foffa, Kitson, Johnson &amp; Steel" authorityYear="2020" box="[1250,1395,175,199]" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7D8208BF0849FF38" box="[1250,1395,175,199]" italics="true" pageId="52" pageNumber="599">Thalattosuchus</emphasis>
</taxonomicName>
was recovered as the sister-taxon to
<taxonomicName id="4C794F93FF84730B7D0908DF0EDEFEF9" authority="sensu Young and Andrade (2009)" authorityName="sensu Young and Andrade" authorityYear="2009" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="family">
<emphasis id="B90DE802FF84730B7D0908DF0875FF19" box="[1129,1359,207,231]" italics="true" pageId="52" pageNumber="599">Metriorhynchidae sensu</emphasis>
<bibRefCitation id="EFE849E1FF84730B7C3508DF0EDEFEF9" author="Young MT &amp; Andrade MB" pageId="52" pageNumber="599" pagination="551 - 85" refId="ref60527" refString="Young MT, Andrade MB. What is Geosaurus? Redescription of Geosaurus giganteus (Thalattosuchia, Metriorhynchidae) from the Upper Jurassic of Bayern, Germany. Zoological Journal of the Linnean Society 2009; 157: 551 - 85." type="journal article" year="2009">Young and Andrade (2009)</bibRefCitation>
</taxonomicName>
. Now, regardless of the phylogenetic position of this taxon,
<taxonomicName id="4C794F93FF84730B7AAF091D095AFEDA" authorityName="Young, Brignon, Sachs, Hornung, Foffa, Kitson, Johnson &amp; Steel" authorityYear="2020" box="[975,1120,269,293]" family="Metriorhynchidae" genus="Thalattosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7AAF091D095AFEDA" box="[975,1120,269,293]" italics="true" pageId="52" pageNumber="599">Thalattosuchus</emphasis>
</taxonomicName>
will always be a metriorhynchid. Moreover, the choice of five internal specifiers ensures consistent usage of the internal specifiers within
<taxonomicName id="4C794F93FF84730B7C76095C08F9FE9B" authorityName="sensu Young and Andrade" authorityYear="2009" box="[1302,1475,332,356]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="family">
<emphasis id="B90DE802FF84730B7C76095C08F9FE9B" box="[1302,1475,332,356]" italics="true" pageId="52" pageNumber="599">Metriorhynchidae</emphasis>
</taxonomicName>
and it’s the major subclades
<taxonomicName id="4C794F93FF84730B7D35097B083BFE7C" authorityName="Lydekker" authorityYear="1888" box="[1109,1281,363,387]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="subFamily" subFamily="Metriorhynchinae">
<emphasis id="B90DE802FF84730B7D35097B083BFE7C" box="[1109,1281,363,387]" italics="true" pageId="52" pageNumber="599">Metriorhynchinae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C794F93FF84730B7C52097C0891FE7C" authorityName="Bonaparte" authorityYear="1845" box="[1330,1451,364,387]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="subFamily" subFamily="Geosaurinae">
<emphasis id="B90DE802FF84730B7C52097C0891FE7C" box="[1330,1451,364,387]" italics="true" pageId="52" pageNumber="599">Geosaurinae</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BC63410FF84730B7A5909B5088CFD09" blockId="52.[825,1475,421,1979]" pageId="52" pageNumber="599">
<emphasis id="B90DE802FF84730B7A5909B50E9DFE43" box="[825,935,421,444]" italics="true" pageId="52" pageNumber="599">Discussion:</emphasis>
Within
<taxonomicName id="4C794F93FF84730B7D6809B509A5FE42" authorityName="sensu Fraas" authorityYear="1901" box="[1032,1183,421,445]" class="Reptilia" higherTaxonomySource="GBIF" infraOrder="Thalattosuchia" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="582" phylum="Chordata" rank="infraOrder">Thalattosuchia</taxonomicName>
,
<taxonomicName id="4C794F93FF84730B7DCF09B50853FE42" authorityName="sensu Young and Andrade" authorityYear="2009" box="[1199,1385,421,445]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="family">Metriorhynchidae</taxonomicName>
are the most diagnostic clade (
<figureCitation id="13422895FF84730B7D5409D4094EFE23" box="[1076,1140,452,476]" captionStart="Figure 1" captionStartId="1.[113,178,1773,1797]" captionTargetBox="[117,1455,145,1742]" captionTargetId="figure-7@1.[114,1458,142,1745]" captionTargetPageId="1" captionText="Figure 1. Comparing the body plans of semi-aquatic thalattosuchians (A–C) with that of a fully aquatic metriorhynchid (D). A, the teleosaurid teleosauroid Platysuchus multiscrobiculatus, holotype SMNS 9930; B, the machimosaurid teleosauroid Macrospondylus bollensis, referred specimen NMW 13445; C, the early diverging metriorhynchoid Pelagosaurus typus, referred specimen FWD 0784; D, the metriorhynchid Cricosaurus suevicus, lectotype SMNS 9808. All scale bars equal 20 cm.Even amongst the semi-aquatic thalattosuchians there is variation, such as the greater size and coverage of osteoderms in Platysuchus, with Pelagosaurus having the most reduced armour of the three." figureDoi="http://doi.org/10.5281/zenodo.11241247" httpUri="https://zenodo.org/record/11241247/files/figure.png" pageId="52" pageNumber="599">Figs 1</figureCitation>
,
<figureCitation id="13422895FF84730B7DE409D409A4FE23" box="[1156,1182,452,476]" captionStart="Figure 22" captionStartId="54.[129,195,1836,1860]" captionTargetBox="[226,1376,147,1806]" captionTargetId="figure-7@54.[223,1379,144,1808]" captionTargetPageId="54" captionText="Figure 22. Comparative plate of metriorhynchid skeletons. A, Thalattosuchus superciliosus, referred specimen GPIT-PV-31379; B, Rhacheosaurus gracilis, referred specimen NHMUK PV R 3948; C, Cricosaurus albersdoerferi, holotype BMMS-BK 1-2. All scale bars equal 30 cm." figureDoi="http://doi.org/10.5281/zenodo.11241289" httpUri="https://zenodo.org/record/11241289/files/figure.png" pageId="52" pageNumber="599">22</figureCitation>
). Unfortunately, the evolutionary timing of many of the postcranial characters that define the group is unknown. This is a consequence of most of the non-metriorhynchid metriorhynchoids having little to no known postcranial skeleton. As such, the diagnostic characters metriorhynchids have in entire regions of the skeleton are still incompletely understood, ranging from their loss of osteoderms (
<figureCitation id="13422895FF84730B7A240A8F0EB1FD48" box="[836,907,671,695]" captionStart="Figure 22" captionStartId="54.[129,195,1836,1860]" captionTargetBox="[226,1376,147,1806]" captionTargetId="figure-7@54.[223,1379,144,1808]" captionTargetPageId="54" captionText="Figure 22. Comparative plate of metriorhynchid skeletons. A, Thalattosuchus superciliosus, referred specimen GPIT-PV-31379; B, Rhacheosaurus gracilis, referred specimen NHMUK PV R 3948; C, Cricosaurus albersdoerferi, holotype BMMS-BK 1-2. All scale bars equal 30 cm." figureDoi="http://doi.org/10.5281/zenodo.11241289" httpUri="https://zenodo.org/record/11241289/files/figure.png" pageId="52" pageNumber="599">Fig. 22</figureCitation>
), modification of the manus into flippers (
<figureCitation id="13422895FF84730B7C280A8F08B5FD48" box="[1352,1423,671,695]" captionStart="Figure 23" captionStartId="55.[115,180,644,668]" captionTargetBox="[117,1455,147,614]" captionTargetId="figure-65@55.[114,1458,144,616]" captionTargetPageId="55" captionText="Figure 23. Comparative plate of thalattosuchian coracoids and forelimbs.A, the teleosaurid teleosauroid Platysuchus multiscrobiculatus, holotype SMNS 9930; B, the machimosaurid teleosauroid Macrospondylus bollensis, referred specimen NMW 13445; C, the early diverging metriorhynchoid Pelagosaurus typus, referred specimen FWD 0784; D, the metriorhynchid Cricosaurus bambergensis, holotype NKMB-P- Watt14/274. All scale bars equal 5 cm.Highlighting the changes as thalattosuchians became increasingly aquatic, such as the flattened and ‘double fan’-shaped coracoids, proportional reduction and flattening of the forelimb elements, and modification of the manus into a flipper." figureDoi="http://doi.org/10.5281/zenodo.11241291" httpUri="https://zenodo.org/record/11241291/files/figure.png" pageId="52" pageNumber="599">Fig. 23</figureCitation>
), reduction of pelvis size, modification of the hindlimbs into paddles (
<figureCitation id="13422895FF84730B7A240ACE0EB3FD09" box="[836,905,734,758]" captionStart="Figure 24" captionStartId="55.[113,178,1725,1749]" captionTargetBox="[117,1455,830,1694]" captionTargetId="figure-156@55.[114,1458,827,1696]" captionTargetPageId="55" captionText="Figure 24. Comparative plate of thalattosuchian pelvic girdles and hindlimbs. A, the teleosaurid teleosauroid Platysuchus multiscrobiculatus, holotype SMNS 9930; B, the machimosaurid teleosauroid Macrospondylus bollensis, referred specimen NMW 13445; C, the early diverging metriorhynchoid Pelagosaurus typus, referred specimen FWD 0784; D, the metriorhynchid Cricosaurus albersdoerferi, holotype BMMS-BK 1-2. All scale bars equal 10 cm. Highlighting the changes as thalattosuchians became increasingly aquatic, such as the loss of osteoderms, proportional reduction of the ilium and tibiae, strongly ventrally curved sacral ribs, and modification of the pes into a paddle." figureDoi="http://doi.org/10.5281/zenodo.11241293" httpUri="https://zenodo.org/record/11241293/files/figure.png" pageId="52" pageNumber="599">Fig. 24</figureCitation>
), and the development of the hypocercal tail (
<figureCitation id="13422895FF84730B7C010ACE089CFD09" box="[1377,1446,734,758]" captionStart="Figure 25" captionStartId="56.[129,194,1011,1035]" captionTargetBox="[132,1470,147,982]" captionTargetId="figure-300@56.[129,1473,144,984]" captionTargetPageId="56" captionText="Figure 25. Comparative plate of thalattosuchian distal tails. A, the machimosaurid teleosauroid Macrospondylus bollensis, referred specimen SMNS 51584; B, the early diverging metriorhynchoid Pelagosaurus typus, referred specimen MTM M62 2516; C, the metriorhynchid Thalattosuchus superciliosus, referred specimen GPIT-PV-31590; D, the metriorhynchid Cricosaurus bambergensis, holotype NKMB-P- Watt14/274. All scale bars equal 10 cm. Highlighting the shift in distal tail morphology as thalattosuchians became increasingly aquatic, culminating in the hypocercal tail of metriorhynchids. Note the variation in tail fluke morphology within Metriorhynchidae." figureDoi="http://doi.org/10.5281/zenodo.11241297" httpUri="https://zenodo.org/record/11241297/files/figure.png" pageId="52" pageNumber="599">Fig. 25</figureCitation>
).
</paragraph>
<paragraph id="8BC63410FF84730B7A350AED09A3FC0F" blockId="52.[825,1475,421,1979]" pageId="52" pageNumber="599">
The body-plan diversity within
<taxonomicName id="4C794F93FF84730B7DC80AED0858FCEA" authorityName="sensu Young and Andrade" authorityYear="2009" box="[1192,1378,765,789]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="family">Metriorhynchidae</taxonomicName>
is as remarkable as their divergence from semi-aquatic thalattosuchians (
<figureCitation id="13422895FF84730B7A240B2C0E45FCAB" box="[836,895,828,852]" captionStart="Figure 1" captionStartId="1.[113,178,1773,1797]" captionTargetBox="[117,1455,145,1742]" captionTargetId="figure-7@1.[114,1458,142,1745]" captionTargetPageId="1" captionText="Figure 1. Comparing the body plans of semi-aquatic thalattosuchians (A–C) with that of a fully aquatic metriorhynchid (D). A, the teleosaurid teleosauroid Platysuchus multiscrobiculatus, holotype SMNS 9930; B, the machimosaurid teleosauroid Macrospondylus bollensis, referred specimen NMW 13445; C, the early diverging metriorhynchoid Pelagosaurus typus, referred specimen FWD 0784; D, the metriorhynchid Cricosaurus suevicus, lectotype SMNS 9808. All scale bars equal 20 cm.Even amongst the semi-aquatic thalattosuchians there is variation, such as the greater size and coverage of osteoderms in Platysuchus, with Pelagosaurus having the most reduced armour of the three." figureDoi="http://doi.org/10.5281/zenodo.11241247" httpUri="https://zenodo.org/record/11241247/files/figure.png" pageId="52" pageNumber="599">Fig. 1</figureCitation>
). There is variation in skull shape, particularly between dakosaurins and other subclades (
<figureCitation id="13422895FF84730B7DEF0B4B09EDFC8C" box="[1167,1239,859,883]" captionStart="Figure 26" captionStartId="57.[113,178,1594,1618]" captionTargetBox="[117,1455,147,1563]" captionTargetId="figure-130@57.[114,1458,144,1566]" captionTargetPageId="57" captionText="Figure 26. Comparative plate of metriorhynchid skulls in dorsal view.A, the early diverging Thalattosuchus superciliosus, lectotype MNHN.F RJN 256; B, the rhacheosaurin Cricosaurus araucanensis, holotype MLP 72-IV-7-1; C, the dakosaurin Dakosaurus andiniensis, referred specimen MOZ-PV 6146.All scale bars equal 10 cm." figureDoi="http://doi.org/10.5281/zenodo.11241299" httpUri="https://zenodo.org/record/11241299/files/figure.png" pageId="52" pageNumber="599">Figs 26</figureCitation>
,
<figureCitation id="13422895FF84730B7D800B4B09C0FC8C" box="[1248,1274,859,883]" captionStart="Figure 27" captionStartId="58.[129,194,1830,1854]" captionTargetBox="[132,1470,147,1800]" captionTargetId="figure-6@58.[129,1473,144,1803]" captionTargetPageId="58" captionText="Figure 27. Comparative plate of metriorhynchid skulls in lateral view. A, the rhacheosaurin Cricosaurus suevicus, lectotype SMNS 9808; B, the early diverging Thalattosuchus superciliosus, lectotype MNHN.F RJN 256; C, the dakosaurin Dakosaurus maximus, neotype SMNS 8203. All scale bars equal 10 cm." figureDoi="http://doi.org/10.5281/zenodo.11241301" httpUri="https://zenodo.org/record/11241301/files/figure.png" pageId="52" pageNumber="599">27</figureCitation>
), and in postcranial morphology, particularly in the angle and morphology of the tail fluke (
<figureCitation id="13422895FF84730B7AB00B8A0926FC4D" box="[976,1052,922,946]" captionStart="Figure 22" captionStartId="54.[129,195,1836,1860]" captionTargetBox="[226,1376,147,1806]" captionTargetId="figure-7@54.[223,1379,144,1808]" captionTargetPageId="54" captionText="Figure 22. Comparative plate of metriorhynchid skeletons. A, Thalattosuchus superciliosus, referred specimen GPIT-PV-31379; B, Rhacheosaurus gracilis, referred specimen NHMUK PV R 3948; C, Cricosaurus albersdoerferi, holotype BMMS-BK 1-2. All scale bars equal 30 cm." figureDoi="http://doi.org/10.5281/zenodo.11241289" httpUri="https://zenodo.org/record/11241289/files/figure.png" pageId="52" pageNumber="599">Figs 22</figureCitation>
,
<figureCitation id="13422895FF84730B7D4A0B8A097EFC4D" box="[1066,1092,922,946]" captionStart="Figure 25" captionStartId="56.[129,194,1011,1035]" captionTargetBox="[132,1470,147,982]" captionTargetId="figure-300@56.[129,1473,144,984]" captionTargetPageId="56" captionText="Figure 25. Comparative plate of thalattosuchian distal tails. A, the machimosaurid teleosauroid Macrospondylus bollensis, referred specimen SMNS 51584; B, the early diverging metriorhynchoid Pelagosaurus typus, referred specimen MTM M62 2516; C, the metriorhynchid Thalattosuchus superciliosus, referred specimen GPIT-PV-31590; D, the metriorhynchid Cricosaurus bambergensis, holotype NKMB-P- Watt14/274. All scale bars equal 10 cm. Highlighting the shift in distal tail morphology as thalattosuchians became increasingly aquatic, culminating in the hypocercal tail of metriorhynchids. Note the variation in tail fluke morphology within Metriorhynchidae." figureDoi="http://doi.org/10.5281/zenodo.11241297" httpUri="https://zenodo.org/record/11241297/files/figure.png" pageId="52" pageNumber="599">25</figureCitation>
). While we have learnt much about metriorhynchids over the past two hundred years, there is still much more we do not understand.
</paragraph>
<paragraph id="8BC63410FF84730B7A350BE80804F99D" blockId="52.[825,1475,421,1979]" pageId="52" pageNumber="599">
Much like the previous use of
<taxonomicName id="4C794F93FF84730B7DEC0BE80830FBF0" box="[1164,1290,1016,1039]" class="Reptilia" family="Teleosauridae" genus="Steneosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7DEC0BE80830FBF0" box="[1164,1290,1016,1039]" italics="true" pageId="52" pageNumber="599">Steneosaurus</emphasis>
</taxonomicName>
as a ‘wastebasket’ taxon, the continued use of
<taxonomicName id="4C794F93FF84730B7D0D0C07083CFBD0" authorityName="von Meyer" authorityYear="1832" box="[1133,1286,1047,1071]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7D0D0C07083CFBD0" box="[1133,1286,1047,1071]" italics="true" pageId="52" pageNumber="599">Metriorhynchus</emphasis>
</taxonomicName>
to place the majority of Middle Jurassic metriorhynchid species (and many Late Jurassic ones) has greatly held back our understanding of the clade. Based on the phylogenetic analyses presented herein (
<figureCitation id="13422895FF84730B7A240C840E90FB53" box="[836,938,1172,1196]" captionStart-0="Figure 3" captionStart-1="Figure 4" captionStart-2="Figure 5" captionStart-3="Figure 6" captionStart-4="Figure 7" captionStart-5="Figure 8" captionStart-6="Figure 9" captionStart-7="Figure 10" captionStart-8="Figure 11" captionStart-9="Figure 12" captionStartId-0="19.[113,178,1185,1209]" captionStartId-1="21.[113,178,1219,1243]" captionStartId-2="22.[128,193,1219,1243]" captionStartId-3="23.[113,178,1244,1268]" captionStartId-4="24.[129,194,1442,1466]" captionStartId-5="25.[113,178,1707,1731]" captionStartId-6="27.[113,178,1246,1270]" captionStartId-7="28.[129,194,1301,1325]" captionStartId-8="29.[113,178,1287,1311]" captionStartId-9="30.[128,193,1319,1343]" captionTargetBox-0="[117,1455,147,1154]" captionTargetBox-1="[117,1455,147,1190]" captionTargetBox-2="[132,1470,147,1188]" captionTargetBox-3="[117,1455,147,1213]" captionTargetBox-4="[132,1470,147,1411]" captionTargetBox-5="[117,1455,553,1676]" captionTargetBox-6="[117,1455,147,1215]" captionTargetBox-7="[132,1470,147,1270]" captionTargetBox-8="[117,1455,147,1256]" captionTargetBox-9="[132,1470,147,1289]" captionTargetId-0="figure-283@19.[114,1458,144,1157]" captionTargetId-1="figure-205@21.[114,1458,144,1192]" captionTargetId-2="figure-220@22.[129,1473,144,1191]" captionTargetId-3="figure-242@23.[114,1458,144,1216]" captionTargetId-4="figure-150@24.[129,1473,144,1414]" captionTargetId-5="figure-191@25.[114,1458,550,1679]" captionTargetId-6="figure-185@27.[114,1458,144,1218]" captionTargetId-7="figure-212@28.[129,1473,144,1273]" captionTargetId-8="figure-196@29.[114,1458,144,1259]" captionTargetId-9="figure-177@30.[129,1473,144,1291]" captionTargetPageId-0="19" captionTargetPageId-1="21" captionTargetPageId-2="22" captionTargetPageId-3="23" captionTargetPageId-4="24" captionTargetPageId-5="25" captionTargetPageId-6="27" captionTargetPageId-7="28" captionTargetPageId-8="29" captionTargetPageId-9="30" captionText-0="Figure 3. The equal weights, no constraints strict consensus topology (C0). Length = 4164, CI = 0.303, RI = 0.831, RC = 0.251, HI = 0.697, number of most parsimonious cladograms hit the hold limit.Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig. S1 for the entire topology). A, Crocodylomorpha; B, Crocodyliformes; C, Thalattosuchia; D, Neothalattosuchia; E, Teleosauridae; F, Teleosaurinae; G, Aeolodontini; H, Machimosauridae; I, Machimosaurinae; J, Machimosaurini; K, Euthalattosuchia; L, Metriorhynchidae; M, Metriorhynchinae; N, Rhacheosaurini; O, Geosaurinae; P, Geosaurini. Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians.Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T. Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons.org/licenses/by/3.0/." captionText-1="Figure 4. The equal weights, crocodyliform constraint strict consensus topology (C1). Length = 4177, CI = 0.302, RI = 0.830, RC = 0.201, HI = 0.698, number of most parsimonious cladograms hit the hold limit. Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig. S2 for the entire topology). A, Crocodylomorpha; B, Thalattosuchia; C, Neothalattosuchia; D, Teleosauridae; E, Teleosaurinae; F, Aeolodontini; G, Machimosauridae; H, Machimosaurinae; I, Machimosaurini; J, Euthalattosuchia; K, Metriorhynchidae; L, Metriorhynchinae; M, Rhacheosaurini; N, Geosaurinae; O, Geosaurini.Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians. Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T. Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons.org/licenses/by/3.0/." captionText-2="Figure 5. The equal weights, mesoeucrocodylian constraint strict consensus topology (C2). Length = 4164, CI = 0.303, RI = 0.831, RC = 0.251, HI = 0.697, number of most parsimonious cladograms hit the hold limit. Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig.S3 for the entire topology). A, Crocodylomorpha; B, Crocodyliformes; C, Thalattosuchia; D, Neothalattosuchia; E, Teleosauridae; F, Teleosaurinae; G, Aeolodontini; H, Machimosauridae; I, Machimosaurinae; J, Machimosaurini; K, Euthalattosuchia; L, Metriorhynchidae; M, Metriorhynchinae; N, Rhacheosaurini; O, Geosaurinae; P, Geosaurini. Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians. Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T.Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons.org/licenses/ by/3.0/." captionText-3="Figure 6. The equal weights, neosuchian constraint strict consensus topology (C3). Length = 4174, CI = 0.302, RI = 0.830, RC = 0.251, HI = 0.698, number of most parsimonious cladograms hit the hold limit. Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig. S4 for the entire topology). A, Crocodylomorpha; B, Crocodyliformes; C, Metasuchia; D, Neosuchia; E, Thalattosuchia; F, Neothalattosuchia; G, Teleosauridae; H, Teleosaurinae; I, Aeolodontini; J, Machimosauridae; K, Machimosaurinae; L, Machimosaurini; M, Euthalattosuchia; N, Metriorhynchidae; O, Metriorhynchinae; P, Rhacheosaurini; Q, Geosaurinae; R, Geosaurini. Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians. Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T. Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons. org/licenses/by/3.0/." captionText-4="Figure 7. The equal weights, tethysuchian constraint strict consensus topology (C4). Length = 4172, CI = 0.302, RI = 0.830, RC = 0.251, HI = 0.698, number of most parsimonious cladograms hit the hold limit. Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig. S5 for the entire topology). A, Crocodylomorpha; B, Crocodyliformes; C, Metasuchia; D, Neosuchia; E, Thalattosuchia; F, Neothalattosuchia; G, Teleosaurinae; H, Aeolodontini; I, Machimosaurinae; J, Machimosaurini; K, Metriorhynchidae; L, Metriorhynchinae; M, Rhacheosaurini; N, Geosaurinae; O, Geosaurini.Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians. Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T.Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons.org/licenses/by/3.0/." captionText-5="Figure 8. The extended implied weighting (k = 15), no constraints strict consensus topology (C0). Length = 4178, fit = 111.578, CI = 0.302, RI = 0.830, RC = 0.250, HI = 0.698, number of most parsimonious cladograms = 4500. Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig.S6 for the entire topology, Appendix Two).A, Crocodylomorpha; B, Crocodyliformes; C, Thalattosuchia; D, Neothalattosuchia; E, Teleosauridae; F, Teleosaurinae; G, Aeolodontini; H, Machimosauridae; I, Machimosaurinae; J, Machimosaurini; K, Euthalattosuchia; L, Metriorhynchidae; M, Metriorhynchinae; N, Rhacheosaurini; O, Geosaurinae; P, Geosaurini. Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians. Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T. Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons. org/licenses/by/3.0/." captionText-6="Figure 9. The extended implied weighting (k = 15), crocodyliform constraint strict consensus topology (C1). Length = 4194, fit = 111.226, CI = 0.300, RI = 0.829, RC = 0.249, HI = 0.700, number of most parsimonious cladograms = 4500.Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig. S7 for the entire topology). A, Crocodylomorpha; B, Thalattosuchia; C, Neothalattosuchia; D, Teleosauridae; E, Teleosaurinae; F, Aeolodontini; G, Machimosauridae; H, Machimosaurinae; I, Machimosaurini; J, Euthalattosuchia; K, Metriorhynchidae; L, Metriorhynchinae; M, Rhacheosaurini; N, Geosaurinae; O, Geosaurini. Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians. Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T.Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons.org/licenses/ by/3.0/." captionText-7="Figure 10. The extended implied weighting (k = 15), mesoeucrocodylian constraint strict consensus topology (C2). Length = 4178, fit = 111.578, CI = 0.302, RI = 0.830, RC = 0.250, HI = 0.698, number of most parsimonious cladograms = 4500.Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig.S8 for the entire topology). A, Crocodylomorpha; B, Crocodyliformes; C, Thalattosuchia; D, Neothalattosuchia; E, Teleosauridae; F, Teleosaurinae; G, Aeolodontini; H, Machimosauridae; I, Machimosaurinae; J, Machimosaurini; K, Euthalattosuchia; L, Metriorhynchidae; M, Metriorhynchinae; N, Rhacheosaurini; O, Geosaurinae; P, Geosaurini. Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians.Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T. Michael Keesey), the only change made was to their colouration, and their license is: https:// creativecommons.org/licenses/by/3.0/." captionText-8="Figure 11. The extended implied weighting (k = 15), neosuchian constraint strict consensus topology (C3). Length = 4192, fit = 112.001, CI = 0.301, RI = 0.829, RC = 0.249, HI = 0.699, number of most parsimonious cladograms = 4500.Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig. S9 for the entire topology, Appendix Two). A, Crocodylomorpha; B, Crocodyliformes; C, Metasuchia; D, Neosuchia; E, Thalattosuchia; F, Neothalattosuchia; G, Teleosauridae; H, Teleosaurinae; I, Aeolodontini; J, Machimosauridae; K, Machimosaurinae; L, Machimosaurini; M, Euthalattosuchia; N, Metriorhynchidae; O, Metriorhynchinae; P, Rhacheosaurini; Q, Geosaurinae; R, Geosaurini. Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians. Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T. Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons.org/licenses/by/3.0/." captionText-9="Figure 12. The extended implied weighting (k = 15), tethysuchian constraint strict consensus topology (C4). Length = 4189, fit = 112.271, CI = 0.301, RI = 0.829, RC = 0.249, HI = 0.699, number of most parsimonious cladograms = 4500.Phylogeny pruned to highlight Thalattosuchia (see Supporting Information, Appendix S2, Fig. S10 for the entire topology). A, Crocodylomorpha; B, Crocodyliformes; C, Metasuchia; D, Neosuchia; E, Thalattosuchia; F, Neothalattosuchia; G, Teleosauridae; H, Teleosaurinae; I, Aeolodontini; J, Machimosauridae; K, Machimosaurinae; L, Machimosaurini; M, Euthalattosuchia; N, Metriorhynchidae; O, Metriorhynchinae; P, Rhacheosaurini; Q, Geosaurinae; R, Geosaurini. Green lines highlight non-neothalattosuchian thalattosuchians, orange lines highlight teleosauroid thalattosuchians, and blue lines highlight metriorhynchoid thalattosuchians. Images from PhyloPics.org, with the teleosauroid by Scott Hartman, and the metriorhynchid by Dmitry Bogdanov (vectorized by T.Michael Keesey), the only change made was to their colouration, and their license is: https://creativecommons.org/licenses/by/3.0/." figureDoi-0="http://doi.org/10.5281/zenodo.11241251" figureDoi-1="http://doi.org/10.5281/zenodo.11241253" figureDoi-2="http://doi.org/10.5281/zenodo.11241255" figureDoi-3="http://doi.org/10.5281/zenodo.11241257" figureDoi-4="http://doi.org/10.5281/zenodo.11241259" figureDoi-5="http://doi.org/10.5281/zenodo.11241261" figureDoi-6="http://doi.org/10.5281/zenodo.11241263" figureDoi-7="http://doi.org/10.5281/zenodo.11241265" figureDoi-8="http://doi.org/10.5281/zenodo.11241267" figureDoi-9="http://doi.org/10.5281/zenodo.11241269" httpUri-0="https://zenodo.org/record/11241251/files/figure.png" httpUri-1="https://zenodo.org/record/11241253/files/figure.png" httpUri-2="https://zenodo.org/record/11241255/files/figure.png" httpUri-3="https://zenodo.org/record/11241257/files/figure.png" httpUri-4="https://zenodo.org/record/11241259/files/figure.png" httpUri-5="https://zenodo.org/record/11241261/files/figure.png" httpUri-6="https://zenodo.org/record/11241263/files/figure.png" httpUri-7="https://zenodo.org/record/11241265/files/figure.png" httpUri-8="https://zenodo.org/record/11241267/files/figure.png" httpUri-9="https://zenodo.org/record/11241269/files/figure.png" pageId="52" pageNumber="599">Figs 3–12</figureCitation>
), the only way to render
<taxonomicName id="4C794F93FF84730B7DD70C84086AFB53" authorityName="von Meyer" authorityYear="1832" box="[1207,1360,1172,1196]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7DD70C84086AFB53" box="[1207,1360,1172,1196]" italics="true" pageId="52" pageNumber="599">Metriorhynchus</emphasis>
</taxonomicName>
monophyletic would be to place all metriorhynchoids more derived than
<taxonomicName id="4C794F93FF84730B7A590CC30EA4FB14" box="[825,926,1235,1259]" genus="Opisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7A590CC30EA4FB14" box="[825,926,1235,1259]" italics="true" pageId="52" pageNumber="599">Opisuchus</emphasis>
</taxonomicName>
into the genus
<taxonomicName id="4C794F93FF84730B7D290CC309D8FB14" authorityName="von Meyer" authorityYear="1832" box="[1097,1250,1235,1259]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7D290CC309D8FB14" box="[1097,1250,1235,1259]" italics="true" pageId="52" pageNumber="599">Metriorhynchus</emphasis>
</taxonomicName>
(as
<taxonomicName id="4C794F93FF84730B7C730CC4084BFB14" authorityName="Young, Brusatte, Ruta &amp; Andrade" authorityYear="2010" box="[1299,1393,1236,1259]" class="Reptilia" genus="Eoneustes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7C730CC4084BFB14" box="[1299,1393,1236,1259]" italics="true" pageId="52" pageNumber="599">Eoneustes</emphasis>
</taxonomicName>
, placed within the genus
<taxonomicName id="4C794F93FF84730B7A8E0CE209BDFAF5" authorityName="von Meyer" authorityYear="1832" box="[1006,1159,1266,1290]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7A8E0CE209BDFAF5" box="[1006,1159,1266,1290]" italics="true" pageId="52" pageNumber="599">Metriorhynchus</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFE849E1FF84730B7DCE0CE20800FAF5" author="Mercier J" box="[1198,1338,1266,1290]" pageId="52" pageNumber="599" pagination="99 - 119" refId="ref56612" refString="Mercier J. Contribution a l'etude des Metriorhynchides (crocodiliens). Annales de Paleontologie 1933; 22: 99 - 119." type="journal article" year="1933">Mercier 1933</bibRefCitation>
, is recovered outside of Euthalattosuchia). The only other way to use the genus
<taxonomicName id="4C794F93FF84730B7A1C0D21092FFAB6" authorityName="von Meyer" authorityYear="1832" box="[892,1045,1329,1353]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7A1C0D21092FFAB6" box="[892,1045,1329,1353]" italics="true" pageId="52" pageNumber="599">Metriorhynchus</emphasis>
</taxonomicName>
as ‘traditionally’ defined would be to accept that it is a paraphyletic taxon (and accept the usage of paraphyletic taxa), one from which the other traditionally accepted metriorhynchid genera (
<taxonomicName id="4C794F93FF84730B7DF90D9F09C5FA59" authorityName="Bonaparte" authorityYear="1850" box="[1177,1279,1423,1446]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7DF90D9F09C5FA59" box="[1177,1279,1423,1446]" italics="true" pageId="52" pageNumber="599">Geosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C794F93FF84730B7C220D9F088DFA58" box="[1346,1463,1423,1447]" class="Reptilia" family="Metriorhynchidae" genus="Dakosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7C220D9F088DFA58" box="[1346,1463,1423,1447]" italics="true" pageId="52" pageNumber="599">Dakosaurus</emphasis>
</taxonomicName>
) evolved. That would place Euthalattosuchia,
<taxonomicName id="4C794F93FF84730B7C630DBE0885FA39" authorityName="sensu Young and Andrade" authorityYear="2009" box="[1283,1471,1454,1478]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="family">Metriorhynchidae</taxonomicName>
,
<taxonomicName id="4C794F93FF84730B7A590DDE0ECFFA19" authorityName="Lydekker" authorityYear="1888" box="[825,1013,1486,1510]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="subFamily" subFamily="Metriorhynchinae">Metriorhynchinae</taxonomicName>
, Rhacheosaurini,
<taxonomicName id="4C794F93FF84730B7D8E0DDE084AFA19" authorityName="Bonaparte" authorityYear="1845" box="[1262,1392,1486,1510]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="subFamily" subFamily="Geosaurinae">Geosaurinae</taxonomicName>
, and
<taxonomicName id="4C794F93FF84730B7A590DFD0E90F9FA" authorityName="Cau and Fanti" authorityYear="2011" box="[825,938,1517,1541]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="tribe" tribe="Geosaurini">Geosaurini</taxonomicName>
as subclades within
<taxonomicName id="4C794F93FF84730B7D1D0DFD082CF9FA" authorityName="von Meyer" authorityYear="1832" box="[1149,1302,1517,1541]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7D1D0DFD082CF9FA" box="[1149,1302,1517,1541]" italics="true" pageId="52" pageNumber="599">Metriorhynchus</emphasis>
</taxonomicName>
. Again, like with the
<taxonomicName id="4C794F93FF84730B7A010E1D0EE5F9DB" box="[865,991,1549,1572]" class="Reptilia" family="Teleosauridae" genus="Steneosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7A010E1D0EE5F9DB" box="[865,991,1549,1572]" italics="true" pageId="52" pageNumber="599">Steneosaurus</emphasis>
</taxonomicName>
example above, this would subsume a remarkable level of morphological variation into a paraphyletic genus, creating a genus defined primarily by what it is not.
</paragraph>
<paragraph id="8BC63410FF84730A7A350E7B0C00FD42" blockId="52.[825,1475,421,1979]" lastBlockId="53.[113,763,144,1264]" lastPageId="53" lastPageNumber="600" pageId="52" pageNumber="599">
It is worth noting that the characters Eudes-Deslongchamps (1867–69: 132–133) used to redefine
<taxonomicName id="4C794F93FF84730B7DA50E990864F95E" authorityName="von Meyer" authorityYear="1832" box="[1221,1374,1673,1697]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7DA50E990864F95E" box="[1221,1374,1673,1697]" italics="true" pageId="52" pageNumber="599">Metriorhynchus</emphasis>
</taxonomicName>
are today known to be present in all metriorhynchids or are variable within the clade (see: Young
<emphasis id="B90DE802FF84730B7D4B0ED90962F91F" box="[1067,1112,1736,1760]" italics="true" pageId="52" pageNumber="599">et al</emphasis>
. 2021: 518). It, therefore, should not be a surprise that if we attempt to apply a 19th century ergotaxonomytoday, aframeworkcreatedwhenalimitednumber of metriorhynchid fossils were known, it will result in a paraphyletic taxon. Moreover, this was widely known and accepted in the early 20th century, as in the hand-drawn phylogenies of
<bibRefCitation id="EFE849E1FF84730B7CEC0F750EB9F863" author="Fraas E" pageId="52" pageNumber="599" pagination="1 - 72" refId="ref51930" refString="Fraas E. Die Meer-Krocodilier (Thalattosuchia) des oberen Jura unter specieller Berucksichtigung von Dacosaurus und Geosaurus. Palaeontographica 1902; 49: 1 - 72." type="journal article" year="1902">Fraas (1902)</bibRefCitation>
and
<bibRefCitation id="EFE849E1FF84730B7ADF0F94095FF863" author="Mercier J" box="[959,1125,1924,1948]" pageId="52" pageNumber="599" pagination="99 - 119" refId="ref56612" refString="Mercier J. Contribution a l'etude des Metriorhynchides (crocodiliens). Annales de Paleontologie 1933; 22: 99 - 119." type="journal article" year="1933">Mercier (1933)</bibRefCitation>
<taxonomicName id="4C794F93FF84730B7D100F940833F863" authorityName="von Meyer" authorityYear="1832" box="[1136,1289,1924,1948]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7D100F940833F863" box="[1136,1289,1924,1948]" italics="true" pageId="52" pageNumber="599">Metriorhynchus</emphasis>
</taxonomicName>
was paraphyletic with regard to
<taxonomicName id="4C794F93FF84730B7ABE0FB4097EF844" authorityName="Bonaparte" authorityYear="1850" box="[990,1092,1956,1979]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7ABE0FB4097EF844" box="[990,1092,1956,1979]" italics="true" pageId="52" pageNumber="599">Geosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C794F93FF84730B7D1F0FB309CEF844" box="[1151,1268,1955,1979]" class="Reptilia" family="Metriorhynchidae" genus="Dakosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="52" pageNumber="599" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF84730B7D1F0FB309CEF844" box="[1151,1268,1955,1979]" italics="true" pageId="52" pageNumber="599">Dakosaurus</emphasis>
</taxonomicName>
. What complicates matters further, is that
<taxonomicName id="4C794F93FF85730A781608800CE6FF58" authorityName="Bonaparte" authorityYear="1850" box="[374,476,144,167]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A781608800CE6FF58" box="[374,476,144,167]" italics="true" pageId="53" pageNumber="600">Geosaurus</emphasis>
</taxonomicName>
was polyphyletic in both ergotaxonomic schemes as well. This resulted in a systematization that placed all Middle Jurassic metriorhynchids in the genus
<taxonomicName id="4C794F93FF85730A791108FE0C30FEF9" authorityName="von Meyer" authorityYear="1832" box="[113,266,238,262]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A791108FE0C30FEF9" box="[113,266,238,262]" italics="true" pageId="53" pageNumber="600">Metriorhynchus</emphasis>
</taxonomicName>
, all ‘robust’ Late Jurassic metriorhynchids in the genus
<taxonomicName id="4C794F93FF85730A79D4091D0C13FEDA" box="[180,297,269,293]" class="Reptilia" family="Metriorhynchidae" genus="Dakosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A79D4091D0C13FEDA" box="[180,297,269,293]" italics="true" pageId="53" pageNumber="600">Dakosaurus</emphasis>
</taxonomicName>
, all ‘gracile’ Late Jurassic metriorhynchids in the genus
<taxonomicName id="4C794F93FF85730A79B7093D0C07FEBB" authorityName="Bonaparte" authorityYear="1850" box="[215,317,301,324]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A79B7093D0C07FEBB" box="[215,317,301,324]" italics="true" pageId="53" pageNumber="600">Geosaurus</emphasis>
</taxonomicName>
, and all other Late Jurassic metriorhynchids were placed in
<taxonomicName id="4C794F93FF85730A787F095C0C82FE9B" authorityName="von Meyer" authorityYear="1832" box="[287,440,332,356]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A787F095C0C82FE9B" box="[287,440,332,356]" italics="true" pageId="53" pageNumber="600">Metriorhynchus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFE849E1FF85730A78AF095C0F7CFE9B" author="Fraas E" box="[463,582,332,356]" pageId="53" pageNumber="600" pagination="1 - 72" refId="ref51930" refString="Fraas E. Die Meer-Krocodilier (Thalattosuchia) des oberen Jura unter specieller Berucksichtigung von Dacosaurus und Geosaurus. Palaeontographica 1902; 49: 1 - 72." type="journal article" year="1902">Fraas 1902</bibRefCitation>
,
<bibRefCitation id="EFE849E1FF85730A7B38095C0FCFFE9B" author="Andrews CW" box="[600,757,332,356]" pageId="53" pageNumber="600" refId="ref48442" refString="Andrews CW. A Descriptive Catalogue of the Marine Reptiles of the Oxford Clay, Part Two. London: British Museum (Natural History), 1913." type="book" year="1913">Andrews 1913</bibRefCitation>
,
<bibRefCitation id="EFE849E1FF85730A7911097B0C30FE7C" author="Mercier J" box="[113,266,363,387]" pageId="53" pageNumber="600" pagination="99 - 119" refId="ref56612" refString="Mercier J. Contribution a l'etude des Metriorhynchides (crocodiliens). Annales de Paleontologie 1933; 22: 99 - 119." type="journal article" year="1933">Mercier 1933</bibRefCitation>
). In these ergotaxonomies the ‘
<taxonomicName id="4C794F93FF85730A7BF4097C0FC0FE7C" authorityName="Bonaparte" authorityYear="1850" box="[660,762,364,387]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A7BF4097C0FC0FE7C" box="[660,762,364,387]" italics="true" pageId="53" pageNumber="600">Geosaurus</emphasis>
</taxonomicName>
morphotype’ evolved independently multiple times [three times in
<bibRefCitation id="EFE849E1FF85730A79B009BA0C63FE3D" author="Fraas E" box="[208,345,426,450]" pageId="53" pageNumber="600" pagination="1 - 72" refId="ref51930" refString="Fraas E. Die Meer-Krocodilier (Thalattosuchia) des oberen Jura unter specieller Berucksichtigung von Dacosaurus und Geosaurus. Palaeontographica 1902; 49: 1 - 72." type="journal article" year="1902">Fraas (1902)</bibRefCitation>
and twice under
<bibRefCitation id="EFE849E1FF85730A7B7709BA0F81FE3D" author="Mercier J" box="[535,699,426,450]" pageId="53" pageNumber="600" pagination="99 - 119" refId="ref56612" refString="Mercier J. Contribution a l'etude des Metriorhynchides (crocodiliens). Annales de Paleontologie 1933; 22: 99 - 119." type="journal article" year="1933">Mercier (1933)</bibRefCitation>
], and most
<taxonomicName id="4C794F93FF85730A79CE09D90CA2FE1E" box="[174,408,457,481]" pageId="53" pageNumber="600">
<emphasis id="B90DE802FF85730A79CE09D90C7DFE1E" box="[174,327,457,481]" italics="true" pageId="53" pageNumber="600">Metriorhynchus</emphasis>
species
</taxonomicName>
were more closely related to one of the
<taxonomicName id="4C794F93FF85730A79D809F90C24FDFF" authorityName="Bonaparte" authorityYear="1850" box="[184,286,489,512]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A79D809F90C24FDFF" box="[184,286,489,512]" italics="true" pageId="53" pageNumber="600">Geosaurus</emphasis>
</taxonomicName>
lineages or
<taxonomicName id="4C794F93FF85730A78C009F90F2FFDFE" box="[416,533,489,513]" class="Reptilia" family="Metriorhynchidae" genus="Dakosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A78C009F90F2FFDFE" box="[416,533,489,513]" italics="true" pageId="53" pageNumber="600">Dakosaurus</emphasis>
</taxonomicName>
than to other species assigned to the genus
<taxonomicName id="4C794F93FF85730A78050A180CC4FDDF" authorityName="von Meyer" authorityYear="1832" box="[357,510,520,544]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A78050A180CC4FDDF" box="[357,510,520,544]" italics="true" pageId="53" pageNumber="600">Metriorhynchus</emphasis>
</taxonomicName>
.
<taxonomicName id="4C794F93FF85730A7B6E0A180FB9FDDF" box="[526,643,520,544]" class="Reptilia" family="Metriorhynchidae" genus="Dakosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A7B6E0A180FB9FDDF" box="[526,643,520,544]" italics="true" pageId="53" pageNumber="600">Dakosaurus</emphasis>
</taxonomicName>
as defined by
<bibRefCitation id="EFE849E1FF85730A79F10A380C22FDC0" author="Fraas E" box="[145,280,551,575]" pageId="53" pageNumber="600" pagination="1 - 72" refId="ref51930" refString="Fraas E. Die Meer-Krocodilier (Thalattosuchia) des oberen Jura unter specieller Berucksichtigung von Dacosaurus und Geosaurus. Palaeontographica 1902; 49: 1 - 72." type="journal article" year="1902">Fraas (1902)</bibRefCitation>
is also polyphyletic, as it included
<taxonomicName id="4C794F93FF85730A7BE30A370DFBFDA1" baseAuthorityName="Hulke" baseAuthorityYear="1870" class="Reptilia" family="Metriorhynchidae" genus="Plesiosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="species" species="manselii">
<emphasis id="B90DE802FF85730A7BE30A370DFBFDA1" italics="true" pageId="53" pageNumber="600">Plesiosuchus manselii</emphasis>
</taxonomicName>
, a Late Jurassic species more closely related to a Middle Jurassic ‘
<taxonomicName id="4C794F93FF85730A79AE0A760C5DFD81" authorityName="von Meyer" authorityYear="1832" box="[206,359,614,638]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A79AE0A760C5DFD81" box="[206,359,614,638]" italics="true" pageId="53" pageNumber="600">Metriorhynchus</emphasis>
</taxonomicName>
’ species (
<taxonomicName id="4C794F93FF85730A78B30A760F82FD81" authorityName="Lydekker" authorityYear="1890" box="[467,696,614,638]" class="Reptilia" family="Metriorhynchidae" genus="Suchodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="species" species="durobrivensis">
<emphasis id="B90DE802FF85730A78B30A760F82FD81" box="[467,696,614,638]" italics="true" pageId="53" pageNumber="600">Suchodus durobrivensis</emphasis>
</taxonomicName>
) than to
<taxonomicName id="4C794F93FF85730A79F10A950C50FD62" baseAuthorityName="Plieninger" baseAuthorityYear="1846" box="[145,362,645,669]" class="Reptilia" family="Metriorhynchidae" genus="Dakosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="species" species="maximus">
<emphasis id="B90DE802FF85730A79F10A950C50FD62" box="[145,362,645,669]" italics="true" pageId="53" pageNumber="600">Dakosaurus maximus</emphasis>
</taxonomicName>
(see phylogenetic results herein, and
<bibRefCitation id="EFE849E1FF85730A79110AB50C10FD42" author="Young MT &amp; Brusatte SL &amp; Andrade MB" box="[113,298,676,701]" pageId="53" pageNumber="600" pagination="44985" refId="ref60761" refString="Young MT, Brusatte SL, Andrade MB et al. The cranial osteology and feeding ecology of the metriorhynchid crocodylomorph genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe. PLoS One 2012 a; 7: e 44985. https: // doi. org / 10.1371 / journal. pone. 0044985" type="journal article" year="2012">
Young
<emphasis id="B90DE802FF85730A79D70AB50DDAFD43" box="[183,224,676,700]" italics="true" pageId="53" pageNumber="600">et al</emphasis>
. 2012a
</bibRefCitation>
).
</paragraph>
<paragraph id="8BC63410FF85730A79ED0AD40DCFFB0F" blockId="53.[113,763,144,1264]" pageId="53" pageNumber="600">
To be clear, we are not criticizing the ergotaxonomic systematizations of earlier generations, not only was there a smaller sample set of thalattosuchian fossils, but a different systematic paradigm was in place. But systematics is a science, and over the past 100 years new data has been accumulated from across the world. Hypotheses have to be re-evaluated in the light of new evidence, and we have to ask ourselves, what is the utility of an ergotaxonomy in which the genera included are paraphyletic and/or polyphyletic? How can we discuss the biodiversity or evolutionary trends within
<taxonomicName id="4C794F93FF85730A78F10BCE0F71FC09" authorityName="sensu Young and Andrade" authorityYear="2009" box="[401,587,990,1014]" class="Reptilia" family="Metriorhynchidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="family">Metriorhynchidae</taxonomicName>
but at the same time have multiple lineages called
<taxonomicName id="4C794F93FF85730A78850BED0F71FBEB" authorityName="Bonaparte" authorityYear="1850" box="[485,587,1021,1044]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A78850BED0F71FBEB" box="[485,587,1021,1044]" italics="true" pageId="53" pageNumber="600">Geosaurus</emphasis>
</taxonomicName>
, or
<taxonomicName id="4C794F93FF85730A7B1C0BED0FCBFBEA" authority="Especially" authorityName="Especially" box="[636,753,1021,1045]" class="Reptilia" family="Metriorhynchidae" genus="Dakosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A7B1C0BED0FCBFBEA" box="[636,753,1021,1045]" italics="true" pageId="53" pageNumber="600">Dakosaurus</emphasis>
</taxonomicName>
? Especially when there are pre-existing generic names that could be ‘resurrected’ and assigned to these other lineages [as was done by
<bibRefCitation id="EFE849E1FF85730A79AB0C4C0CDDFB8C" author="Young MT &amp; Andrade MB" box="[203,487,1115,1140]" pageId="53" pageNumber="600" pagination="551 - 85" refId="ref60527" refString="Young MT, Andrade MB. What is Geosaurus? Redescription of Geosaurus giganteus (Thalattosuchia, Metriorhynchidae) from the Upper Jurassic of Bayern, Germany. Zoological Journal of the Linnean Society 2009; 157: 551 - 85." type="journal article" year="2009">Young and Andrade (2009)</bibRefCitation>
and
<bibRefCitation id="EFE849E1FF85730A7B790C4C0FD0FB8C" author="Young MT &amp; Brusatte SL &amp; Andrade MB" box="[537,746,1115,1140]" pageId="53" pageNumber="600" pagination="44985" refId="ref60761" refString="Young MT, Brusatte SL, Andrade MB et al. The cranial osteology and feeding ecology of the metriorhynchid crocodylomorph genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe. PLoS One 2012 a; 7: e 44985. https: // doi. org / 10.1371 / journal. pone. 0044985" type="journal article" year="2012">
Young
<emphasis id="B90DE802FF85730A7B000C4C0FB0FB8C" box="[608,650,1115,1139]" italics="true" pageId="53" pageNumber="600">et al</emphasis>
. (2012a)
</bibRefCitation>
]. Using
<taxonomicName id="4C794F93FF85730A79D40C6A0C77FB6D" authorityName="von Meyer" authorityYear="1832" box="[180,333,1146,1170]" class="Reptilia" family="Metriorhynchidae" genus="Metriorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A79D40C6A0C77FB6D" box="[180,333,1146,1170]" italics="true" pageId="53" pageNumber="600">Metriorhynchus</emphasis>
</taxonomicName>
as a ‘wastebasket’ for all metriorhynchids that do not fall within
<taxonomicName id="4C794F93FF85730A78350C8A0C81FB4E" authorityName="Bonaparte" authorityYear="1850" box="[341,443,1178,1201]" class="Reptilia" family="Metriorhynchidae" genus="Geosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A78350C8A0C81FB4E" box="[341,443,1178,1201]" italics="true" pageId="53" pageNumber="600">Geosaurus</emphasis>
</taxonomicName>
or
<taxonomicName id="4C794F93FF85730A78BD0C8A0F68FB4D" box="[477,594,1178,1202]" class="Reptilia" family="Metriorhynchidae" genus="Dakosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="53" pageNumber="600" phylum="Chordata" rank="genus">
<emphasis id="B90DE802FF85730A78BD0C8A0F68FB4D" box="[477,594,1178,1202]" italics="true" pageId="53" pageNumber="600">Dakosaurus</emphasis>
</taxonomicName>
simply obscures the morphological diversity present (e.g. see:
<bibRefCitation id="EFE849E1FF85730A7B240CAA0FCFFB2E" author="Young MT &amp; Brusatte SL &amp; Ruta M" box="[580,757,1209,1234]" pageId="53" pageNumber="600" pagination="801 - 59" refId="ref60618" refString="Young MT, Brusatte SL, Ruta M et al. The evolution of Metriorhynchoidea (Mesoeucrocodylia, Thalattosuchia): an integrated approach using geometrics morphometrics, analysis of disparity and biomechanics. Zoological Journal of the Linnean Society 2010; 158: 801 - 59." type="journal article" year="2010">
Young
<emphasis id="B90DE802FF85730A7BEB0CAA0F8EFB2E" box="[651,692,1209,1233]" italics="true" pageId="53" pageNumber="600">et al</emphasis>
. 2010
</bibRefCitation>
, 2013, 2021).
</paragraph>
</subSubSection>
</treatment>
</document>
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