<documentid="C9354DF97EEC6A5CF7D7478E53C8B268"ID-CLB-Dataset="55595"ID-DOI="10.1093/zoolinnean/zlab118"ID-GBIF-Dataset="e285e5e2-a1c5-4a7e-ae03-01a19f9867c2"ID-ISSN="0024-4082"ID-Zenodo-Dep="6536781"ID-ZooBank="DACBA4A5-8D5F-479B-AFA5-E38B380F8962"IM.materialsCitations_approvedBy="felipe"IM.metadata_approvedBy="felipe"IM.tables_approvedBy="jonas"IM.taxonomicNames_approvedBy="jonas"IM.treatments_approvedBy="jonas"checkinTime="1652084918445"checkinUser="plazi"docAuthor="Laboury, Antoine, Bennion, Rebecca F., Thuy, Ben, Weis, Robert & Fischer, Valentin"docDate="2022"docId="2663F827FFDFAC54121EF54EFDEFFB33"docLanguage="en"docName="zlab118.pdf"docOrigin="Zoological Journal of the Linnean Society 195"docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-2023.journal_article"docStyleId="36B3BD6A90C22AB4F7F465C853188CC8"docStyleName="ZoolJLinnSoc.2017-2023.journal_article"docStyleVersion="7"docTitle="Temnodontosaurus cf. zetlandicus"docType="treatment"docVersion="7"lastPageNumber="184"masterDocId="DA5A805FFFD6AC581164F30AFFBFFFA8"masterDocTitle="Anatomy and phylogenetic relationships of Temnodontosaurus zetlandicus (Reptilia: Ichthyosauria)"masterLastPageNumber="194"masterPageNumber="172"pageNumber="181"updateTime="1731589745639"updateUser="jonas">
<mods:titleid="3663449BAE7601B47DEA42CC5498089A">Anatomy and phylogenetic relationships of Temnodontosaurus zetlandicus (Reptilia: Ichthyosauria)</mods:title>
MNHNL TU885, a partial skull from the Schistes Carton Formation, Lower Toarcian, Schouweiler,
<collectingCountryid="D6DD09A1FFDFAC5112DAF5C3FAB2F977"box="[958,1293,1737,1759]"name="Luxembourg"pageId="9"pageNumber="181">Grand Duchy of Luxembourg</collectingCountry>
<emphasisid="9CBE9523FFDFAC51124DF5F7FAF3F8BA"box="[809,1356,1789,1811]"italics="true"pageId="9"pageNumber="181">Comparative description of MNHNL TU885:</emphasis>
long and consists of a cranium lacking the snout and a fragmentary surangular (
<figureCitationid="36F155B4FFDCAC52137FF3CFFDDBFF73"box="[539,612,197,219]"captionStart="Figure 5"captionStartId="10.[164,242,1846,1868]"captionTargetBox="[166,1446,867,1803]"captionTargetId="figure-381@10.[163,1449,864,1806]"captionTargetPageId="10"captionText="Figure 5. Photograph and interpretation of the specimen MNHNL TU885 Temnodontosaurus cf. zetlandicus A, B, in right lateral view. C, D, in dorsal view."figureDoi="http://doi.org/10.5281/zenodo.6536803"httpUri="https://zenodo.org/record/6536803/files/figure.png"pageId="10"pageNumber="182">Figs 5</figureCitation>
,
<figureCitationid="36F155B4FFDCAC521316F3CFFD3FFF73"box="[626,640,197,219]"captionStart="Figure 6"captionStartId="11.[145,223,1284,1306]"captionTargetBox="[148,1429,198,1241]"captionTargetId="figure-281@11.[145,1432,195,1244]"captionTargetPageId="11"captionText="Figure 6. Photograph and interpretation of the specimen MNHNL TU885 Temnodontosaurus cf.zetlandicus A, B, palate in ventral view. C, D, Basicranium in posterior view. Abbreviation: ECA, extracondylar area of the basioccipital."figureDoi="http://doi.org/10.5281/zenodo.6536807"httpUri="https://zenodo.org/record/6536807/files/figure.png"pageId="10"pageNumber="182">6</figureCitation>
). Due to its preservation state, the orbits and temporal fenestrae are not fully delimited and the right vomer and the left palatine have been slightly displaced (
<figureCitationid="36F155B4FFDCAC52133EF22BFD75FE9F"box="[602,714,289,311]"captionStart="Figure 6"captionStartId="11.[145,223,1284,1306]"captionTargetBox="[148,1429,198,1241]"captionTargetId="figure-281@11.[145,1432,195,1244]"captionTargetPageId="11"captionText="Figure 6. Photograph and interpretation of the specimen MNHNL TU885 Temnodontosaurus cf.zetlandicus A, B, palate in ventral view. C, D, Basicranium in posterior view. Abbreviation: ECA, extracondylar area of the basioccipital."figureDoi="http://doi.org/10.5281/zenodo.6536807"httpUri="https://zenodo.org/record/6536807/files/figure.png"pageId="10"pageNumber="182">Fig. 6A, B</figureCitation>
). The sutures between the nasals, frontals, right prefrontal and postfrontals are often difficult to discern (
<figureCitationid="36F155B4FFDCAC5213BAF254FF59FE3B"captionStart="Figure 5"captionStartId="10.[164,242,1846,1868]"captionTargetBox="[166,1446,867,1803]"captionTargetId="figure-381@10.[163,1449,864,1806]"captionTargetPageId="10"captionText="Figure 5. Photograph and interpretation of the specimen MNHNL TU885 Temnodontosaurus cf. zetlandicus A, B, in right lateral view. C, D, in dorsal view."figureDoi="http://doi.org/10.5281/zenodo.6536803"httpUri="https://zenodo.org/record/6536803/files/figure.png"pageId="10"pageNumber="182">Fig. 5C, D</figureCitation>
). The parietal foramen is entirely preserved and appears unusually large, with a markedly raised rim; it is presently unclear whether this condition is pathological or not, as the parietal foramen is not located on the cranial midline. The postorbital region is only preserved on the left side and is anteroposteriorly long (
<figureCitationid="36F155B4FFDCAC521187F13FFEEAFDE3"box="[227,341,565,587]"captionStart="Figure 5"captionStartId="10.[164,242,1846,1868]"captionTargetBox="[166,1446,867,1803]"captionTargetId="figure-381@10.[163,1449,864,1806]"captionTargetPageId="10"captionText="Figure 5. Photograph and interpretation of the specimen MNHNL TU885 Temnodontosaurus cf. zetlandicus A, B, in right lateral view. C, D, in dorsal view."figureDoi="http://doi.org/10.5281/zenodo.6536803"httpUri="https://zenodo.org/record/6536803/files/figure.png"pageId="10"pageNumber="182">Fig. 5A, B</figureCitation>
<figureCitationid="36F155B4FFDCAC5210A1F15EFD86FDC1"box="[453,569,596,618]"captionStart="Figure 6"captionStartId="11.[145,223,1284,1306]"captionTargetBox="[148,1429,198,1241]"captionTargetId="figure-281@11.[145,1432,195,1244]"captionTargetPageId="11"captionText="Figure 6. Photograph and interpretation of the specimen MNHNL TU885 Temnodontosaurus cf.zetlandicus A, B, palate in ventral view. C, D, Basicranium in posterior view. Abbreviation: ECA, extracondylar area of the basioccipital."figureDoi="http://doi.org/10.5281/zenodo.6536807"httpUri="https://zenodo.org/record/6536807/files/figure.png"pageId="10"pageNumber="182">Fig. 6C, D</figureCitation>
), only lacking the left opisthotic. Hereafter, we compare salient features of MNHNL TU885 with early neoichthyosaurians.
This specimen shares many features with early neoichthyosaurians such as a frontal without a temporal process, a supraoccipital with a shallow ventral notch, a robust stapes with a prominent proximal head, a stout opisthotic with a short paroccipital process, a basioccipital with a well-demarcated extracondylar area, which is ventrally saddle-shaped and marked by the presence of a basal tubera, and a paired carotid foramen as suggested by the morphology of the parasphenoid, even if the posterior part mostly seems to be broken (
<bibRefCitationid="CA5B34C0FFDCAC521557F296FB4FFE19"author="McGowan C"box="[1075,1264,412,433]"pageId="10"pageNumber="182"pagination="1 - 109"refId="ref14201"refString="McGowan C. 1973. The cranial morphology of the Lower Liassic latipinnate ichthyosaurs of England. Bulletin of the British Museum (Natural History) Geology 24: 1 - 109."type="journal article"year="1973">McGowan, 1973</bibRefCitation>
;
<bibRefCitationid="CA5B34C0FFDCAC52159BF296FCCCFE67"author="Fischer V & Masure E & Arkhangelsky MSMS & Godefroit P"pageId="10"pageNumber="182"pagination="1010 - 1025"refId="ref12482"refString="Fischer V, Masure E, Arkhangelsky MSMS, Godefroit P. 2011. A new Barremian (Early Cretaceous) ichthyosaur from western Russia. Journal of Vertebrate Paleontology 31: 1010 - 1025."type="journal article"year="2011">
<bibRefCitationid="CA5B34C0FFDCAC5212E4F2B0FBE6FE67"author="Martin JE & Fischer V & Vincent P & Suan G"box="[896,1113,442,464]"pageId="10"pageNumber="182"pagination="995 - 1005"refId="ref13769"refString="Martin JE, Fischer V, Vincent P, Suan G. 2012. A longirostrine Temnodontosaurus (Ichthyosauria) with comments on Early Jurassic ichthyosaur niche partitioning and disparity. Palaeontology 55: 995 - 1005."type="journal article"year="2012">
<bibRefCitationid="CA5B34C0FFDCAC521502F2B0FA85FE78"author="Marek RD & Moon BC & Williams M & Benton MJ"box="[1126,1338,442,464]"pageId="10"pageNumber="182"pagination="723 - 742"refId="ref13734"refString="Marek RD, Moon BC, Williams M, Benton MJ. 2015. The skull and endocranium of a Lower Jurassic ichthyosaur based on digital reconstructions. Palaeontology 58: 723 - 742."type="journal article"year="2015">
<bibRefCitationid="CA5B34C0FFDCAC521423F2B1FC76FE47"author="Moon BC & Kirton AM"pageId="10"pageNumber="182"pagination="1 - 84"refId="ref14562"refString="Moon BC, Kirton AM. 2016. Ichthyosaurs of the British Middle and Upper Jurassic Part 1, Ophthalmosaurus. Monographs of the Palaeontographical Society 170: 1 - 84."type="journal article"year="2016">Moon & Kirton, 2016</bibRefCitation>
;
<bibRefCitationid="CA5B34C0FFDCAC5212B1F2D3FB1BFE46"author="Lomax DR & Porro LB & Larkin NR"box="[981,1188,473,495]"pageId="10"pageNumber="182"pagination="6112"refId="ref13209"refString="Lomax DR, Porro LB, Larkin NR. 2019. Descriptive anatomy of the largest known specimen of Protoichthyosaurus prostaxalis (Reptilia: Ichthyosauria) including computed tomography and digital reconstruction of a three-dimensional skull. PeerJ 7: e 6112."type="journal article"year="2019">
<bibRefCitationid="CA5B34C0FFDCAC5215D5F2D3FCCBFDA5"author="Miedema F & Maxwell EE"pageId="10"pageNumber="182"pagination="1675164"refId="ref14442"refString="Miedema F, Maxwell EE. 2019. Ontogeny of the braincase in Stenopterygius (Reptilia, Ichthyosauria) from the Lower Jurassic of Germany. Journal of Vertebrate Paleontology 39: e 1675164."type="journal article"year="2019">Miedema & Maxwell, 2019</bibRefCitation>
). In posterior view (
<figureCitationid="36F155B4FFDCAC521515F2F2FB54FDA5"box="[1137,1259,504,526]"captionStart="Figure 6"captionStartId="11.[145,223,1284,1306]"captionTargetBox="[148,1429,198,1241]"captionTargetId="figure-281@11.[145,1432,195,1244]"captionTargetPageId="11"captionText="Figure 6. Photograph and interpretation of the specimen MNHNL TU885 Temnodontosaurus cf.zetlandicus A, B, palate in ventral view. C, D, Basicranium in posterior view. Abbreviation: ECA, extracondylar area of the basioccipital."figureDoi="http://doi.org/10.5281/zenodo.6536807"httpUri="https://zenodo.org/record/6536807/files/figure.png"pageId="10"pageNumber="182">Fig. 6C, D</figureCitation>
), the quadrate is a robust and large element that has a straight occipital lamella unlike
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<taxonomicNameid="69CA32B2FFDCAC5212DAF15EFA1DFDC1"authority="(Swaby & Lomax, 2020)"baseAuthorityName="Swaby & Lomax"baseAuthorityYear="2020"box="[958,1442,596,618]"class="Reptilia"family="Temnodontosauridae"genus="Temnodontosaurus"higherTaxonomySource="GBIF"kingdom="Animalia"order="Ichthyosauria"pageId="10"pageNumber="182"phylum="Chordata"rank="species"species="crassimanus">
<bibRefCitationid="CA5B34C0FFDCAC5215ECF15EFA26FDC1"author="Swaby EJ & Lomax DR"box="[1160,1433,596,618]"pageId="10"pageNumber="182"pagination="2715 - 2731"refId="ref15048"refString="Swaby EJ, Lomax DR. 2020. A revision of Temnodontosaurus crassimanus (Reptilia: Ichthyosauria) from the Lower Jurassic (Toarcian) of Whitby, Yorkshire, UK. Historical Biology 33: 2715 - 2731."type="journal article"year="2020">Swaby & Lomax, 2020</bibRefCitation>
)
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and with a massive and well-developed condyle as in species currently referred to as
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<taxonomicNameid="69CA32B2FFDCAC5212B9F1C5FA21FD4C"authority="(Martin et al., 2012)"baseAuthorityName="Martin"baseAuthorityYear="2012"box="[989,1438,718,740]"class="Reptilia"family="Temnodontosauridae"genus="Temnodontosaurus"higherTaxonomySource="GBIF"kingdom="Animalia"order="Ichthyosauria"pageId="10"pageNumber="182"phylum="Chordata"rank="species"species="azerguensis">
<bibRefCitationid="CA5B34C0FFDCAC5215C7F1C4FA2CFD4B"author="Martin JE & Fischer V & Vincent P & Suan G"box="[1187,1427,718,740]"pageId="10"pageNumber="182"pagination="995 - 1005"refId="ref13769"refString="Martin JE, Fischer V, Vincent P, Suan G. 2012. A longirostrine Temnodontosaurus (Ichthyosauria) with comments on Early Jurassic ichthyosaur niche partitioning and disparity. Palaeontology 55: 995 - 1005."type="journal article"year="2012">
<bibRefCitationid="CA5B34C0FFDDAC5311ABF6BAFE38FA6D"author="McGowan C"box="[207,391,1456,1477]"pageId="11"pageNumber="183"pagination="1 - 37"refId="ref14234"refString="McGowan C. 1974. A revision of the longipinnate ichthyosaurs of the Lower Jurassic of England, with description of the new species (Reptilia, Ichthyosauria). Life Science Contributions, Royal Ontario Museum 97: 1 - 37."type="journal article"year="1974">McGowan, 1974</bibRefCitation>
,
<bibRefCitationid="CA5B34C0FFDDAC5310F1F6BAFE72FA6D"author="McGowan C"box="[405,461,1456,1477]"pageId="11"pageNumber="183"pagination="472 - 479"refId="ref14301"refString="McGowan C. 1994. Temnodontosaurus risor is a juvenile of T. platyodon (Reptilia: Ichthyosauria). Journal of Vertebrate Paleontology 14: 472 - 479."type="journal article"year="1994">1994</bibRefCitation>
;
<bibRefCitationid="CA5B34C0FFDDAC5310BEF6A5FF77FA4B"author="Maisch MW & Hungerbuhler A"pageId="11"pageNumber="183"pagination="335 - 345"refId="ref13598"refString="Maisch MW, Hungerbuhler A. 2001. New evidence for a discrete supratemporal bone in the Jurassic Ichthyosaur Temnodontosaurus. Historical Biology 15: 335 - 345."type="journal article"year="2001">Maisch & Hungerbühler, 2001</bibRefCitation>
;
<bibRefCitationid="CA5B34C0FFDDAC5311B0F6C4FE6FFA4B"author="Swaby EJ & Lomax DR"box="[212,464,1486,1508]"pageId="11"pageNumber="183"pagination="2715 - 2731"refId="ref15048"refString="Swaby EJ, Lomax DR. 2020. A revision of Temnodontosaurus crassimanus (Reptilia: Ichthyosauria) from the Lower Jurassic (Toarcian) of Whitby, Yorkshire, UK. Historical Biology 33: 2715 - 2731."type="journal article"year="2020">Swaby & Lomax, 2020</bibRefCitation>
) but unlike in other Early Jurassic parvipelvians (Maisch, 2001, 2008;
<bibRefCitationid="CA5B34C0FFDDAC5313C0F6E7FEF8F989"author="Maisch MW & Matzke AT"pageId="11"pageNumber="183"pagination="116 - 127"refId="ref13701"refString="Maisch MW, Matzke AT. 2003. The cranial osteology of the ichthyosaur Leptonectes tenuirostris from the Lower Jurassic of England. Journal of Vertebrate Paleontology 23: 116 - 127."type="journal article"year="2003">Maisch & Matzke, 2003</bibRefCitation>
;
<bibRefCitationid="CA5B34C0FFDDAC531031F506FDD2F989"author="Massare JA & Lomax DR"box="[341,621,1547,1569]"pageId="11"pageNumber="183"pagination="5"refId="ref13889"refString="Massare JA, Lomax DR. 2016. A new specimen of Ichthyosaurus conybeari (Reptilia, Ichthyosauria) from Watchet, Somerset, England, U. K., and a re-examination of the species. Journal of Vertebrate Paleontology 36: 5."type="journal article"year="2016">Lomax & Massare, 2016</bibRefCitation>
;
<bibRefCitationid="CA5B34C0FFDDAC53131FF501FEA1F997"author="Maxwell EE & Cortes D"pageId="11"pageNumber="183"pagination="31"refId="ref14039"refString="Maxwell EE, Cortes D. 2020. A revision of the Early Jurassic ichthyosaur Hauffiopteryx (Reptilia: Ichthyosauria), and description of a new species from southwestern Germany. Palaeontologia Electronica 23: a 31."type="journal article"year="2020">Maxwell & Cortés, 2020</bibRefCitation>
). The contribution of the supraoccipital to the dorsal margin of the foramen magnum is limited and nearly absent, as in
<taxonomicNameid="69CA32B2FFDDAC531033F5CEFDB7F970"authorityName="Laboury & Bennion & Thuy & Weis & Fischer"authorityYear="2022"baseAuthorityName="Laboury & Bennion & Thuy & Weis & Fischer"baseAuthorityYear="2022"box="[343,520,1731,1753]"class="Reptilia"family="Temnodontosauridae"genus="Temnodontosaurus"higherTaxonomySource="GBIF"kingdom="Animalia"order="Ichthyosauria"pageId="11"pageNumber="183"phylum="Chordata"rank="species"species="zetlandicus">
<bibRefCitationid="CA5B34C0FFDDAC5311FDF5E8FE82F950"author="Maisch MW"box="[153,317,1762,1784]"pageId="11"pageNumber="183"pagination="401 - 431"refId="ref13373"refString="Maisch MW. 1998 b. Short review of the ichthyosaurs of the Posidonienschiefer with remarks on the taxonomy of the Stenopterygiidae and Temnodontosauridae. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 209: 401 - 431."type="journal article"year="1998">Maisch, 1998b</bibRefCitation>
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<taxonomicNameid="69CA32B2FFDDAC531223F69BFC54FA0E"authorityName="Laboury & Bennion & Thuy & Weis & Fischer"authorityYear="2022"baseAuthorityName="Laboury & Bennion & Thuy & Weis & Fischer"baseAuthorityYear="2022"box="[839,1003,1425,1446]"class="Reptilia"family="Temnodontosauridae"genus="Temnodontosaurus"higherTaxonomySource="GBIF"kingdom="Animalia"order="Ichthyosauria"pageId="11"pageNumber="183"phylum="Chordata"rank="species"species="zetlandicus">
<figureCitationid="36F155B4FFDDAC531299F69BFBFFFA0E"box="[1021,1088,1425,1447]"captionStart="Figure 7"captionStartId="12.[163,241,1052,1074]"captionTargetBox="[166,1446,198,1009]"captionTargetId="figure-457@12.[163,1449,195,1012]"captionTargetPageId="12"captionText="Figure 7. Braincase comparison of the holotype of Temnodontosaurus zetlandicus comb. nov. (CAMSM J35176) (A) and the referred specimen of Temnodontosaurus cf. zetlandicus (MNHNL TU885) (B)."figureDoi="http://doi.org/10.5281/zenodo.6536812"httpUri="https://zenodo.org/record/6536812/files/figure.png"pageId="11"pageNumber="183">Fig. 7</figureCitation>
), such as: (1) the bifurcated posterior processes of the nasal which partially covers the postfrontal which anteriorly appears in the medial and anterolateral processes; (2) the presence of a prominent mediolaterally oriented ridge on the postfrontal which forms the anterior margin of the supratemporal fenestra; and (3) a lanceolate frontal posteriorly covered by a slender forked process of the parietal. The skull roof of MNHNL TU885 also resembles that of
<bibRefCitationid="CA5B34C0FFDDAC53150EF5E8FAACF950"author="Maisch MW"box="[1130,1299,1762,1784]"pageId="11"pageNumber="183"pagination="401 - 431"refId="ref13373"refString="Maisch MW. 1998 b. Short review of the ichthyosaurs of the Posidonienschiefer with remarks on the taxonomy of the Stenopterygiidae and Temnodontosauridae. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 209: 401 - 431."type="journal article"year="1998">Maisch, 1998b</bibRefCitation>
)
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. MNHNL TU885 is also similar in size and in cranial proportions to
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<taxonomicNameid="69CA32B2FFDDAC53124DF435FC5AF8FB"authorityName="Laboury & Bennion & Thuy & Weis & Fischer"authorityYear="2022"baseAuthorityName="Laboury & Bennion & Thuy & Weis & Fischer"baseAuthorityYear="2022"box="[809,997,1854,1876]"class="Reptilia"family="Temnodontosauridae"genus="Temnodontosaurus"higherTaxonomySource="GBIF"isUncertain="true"kingdom="Animalia"order="Ichthyosauria"pageId="11"pageNumber="183"phylum="Chordata"rank="species"species="zetlandicus">
<taxonomicNameid="69CA32B2FFDAAC5413DCF716FC44FB9A"authorityName="Laboury & Bennion & Thuy & Weis & Fischer"authorityYear="2022"baseAuthorityName="Laboury & Bennion & Thuy & Weis & Fischer"baseAuthorityYear="2022"box="[696,1019,1052,1074]"class="Reptilia"family="Temnodontosauridae"genus="Temnodontosaurus"higherTaxonomySource="GBIF"kingdom="Animalia"order="Ichthyosauria"pageId="12"pageNumber="184"phylum="Chordata"rank="species"species="zetlandicus"status="comb. nov.">
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