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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Revision of the Late Jurassic teleosaurid genus <italic>Machimosaurus</italic> (Crocodylomorpha, Thalattosuchia)</title><author><name sortKey="Young, Mark T" sort="Young, Mark T" uniqKey="Young M" first="Mark T." last="Young">Mark T. Young</name><affiliation><nlm:aff id="aff1"><addr-line>School of GeoSciences</addr-line>,<institution>University of Edinburgh, The King's Buildings</institution>,<addr-line>Edinburgh EH9 3JW, UK</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><addr-line>School of Ocean and Earth Science, National Oceanography Centre</addr-line>,<institution>University of Southampton</institution>,<addr-line>Southampton SO14 3ZH, UK</addr-line></nlm:aff></affiliation></author><author><name sortKey="Hua, Stephane" sort="Hua, Stephane" uniqKey="Hua S" first="Stéphane" last="Hua">Stéphane Hua</name><affiliation><nlm:aff id="aff3"><addr-line>Le Musée des Dinosaures d'Espéraza</addr-line>,<addr-line>Espéraza 11260, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Steel, Lorna" sort="Steel, Lorna" uniqKey="Steel L" first="Lorna" last="Steel">Lorna Steel</name><affiliation><nlm:aff id="aff4"><addr-line>Department of Earth Sciences</addr-line>,<institution>Natural History Museum</institution>,<addr-line>London SW7 5BD, UK</addr-line></nlm:aff></affiliation></author><author><name sortKey="Foffa, Davide" sort="Foffa, Davide" uniqKey="Foffa D" first="Davide" last="Foffa">Davide Foffa</name><affiliation><nlm:aff id="aff1"><addr-line>School of GeoSciences</addr-line>,<institution>University of Edinburgh, The King's Buildings</institution>,<addr-line>Edinburgh EH9 3JW, UK</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff5"><addr-line>School of Earth Sciences, Wills Memorial Building</addr-line>,<institution>University of Bristol</institution>,<addr-line>Bristol BS8 1RJ, UK</addr-line></nlm:aff></affiliation></author><author><name sortKey="Brusatte, Stephen L" sort="Brusatte, Stephen L" uniqKey="Brusatte S" first="Stephen L." last="Brusatte">Stephen L. Brusatte</name><affiliation><nlm:aff id="aff1"><addr-line>School of GeoSciences</addr-line>,<institution>University of Edinburgh, The King's Buildings</institution>,<addr-line>Edinburgh EH9 3JW, UK</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff6"><addr-line>National Museums Scotland</addr-line>,<institution>Chambers Street, Edinburgh</institution>,<addr-line>EH1 1JF, UK</addr-line></nlm:aff></affiliation></author><author><name sortKey="Thuring, Silvan" sort="Thuring, Silvan" uniqKey="Thuring S" first="Silvan" last="Thüring">Silvan Thüring</name><affiliation><nlm:aff id="aff7"><addr-line>Naturmuseum Solothurn</addr-line>,<institution>Klosterplatz 2</institution>,<addr-line>Solothurn 4500, Switzerland</addr-line></nlm:aff></affiliation></author><author><name sortKey="Mateus, Octavio" sort="Mateus, Octavio" uniqKey="Mateus O" first="Octávio" last="Mateus">Octávio Mateus</name><affiliation><nlm:aff id="aff8"><addr-line>Departamento de Ciências da Terra, Faculdade de Ciências e Tecnologia</addr-line>,<institution>Universidade Nova de Lisboa, GeoBioTec, Quinta da Torre</institution>,<addr-line>Caparica 2829-516, Portugal</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff9"><addr-line>Museu da Lourinhã</addr-line>,<institution>Rua João Luis de Moura 9</institution>,<addr-line>Lourinhã 2530-158, Portugal</addr-line></nlm:aff></affiliation></author><author><name sortKey="Ruiz Ome Aca, Jose Ignacio" sort="Ruiz Ome Aca, Jose Ignacio" uniqKey="Ruiz Ome Aca J" first="José Ignacio" last="Ruiz-Ome Aca">José Ignacio Ruiz-Ome Aca</name><affiliation><nlm:aff id="aff10"><addr-line>Museo del Jurásico de Asturias (MUJA)</addr-line>,<addr-line>Colunga 33328, Spain</addr-line></nlm:aff></affiliation></author><author><name sortKey="Havlik, Philipe" sort="Havlik, Philipe" uniqKey="Havlik P" first="Philipe" last="Havlik">Philipe Havlik</name><affiliation><nlm:aff id="aff11"><addr-line>Paläontologische Sammlung, Senckenberg Center for Human Evolution and Palaeoenvironment</addr-line>,<institution>Eberhard Karls Universität, Sigwartstrasse 10</institution>,<addr-line>Tübingen 72072, Germany</addr-line></nlm:aff></affiliation></author><author><name sortKey="Lepage, Yves" sort="Lepage, Yves" uniqKey="Lepage Y" first="Yves" last="Lepage">Yves Lepage</name><affiliation><nlm:aff id="aff12"><addr-line>Sciences et Géologie Normandes</addr-line>,<addr-line>Le Havre 76620, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="De Andrade, Marco Brandalise" sort="De Andrade, Marco Brandalise" uniqKey="De Andrade M" first="Marco Brandalise" last="De Andrade">Marco Brandalise De Andrade</name><affiliation><nlm:aff id="aff13"><addr-line>Faculdade de Biociências</addr-line>,<institution>Pontifícia Universidade Católica do Rio Grande do Sul, Avenida Ipiranga 6681</institution>,<addr-line>Porto Alegre 90619-900, Brazil</addr-line></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">26064545</idno><idno type="pmc">4448888</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4448888</idno><idno type="RBID">PMC:4448888</idno><idno type="doi">10.1098/rsos.140222</idno><date when="2014">2014</date><idno type="wicri:Area/Pmc/Corpus">000014</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Revision of the Late Jurassic teleosaurid genus <italic>Machimosaurus</italic> (Crocodylomorpha, Thalattosuchia)</title><author><name sortKey="Young, Mark T" sort="Young, Mark T" uniqKey="Young M" first="Mark T." last="Young">Mark T. Young</name><affiliation><nlm:aff id="aff1"><addr-line>School of GeoSciences</addr-line>,<institution>University of Edinburgh, The King's Buildings</institution>,<addr-line>Edinburgh EH9 3JW, UK</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><addr-line>School of Ocean and Earth Science, National Oceanography Centre</addr-line>,<institution>University of Southampton</institution>,<addr-line>Southampton SO14 3ZH, UK</addr-line></nlm:aff></affiliation></author><author><name sortKey="Hua, Stephane" sort="Hua, Stephane" uniqKey="Hua S" first="Stéphane" last="Hua">Stéphane Hua</name><affiliation><nlm:aff id="aff3"><addr-line>Le Musée des Dinosaures d'Espéraza</addr-line>,<addr-line>Espéraza 11260, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="Steel, Lorna" sort="Steel, Lorna" uniqKey="Steel L" first="Lorna" last="Steel">Lorna Steel</name><affiliation><nlm:aff id="aff4"><addr-line>Department of Earth Sciences</addr-line>,<institution>Natural History Museum</institution>,<addr-line>London SW7 5BD, UK</addr-line></nlm:aff></affiliation></author><author><name sortKey="Foffa, Davide" sort="Foffa, Davide" uniqKey="Foffa D" first="Davide" last="Foffa">Davide Foffa</name><affiliation><nlm:aff id="aff1"><addr-line>School of GeoSciences</addr-line>,<institution>University of Edinburgh, The King's Buildings</institution>,<addr-line>Edinburgh EH9 3JW, UK</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff5"><addr-line>School of Earth Sciences, Wills Memorial Building</addr-line>,<institution>University of Bristol</institution>,<addr-line>Bristol BS8 1RJ, UK</addr-line></nlm:aff></affiliation></author><author><name sortKey="Brusatte, Stephen L" sort="Brusatte, Stephen L" uniqKey="Brusatte S" first="Stephen L." last="Brusatte">Stephen L. Brusatte</name><affiliation><nlm:aff id="aff1"><addr-line>School of GeoSciences</addr-line>,<institution>University of Edinburgh, The King's Buildings</institution>,<addr-line>Edinburgh EH9 3JW, UK</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff6"><addr-line>National Museums Scotland</addr-line>,<institution>Chambers Street, Edinburgh</institution>,<addr-line>EH1 1JF, UK</addr-line></nlm:aff></affiliation></author><author><name sortKey="Thuring, Silvan" sort="Thuring, Silvan" uniqKey="Thuring S" first="Silvan" last="Thüring">Silvan Thüring</name><affiliation><nlm:aff id="aff7"><addr-line>Naturmuseum Solothurn</addr-line>,<institution>Klosterplatz 2</institution>,<addr-line>Solothurn 4500, Switzerland</addr-line></nlm:aff></affiliation></author><author><name sortKey="Mateus, Octavio" sort="Mateus, Octavio" uniqKey="Mateus O" first="Octávio" last="Mateus">Octávio Mateus</name><affiliation><nlm:aff id="aff8"><addr-line>Departamento de Ciências da Terra, Faculdade de Ciências e Tecnologia</addr-line>,<institution>Universidade Nova de Lisboa, GeoBioTec, Quinta da Torre</institution>,<addr-line>Caparica 2829-516, Portugal</addr-line></nlm:aff></affiliation><affiliation><nlm:aff id="aff9"><addr-line>Museu da Lourinhã</addr-line>,<institution>Rua João Luis de Moura 9</institution>,<addr-line>Lourinhã 2530-158, Portugal</addr-line></nlm:aff></affiliation></author><author><name sortKey="Ruiz Ome Aca, Jose Ignacio" sort="Ruiz Ome Aca, Jose Ignacio" uniqKey="Ruiz Ome Aca J" first="José Ignacio" last="Ruiz-Ome Aca">José Ignacio Ruiz-Ome Aca</name><affiliation><nlm:aff id="aff10"><addr-line>Museo del Jurásico de Asturias (MUJA)</addr-line>,<addr-line>Colunga 33328, Spain</addr-line></nlm:aff></affiliation></author><author><name sortKey="Havlik, Philipe" sort="Havlik, Philipe" uniqKey="Havlik P" first="Philipe" last="Havlik">Philipe Havlik</name><affiliation><nlm:aff id="aff11"><addr-line>Paläontologische Sammlung, Senckenberg Center for Human Evolution and Palaeoenvironment</addr-line>,<institution>Eberhard Karls Universität, Sigwartstrasse 10</institution>,<addr-line>Tübingen 72072, Germany</addr-line></nlm:aff></affiliation></author><author><name sortKey="Lepage, Yves" sort="Lepage, Yves" uniqKey="Lepage Y" first="Yves" last="Lepage">Yves Lepage</name><affiliation><nlm:aff id="aff12"><addr-line>Sciences et Géologie Normandes</addr-line>,<addr-line>Le Havre 76620, France</addr-line></nlm:aff></affiliation></author><author><name sortKey="De Andrade, Marco Brandalise" sort="De Andrade, Marco Brandalise" uniqKey="De Andrade M" first="Marco Brandalise" last="De Andrade">Marco Brandalise De Andrade</name><affiliation><nlm:aff id="aff13"><addr-line>Faculdade de Biociências</addr-line>,<institution>Pontifícia Universidade Católica do Rio Grande do Sul, Avenida Ipiranga 6681</institution>,<addr-line>Porto Alegre 90619-900, Brazil</addr-line></nlm:aff></affiliation></author></analytic><series><title level="j">Royal Society Open Science</title><idno type="eISSN">2054-5703</idno><imprint><date when="2014">2014</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p><italic>Machimosaurus</italic> was a large-bodied genus of teleosaurid crocodylomorph, considered to have been durophagous/chelonivorous, and which frequented coastal marine/estuarine ecosystems during the Late Jurassic. Here, we revise the genus based on previously described specimens and revise the species within this genus. We conclude that there were three European <italic>Machimosaurus</italic> species and another taxon in Ethiopia. This conclusion is based on numerous lines of evidence: craniomandibular, dental and postcranial morphologies; differences in estimated total body length; geological age; geographical distribution; and hypothetical lifestyle. We re-diagnose the type species <italic>Machimosaurus hugii</italic> and limit referred specimens to only those from Upper Kimmeridgian–Lower Tithonian of Switzerland, Portugal and Spain. We also re-diagnose <italic>Machimosaurus mosae</italic>, demonstrate that it is an available name and restrict the species to the uppermost Kimmeridgian–lowermost Tithonian of northeastern France. We re-diagnose and validate the species <italic>Machimosaurus nowackianus</italic> from Harrar, Ethiopia. Finally, we establish a new species, <italic>Machimosaurus buffetauti</italic>, for the Lower Kimmeridgian specimens of France and Germany (and possibly England and Poland). We hypothesize that <italic>Machimosaurus</italic> may have been analogous to the Pliocene–Holocene genus <italic>Crocodylus</italic> in having one large-bodied taxon suited to traversing marine barriers and additional, geographically limited taxa across its range.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Andrews, Cw" uniqKey="Andrews C">CW Andrews</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Buffetaut, E" uniqKey="Buffetaut E">E Buffetaut</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Buffetaut, E" uniqKey="Buffetaut E">E Buffetaut</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hua, S" uniqKey="Hua S">S Hua</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Young, Mt" uniqKey="Young M">MT Young</name></author><author><name sortKey="Steel, L" uniqKey="Steel L">L 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<front><journal-meta><journal-id journal-id-type="nlm-ta">R Soc Open Sci</journal-id><journal-id journal-id-type="iso-abbrev">R Soc Open Sci</journal-id><journal-id journal-id-type="publisher-id">RSOS</journal-id><journal-id journal-id-type="hwp">royopensci</journal-id><journal-title-group><journal-title>Royal Society Open Science</journal-title></journal-title-group><issn pub-type="epub">2054-5703</issn><publisher><publisher-name>The Royal Society Publishing</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">26064545</article-id><article-id pub-id-type="pmc">4448888</article-id><article-id pub-id-type="doi">10.1098/rsos.140222</article-id><article-id pub-id-type="publisher-id">rsos140222</article-id><article-categories><subj-group subj-group-type="hwp-journal-coll"><subject>183</subject><subject>1001</subject><subject>70</subject><subject>144</subject></subj-group><subj-group subj-group-type="heading"><subject>Research Articles</subject></subj-group></article-categories><title-group><article-title>Revision of the Late Jurassic teleosaurid genus <italic>Machimosaurus</italic> (Crocodylomorpha, Thalattosuchia)</article-title><alt-title alt-title-type="short">Revision of the genus Machimosaurus</alt-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Young</surname><given-names>Mark T.</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="aff" rid="aff2">2</xref><xref ref-type="corresp" rid="cor1"></xref></contrib><contrib contrib-type="author"><name><surname>Hua</surname><given-names>Stéphane</given-names></name><xref ref-type="aff" rid="aff3">3</xref></contrib><contrib contrib-type="author"><name><surname>Steel</surname><given-names>Lorna</given-names></name><xref ref-type="aff" rid="aff4">4</xref></contrib><contrib contrib-type="author"><name><surname>Foffa</surname><given-names>Davide</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="aff" rid="aff5">5</xref></contrib><contrib contrib-type="author"><name><surname>Brusatte</surname><given-names>Stephen L.</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="aff" rid="aff6">6</xref></contrib><contrib contrib-type="author"><name><surname>Thüring</surname><given-names>Silvan</given-names></name><xref ref-type="aff" rid="aff7">7</xref></contrib><contrib contrib-type="author"><name><surname>Mateus</surname><given-names>Octávio</given-names></name><xref ref-type="aff" rid="aff8">8</xref><xref ref-type="aff" rid="aff9">9</xref></contrib><contrib contrib-type="author"><name><surname>Ruiz-Omeñaca</surname><given-names>José Ignacio</given-names></name><xref ref-type="aff" rid="aff10">10</xref></contrib><contrib contrib-type="author"><name><surname>Havlik</surname><given-names>Philipe</given-names></name><xref ref-type="aff" rid="aff11">11</xref></contrib><contrib contrib-type="author"><name><surname>Lepage</surname><given-names>Yves</given-names></name><xref ref-type="aff" rid="aff12">12</xref></contrib><contrib contrib-type="author"><name><surname>De Andrade</surname><given-names>Marco Brandalise</given-names></name><xref ref-type="aff" rid="aff13">13</xref></contrib></contrib-group><aff id="aff1"><label>1</label><addr-line>School of GeoSciences</addr-line>,<institution>University of Edinburgh, The King's Buildings</institution>,<addr-line>Edinburgh EH9 3JW, UK</addr-line></aff><aff id="aff2"><label>2</label><addr-line>School of Ocean and Earth Science, National Oceanography Centre</addr-line>,<institution>University of Southampton</institution>,<addr-line>Southampton SO14 3ZH, UK</addr-line></aff><aff id="aff3"><label>3</label><addr-line>Le Musée des Dinosaures d'Espéraza</addr-line>,<addr-line>Espéraza 11260, France</addr-line></aff><aff id="aff4"><label>4</label><addr-line>Department of Earth Sciences</addr-line>,<institution>Natural History Museum</institution>,<addr-line>London SW7 5BD, UK</addr-line></aff><aff id="aff5"><label>5</label><addr-line>School of Earth Sciences, Wills Memorial Building</addr-line>,<institution>University of Bristol</institution>,<addr-line>Bristol BS8 1RJ, UK</addr-line></aff><aff id="aff6"><label>6</label><addr-line>National Museums Scotland</addr-line>,<institution>Chambers Street, Edinburgh</institution>,<addr-line>EH1 1JF, UK</addr-line></aff><aff id="aff7"><label>7</label><addr-line>Naturmuseum Solothurn</addr-line>,<institution>Klosterplatz 2</institution>,<addr-line>Solothurn 4500, Switzerland</addr-line></aff><aff id="aff8"><label>8</label><addr-line>Departamento de Ciências da Terra, Faculdade de Ciências e Tecnologia</addr-line>,<institution>Universidade Nova de Lisboa, GeoBioTec, Quinta da Torre</institution>,<addr-line>Caparica 2829-516, Portugal</addr-line></aff><aff id="aff9"><label>9</label><addr-line>Museu da Lourinhã</addr-line>,<institution>Rua João Luis de Moura 9</institution>,<addr-line>Lourinhã 2530-158, Portugal</addr-line></aff><aff id="aff10"><label>10</label><addr-line>Museo del Jurásico de Asturias (MUJA)</addr-line>,<addr-line>Colunga 33328, Spain</addr-line></aff><aff id="aff11"><label>11</label><addr-line>Paläontologische Sammlung, Senckenberg Center for Human Evolution and Palaeoenvironment</addr-line>,<institution>Eberhard Karls Universität, Sigwartstrasse 10</institution>,<addr-line>Tübingen 72072, Germany</addr-line></aff><aff id="aff12"><label>12</label><addr-line>Sciences et Géologie Normandes</addr-line>,<addr-line>Le Havre 76620, France</addr-line></aff><aff id="aff13"><label>13</label><addr-line>Faculdade de Biociências</addr-line>,<institution>Pontifícia Universidade Católica do Rio Grande do Sul, Avenida Ipiranga 6681</institution>,<addr-line>Porto Alegre 90619-900, Brazil</addr-line></aff><author-notes><corresp id="cor1">Author for correspondence: Mark T. Young e-mail: <email>zoologika@gmail.com</email></corresp></author-notes><pub-date pub-type="collection"><month>10</month><year>2014</year></pub-date><pub-date pub-type="epub"><day>15</day><month>10</month><year>2014</year></pub-date><pub-date pub-type="pmc-release"><day>15</day><month>10</month><year>2014</year></pub-date><pmc-comment> PMC Release delay is 0 months and 0 days and was based on the
<volume>1</volume><issue>2</issue><elocation-id>140222</elocation-id><history><date date-type="received"><day>13</day><month>8</month><year>2014</year></date><date date-type="accepted"><day>16</day><month>9</month><year>2014</year></date></history><permissions><copyright-statement>© 2014 The Authors.</copyright-statement><copyright-year>2014</copyright-year><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/4.0/"><license-p>Published by the Royal Society under the terms of the Creative Commons Attribution License <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">http://creativecommons.org/licenses/by/4.0/</ext-link>, which permits unrestricted use, provided the original author and source are credited.</license-p></license></permissions><self-uri content-type="pdf" xlink:href="rsos140222.pdf"></self-uri><self-uri content-type="reviewer_comments" xlink:href="rsos140222_review_history.pdf"></self-uri><abstract><p><italic>Machimosaurus</italic> was a large-bodied genus of teleosaurid crocodylomorph, considered to have been durophagous/chelonivorous, and which frequented coastal marine/estuarine ecosystems during the Late Jurassic. Here, we revise the genus based on previously described specimens and revise the species within this genus. We conclude that there were three European <italic>Machimosaurus</italic> species and another taxon in Ethiopia. This conclusion is based on numerous lines of evidence: craniomandibular, dental and postcranial morphologies; differences in estimated total body length; geological age; geographical distribution; and hypothetical lifestyle. We re-diagnose the type species <italic>Machimosaurus hugii</italic> and limit referred specimens to only those from Upper Kimmeridgian–Lower Tithonian of Switzerland, Portugal and Spain. We also re-diagnose <italic>Machimosaurus mosae</italic>, demonstrate that it is an available name and restrict the species to the uppermost Kimmeridgian–lowermost Tithonian of northeastern France. We re-diagnose and validate the species <italic>Machimosaurus nowackianus</italic> from Harrar, Ethiopia. Finally, we establish a new species, <italic>Machimosaurus buffetauti</italic>, for the Lower Kimmeridgian specimens of France and Germany (and possibly England and Poland). We hypothesize that <italic>Machimosaurus</italic> may have been analogous to the Pliocene–Holocene genus <italic>Crocodylus</italic> in having one large-bodied taxon suited to traversing marine barriers and additional, geographically limited taxa across its range.</p></abstract><kwd-group><kwd>Africa</kwd><kwd>Europe</kwd><kwd>Kimmeridgian</kwd><kwd><italic>Machimosaurus</italic></kwd><kwd>Teleosauridae</kwd><kwd>Tithonian</kwd></kwd-group><custom-meta-group><custom-meta><meta-name>cover-date</meta-name><meta-value>October, 2014</meta-value></custom-meta></custom-meta-group></article-meta></front><body><sec sec-type="intro" id="s1"><label>2.</label><title>Introduction</title><p>Teleosaurids were a successful and diverse group of marine crocodylomorphs that lived during the Jurassic. Most teleosaurids are often considered to be marine analogues to extant gavials, due to their elongate, tubular, polydont snout, presumed primarily piscivorous diet and dorsally directed orbits [<xref rid="RSOS140222C1" ref-type="bibr">1</xref>–<xref rid="RSOS140222C5" ref-type="bibr">5</xref>]. However, there is great confusion surrounding the taxonomy of one of the most characteristic teleosaurid genera: <italic>Machimosaurus</italic>. This genus is often considered to be durophagous/chelonivorous due to a suite of craniodental morphologies that would have been well suited for feeding on hard-shelled turtles or thick-scaled fish: i.e. a foreshortened snout, proportionally enlarged supratemporal fenestrae and blunt, heavily ornamented dentition [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>,<xref rid="RSOS140222C5" ref-type="bibr">5</xref>–<xref rid="RSOS140222C8" ref-type="bibr">8</xref>]. As such, it is one of the more unusual crocodylomorphs of the Jurassic.</p><p>Two recent papers have hypothesized some unusual subjective species synonymies for the type species <italic>Machimosaurus hugii</italic> and made confusing statements about the type specimen of this species [<xref rid="RSOS140222C9" ref-type="bibr">9</xref>,<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], while one has questioned the availability of a second species, <italic>Machimosaurus mosae</italic>, as a taxonomic name [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>]. This is the impetus for this study. Here, we undertake a systematic revision of <italic>Machimosaurus</italic> and demonstrate that there were three species in the Kimmeridgian–Tithonian of Europe and a fourth species in Ethiopia. The third European species is a new taxon we name herein for the Lower Kimmeridgian specimens from France and Germany. The three European species were non-sympatric and differed in craniomandibular, dental and postcranial morphologies, total body length, geological age, geographical distribution and hypothetical lifestyle. We also address the issues surrounding the type specimens of these species and demonstrate that <italic>M. mosae</italic> is indeed an available name.</p><sec id="s1a"><label>2.1</label><title>Institutional abbreviations</title><p>BHN2R, Muséum d'Histoire Naturelle de Boulogne-sur-Mer, France (closed in 2003); DFMMh, Dinosaurier-Freilichtmuseum Münchehagen, Lower Saxony, Germany; GPIT, Paläontologische Sammlung der Eberhard Karls Universität Tübingen, Germany; MCNV, Museo de Ciencias Naturales de Valencia, Spain; MG, Museu Geológico, Lisbon, Portugal; ML, Museu da Lourinhã, Portugal; MPV, Musée paléontologique (Paléospace) de Villers-sur-Mer, Normandy, France; MUJA, Museo del Jurásico de Asturias, Colunga, Spain; NHMUK, Natural History Museum, London, United Kingdom; NMS, Naturmuseum Solothurn, Switzerland; OUMNH, Oxford University Museum of Natural History, United Kingdom; RBINS, Royal Belgian Institute of Natural Sciences, Brussels, Belgium; SMNS, Staatliches Museum für Naturkunde Stuttgart, Germany; TWCMS, Sunderland Museum and Art Gallery, United Kingdom.</p></sec></sec><sec id="s2"><label>3.</label><title><italic>Machimosaurus</italic> through time</title><sec id="s2a"><label>3.1</label><title>Bathonian</title><p>Based upon isolated tooth crowns from the Bathonian of France, Sauvage [<xref rid="RSOS140222C11" ref-type="bibr">11</xref>] established two species: <italic>Machimosaurus bathonicus</italic> and <italic>Machimosaurus rigauxi</italic>. Krebs [<xref rid="RSOS140222C7" ref-type="bibr">7</xref>, p. 48] considered these teeth more likely to be from <italic>Steneosaurus</italic>, as their apices are too pointed and the enamel surfaces too smooth to belong to <italic>Machimosaurus</italic>. As blunt apices and numerous apicobasal enamel ridges are apomorphies of <italic>Machimosaurus</italic> (see below), these tooth taxa cannot be referred to this genus.</p><p><italic>Machimosaurus</italic>/‘<italic>Steneosaurus</italic>’ <italic>obtusidens</italic>-like tooth crowns are also found in Bathonian deposits of the Great Oolite Group from England. One such tooth (TWCMS K1239) from Maidford in Northamptonshire has a blunt apex, and numerous apicobasal enamel ridges on the lingual and labial surface. This tooth crown is similar to the posterior-most tooth crowns of <italic>Steneosaurus larteti</italic> skulls (OUMNH J.29850 and OUMNH J.29851) from the Great Oolite Group of England (<xref ref-type="fig" rid="RSOS140222F1">figure 1</xref>). It is therefore possible that the ‘<italic>Machimosaurus</italic>’ tooth taxa from France are also referable to <italic>S. larteti</italic>; however, investigating that hypothesis is beyond the scope of this study.
<fig id="RSOS140222F1" orientation="portrait" position="float"><label>Figure 1.</label><caption><p><italic>Steneosaurus larteti</italic>, OUMNH J.29850 and OUMNH J.29851, referred specimens. (<italic>a</italic>) Photograph and (<italic>b</italic>) line drawing of the posterior end of the left mandibular rami in medial view. (<italic>c</italic>) Photograph of the skull in palatal view. ang, angular; ar, articular; pra, prearticular; sur, surangular.</p></caption><graphic xlink:href="rsos140222-g1"></graphic></fig></p></sec><sec id="s2b"><label>3.2</label><title>Callovian</title><p>‘<italic>Steneosaurus</italic>’<italic>obtusidens</italic> has been considered to be a subjective junior synonym of the Kimmeridgian taxon <italic>M. hugii</italic>, although most studies which drew this conclusion did note that further study was necessary on the anatomy and taxonomy of blunt-toothed teleosaurids [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C12" ref-type="bibr">12</xref>,<xref rid="RSOS140222C13" ref-type="bibr">13</xref>]. ‘<italic>Steneosaurus</italic>’ <italic>obtusidens</italic> is known from the Oxford Clay Formation of central England, and a specimen from the Marnes de Dives Formation of northern France has been referred to this species [<xref rid="RSOS140222C1" ref-type="bibr">1</xref>,<xref rid="RSOS140222C13" ref-type="bibr">13</xref>,<xref rid="RSOS140222C14" ref-type="bibr">14</xref>]. Recent studies, however, have considered ‘<italic>S</italic>.’ <italic>obtusidens</italic> to be distinct enough to warrant its own genus [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>,<xref rid="RSOS140222C15" ref-type="bibr">15</xref>], which we agree with. The holotype of this species is currently being re-described, which will help elucidate its anatomy and evolutionary relationships.</p><p>An isolated tooth crown (OUMNH J.14464) referred to <italic>M. rigauxi</italic> was found at Hanborough railway station, Oxfordshire, England (Cornbrash Formation) [<xref rid="RSOS140222C16" ref-type="bibr">16</xref>]. The description and figures of this specimen [<xref rid="RSOS140222C16" ref-type="bibr">16</xref>, p. 26–27, plate 1 fig. 5<italic>a</italic>–<italic>c</italic>] match the posterior dentition of the ‘<italic>S</italic>.’ <italic>obtusidens</italic> holotype, as the carinal keels are prominent and the apicobasal enamel ridges near the keels converge and contact the keel itself [<xref rid="RSOS140222C1" ref-type="bibr">1</xref>,<xref rid="RSOS140222C15" ref-type="bibr">15</xref>].</p><p>Based upon the Bathonian tooth taxa being more similar to <italic>Steneosaurus</italic> (in particular <italic>S. larteti</italic>) and ‘<italic>S</italic>.’ <italic>obtusidens</italic> being a distinct taxon, <italic>Machimosaurus</italic> is therefore unknown in the Middle Jurassic.</p></sec><sec id="s2c"><label>3.3</label><title>Oxfordian</title><p>An almost complete <italic>Machimosaurus</italic> mandible with isolated tooth crowns is known from the Upper Oxfordian (<italic>Perisphinctes</italic><italic>cautisnigrae</italic> NW European ammonite zone, <italic>Pe</italic>. <italic>variocostatus</italic> subzone) of Haudainville near Verdun (Département de la Meuse, northeastern France) (<xref ref-type="fig" rid="RSOS140222F2">figure 2</xref>). This mandible has previously been assigned to <italic>Steneosaurus</italic> cf. <italic>obtusidens</italic> and <italic>Machimosaurus</italic> cf. <italic>hugii</italic> (see [<xref rid="RSOS140222C12" ref-type="bibr">12</xref>] and the references therein). However, here we consider this specimen as <italic>Machimosaurus</italic> sp.
<fig id="RSOS140222F2" orientation="portrait" position="float"><label>Figure 2.</label><caption><p><italic>Machimosaurus</italic> sp., Musée de la Princerie (Verdun, France) 2007.0.14. Incomplete lower jaw in (<italic>a</italic>) dorsal view, (<italic>b</italic>) right oblique view and (<italic>c</italic>) left oblique view.</p></caption><graphic xlink:href="rsos140222-g2"></graphic></fig></p><p>Moreover, two isolated tooth crowns are known from the Upper Oxfordian Calcaire gréseux d'Hennequeville Formation. These teeth were found at Villerville, Département du Calvados, Basse-Normandie, France [<xref rid="RSOS140222C17" ref-type="bibr">17</xref>, p. 97–98, fig. 2].</p><p>Sauvage [<xref rid="RSOS140222C18" ref-type="bibr">18</xref>] reported and figured the first <italic>M. hugii</italic> specimens from Portugal. These included Oxfordian specimens, such as an isolated tooth from the Upper Oxfordian of Cesareda ‘Couches à <italic>Cidaris chofatti</italic>’ [<xref rid="RSOS140222C18" ref-type="bibr">18</xref>, plate 3 fig. 10] and a partial snout from Malhão, Algarve (one specimen label has: ‘Entre Amendoeira et Azinhal, Flanc nord de Malhão prés Estoy’) which is from the same horizon as <italic>Perisphinctes effrenatus</italic> [<xref rid="RSOS140222C18" ref-type="bibr">18</xref>, plate 3 fig. 9, and plate 5 figs 6 and 7].</p><p>Moreover, an isolated <italic>Machimosaurus</italic> sp. tooth crown (ML1208) was collected from Middle Oxfordian deposits at Cesaredas (39°N, 9°W) in central west Portugal.</p></sec><sec id="s2d"><label>3.4</label><title>Oxfordian–Kimmeridgian</title><p>From near Harrar in Ethiopia (Oxfordian or Kimmeridgian aged deposits), an anterior region of dentary was described as a new species of pliosaurid sauropterygian, cf. <italic>Simolestes nowackianus</italic> [<xref rid="RSOS140222C19" ref-type="bibr">19</xref>] (<xref ref-type="fig" rid="RSOS140222F3">figure 3</xref>). However, based on the dental morphology, the spatulate anterior region, arrangement of the dentary alveoli and thecodont tooth replacement, Bardet & Hua [<xref rid="RSOS140222C20" ref-type="bibr">20</xref>] demonstrated that it is in fact a large specimen of <italic>Machimosaurus</italic>. Currently, this is the only <italic>Machimosaurus</italic> specimen known from outside Europe.
<fig id="RSOS140222F3" orientation="portrait" position="float"><label>Figure 3.</label><caption><p><italic>Machimosaurus nowackianus</italic> comb. nov., GPIT Orig. Huene 1938 fig. 1–4, holotype. Incomplete dentary in dorsal view, (<italic>a</italic>) photograph and (<italic>b</italic>) line drawing.</p></caption><graphic xlink:href="rsos140222-g3"></graphic></fig></p></sec><sec id="s2e"><label>3.5</label><title>Kimmeridgian</title><p>Prior to this study, two valid species of <italic>Machimosaurus</italic> were recognized in the Kimmeridgian of Europe: the type species <italic>M. hugii</italic> and <italic>M. mosae</italic> [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C21" ref-type="bibr">21</xref>]. Based on the numerous European localities outlined below, during the Kimmeridgian <italic>Machimosaurus</italic> commonly frequented shallow marine ecosystems, with the occasional individual known from brackish and open-shelf environments, and possibly also freshwater environments.</p><p>From Portugal, <italic>Machimosaurus</italic> is known from two sites:
<list list-type="simple"><list-item><p>— In 1943, the geologist Carlos Teixeira reported an isolated <italic>M. hugii</italic> tooth from Lagares (Colmeias, near Leiria), Portugal [<xref rid="RSOS140222C22" ref-type="bibr">22</xref>]. This tooth (MG 25) is from the Alcobaça Beds Formation (Upper Kimmeridgian).</p></list-item><list-item><p>— The largest known specimen of <italic>Machimosaurus</italic> is known from the Guimarota site, Leiria, Portugal (Alcobaça Beds Formation) [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>] (<xref ref-type="fig" rid="RSOS140222F4">figures 4</xref>–<xref ref-type="fig" rid="RSOS140222F10">10</xref>). The Guimarota site was deposited in either a lagoonal environment with some freshwater influx or a wooded swamp similar to extant mangrove forests. This locality has yielded several species of crocodylomorphs such as <italic>Lusitanisuchus mitracostatus</italic>, <italic>Goniopholis baryglyphaeus</italic> and <italic>Theriosuchus guimarotae</italic> [<xref rid="RSOS140222C23" ref-type="bibr">23</xref>,<xref rid="RSOS140222C24" ref-type="bibr">24</xref>].</p></list-item></list><fig id="RSOS140222F4" orientation="portrait" position="float"><label>Figure 4.</label><caption><p><italic>Machimosaurus hugii</italic>, MG-8730-1, referred specimen. Incomplete snout (fragment of maxilla), (<italic>a</italic>,<italic>c</italic>) photographs in both lateral views and (<italic>b</italic>,<italic>d</italic>) the corresponding line drawings in both lateral views.</p></caption><graphic xlink:href="rsos140222-g4"></graphic></fig><fig id="RSOS140222F5" orientation="portrait" position="float"><label>Figure 5.</label><caption><p><italic>Machimosaurus hugii</italic>, MG-8730-1, referred specimen. Incomplete snout (fragment of maxilla), (<italic>a</italic>) photograph in ventral view, (<italic>b</italic>) line drawing in ventral view, (<italic>c</italic>) photograph in posterior view and (<italic>d</italic>) line drawing in posterior view. mx, maxilla.</p></caption><graphic xlink:href="rsos140222-g5"></graphic></fig><fig id="RSOS140222F6" orientation="portrait" position="float"><label>Figure 6.</label><caption><p><italic>Machimosaurus hugii</italic>, MG-8730-1, referred specimen. Incomplete snout (fragment of maxilla, and possibly nasals), (<italic>a</italic>) photograph in right lateral view, (<italic>b</italic>) line drawing in right lateral view, (<italic>c</italic>) photograph in left lateral view and (<italic>d</italic>) line drawing in left lateral view. mx, maxilla.</p></caption><graphic xlink:href="rsos140222-g6"></graphic></fig><fig id="RSOS140222F7" orientation="portrait" position="float"><label>Figure 7.</label><caption><p><italic>Machimosaurus hugii</italic>, MG-8730-1, referred specimen. Incomplete snout (fragment of maxilla, and possibly nasals), (<italic>a</italic>) photograph in dorsal view, (<italic>b</italic>) line drawing in dorsal view, (<italic>c</italic>) photograph in ventral view and (<italic>d</italic>) line drawing in ventral view. mx, maxilla.</p></caption><graphic xlink:href="rsos140222-g7"></graphic></fig><fig id="RSOS140222F8" orientation="portrait" position="float"><label>Figure 8.</label><caption><p><italic>Machimosaurus hugii</italic>, MG-8730-2, referred specimen. Incomplete braincase, (<italic>a</italic>) photograph in dorsal view, (<italic>b</italic>) line drawing in dorsal view, (<italic>c</italic>) photograph in ventral view, (<italic>d</italic>) line drawing in ventral view, (<italic>e</italic>) photograph in occipital view and (<italic>f</italic>) line drawing in occipital view. bo, basioccipital; bt, basioccipital tuberosities; eo, exoccipital; pop, paroccipital process of the opisthotic; qu, quadrate.</p></caption><graphic xlink:href="rsos140222-g8"></graphic></fig><fig id="RSOS140222F9" orientation="portrait" position="float"><label>Figure 9.</label><caption><p><italic>Machimosaurus hugii</italic>, MG unnumbered, referred specimen. Isolated tooth crown in (<italic>a</italic>) right lateral view and (<italic>b</italic>) apical view.</p></caption><graphic xlink:href="rsos140222-g9"></graphic></fig><fig id="RSOS140222F10" orientation="portrait" position="float"><label>Figure 10.</label><caption><p><italic>Machimosaurus hugii</italic>, MG unnumbered, referred specimen. First sacral vertebra in (<italic>a</italic>) posterior view, (<italic>b</italic>) anterior view, (<italic>c</italic>) dorsal view and (<italic>d</italic>) ventral view.</p></caption><graphic xlink:href="rsos140222-g10"></graphic></fig></p><p>From Spain:
<list list-type="simple"><list-item><p>— Isolated cf. <italic>Machimosaurus</italic> teeth have been reported from the Tereñes Formation of the Asturias coast, Northern Spain [<xref rid="RSOS140222C25" ref-type="bibr">25</xref>]. This formation is considered to represent a shallow tide-less sea [<xref rid="RSOS140222C26" ref-type="bibr">26</xref>]. Teeth are also known from the Lastres Formation in Asturias [<xref rid="RSOS140222C25" ref-type="bibr">25</xref>], a fluvial-dominated deltaic system in origin [<xref rid="RSOS140222C26" ref-type="bibr">26</xref>] (<xref ref-type="fig" rid="RSOS140222F11">figure 11</xref>).</p></list-item><list-item><p>— The Kimmeridgian ichnogenus <italic>Hatcherichnus</italic> is known from coastal and deltaic units of Asturias, Spain. It has been suggested that these track ways were made by either <italic>Machimosaurus</italic> or a large goniopholidid [<xref rid="RSOS140222C27" ref-type="bibr">27</xref>].</p></list-item></list><fig id="RSOS140222F11" orientation="portrait" position="float"><label>Figure 11.</label><caption><p><italic>Machimosaurus hugii</italic>, MUJA-1298 and MUJA-1008, referred specimens. Isolated tooth crowns, (<italic>a</italic>,<italic>b</italic>) MUJA-1298 and (<italic>c</italic>–<italic>e</italic>) MUJA-1008.</p></caption><graphic xlink:href="rsos140222-g11"></graphic></fig></p><p>From France:
<list list-type="simple"><list-item><p>— The anterior half of a rostrum and mandible in occlusion (premaxilla, maxilla and dentary) that has been attributed to <italic>M. hugii</italic> is known from the Calcaires Coquilliers Formation (<italic>Pictonia baylei</italic> Sub-Boreal ammonite zone, lowermost Kimmeridgian) of Cricqueboeuf, Normandy, Northern France [<xref rid="RSOS140222C2" ref-type="bibr">2</xref>,<xref rid="RSOS140222C17" ref-type="bibr">17</xref>] (<xref ref-type="fig" rid="RSOS140222F12">figures 12</xref>–<xref ref-type="fig" rid="RSOS140222F14">14</xref>). During the Early Kimmeridgian, the Calcaires Coquilliers Formation was deposited in a homoclinal mid-ramp with significant storm-wave reworking [<xref rid="RSOS140222C28" ref-type="bibr">28</xref>].</p></list-item><list-item><p>— The most complete skull of <italic>M. hugii</italic> was discovered from Ain, France (Lower Kimmeridgian) [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>]. During the Kimmeridgian–Tithonian this region was a lagoonal environment [<xref rid="RSOS140222C29" ref-type="bibr">29</xref>].</p></list-item><list-item><p>— An almost complete skeleton of <italic>M. mosae</italic> was discovered near Ambleteuse, Boulonnais, France (Argiles de Châtillon Formation, either the <italic>Aulacostephanus</italic><italic>autissiodorensis</italic> Sub-Boreal ammonite zone, uppermost Kimmeridgian or the <italic>Gravesia gigas</italic>/<italic>Pectinatites elegans</italic> Sub-Boreal ammonite zone, lowermost Tithonian) [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C21" ref-type="bibr">21</xref>] (<xref ref-type="fig" rid="RSOS140222F15">figures 15</xref>–<xref ref-type="fig" rid="RSOS140222F20">20</xref>). The Argiles de Châtillon Formation was deposited in a nearshore or shallow-shelf marine environment off the west coast of the London–Brabant Massive [<xref rid="RSOS140222C30" ref-type="bibr">30</xref>].</p></list-item></list><fig id="RSOS140222F12" orientation="portrait" position="float"><label>Figure 12.</label><caption><p><italic>Machimosaurus buffetauti</italic> sp. nov., MPV V1600.Bo, referred specimen. Anterior region of the snout and lower jaw, (<italic>a</italic>) photograph in left lateral view, (<italic>b</italic>) line drawing in left lateral view, (<italic>c</italic>) photograph in right lateral view and (<italic>d</italic>) line drawing in right lateral view. den, dentary; mx, maxilla; pmx, premaxilla.</p></caption><graphic xlink:href="rsos140222-g12"></graphic></fig><fig id="RSOS140222F13" orientation="portrait" position="float"><label>Figure 13.</label><caption><p><italic>Machimosaurus buffetauti</italic> sp. nov., MPV V1600.Bo, referred specimen. Anterior region of the snout and lower jaw, (<italic>a</italic>) photograph in dorsal view, (<italic>b</italic>) line drawing in dorsal view, (<italic>c</italic>) photograph in ventral view and (<italic>d</italic>) line drawing in ventral view. mx, maxilla; pmx, premaxilla.</p></caption><graphic xlink:href="rsos140222-g13"></graphic></fig><fig id="RSOS140222F14" orientation="portrait" position="float"><label>Figure 14.</label><caption><p><italic>Machimosaurus buffetauti</italic> sp. nov., MPV V1601.Bo, referred specimen. Middle region of the snout, (<italic>a</italic>) photograph in dorsal view, (<italic>b</italic>) line drawing in dorsal view, (<italic>c</italic>) photograph in ventral view and (<italic>d</italic>) line drawing in ventral view. mx, maxilla; na, nasals.</p></caption><graphic xlink:href="rsos140222-g14"></graphic></fig><fig id="RSOS140222F15" orientation="portrait" position="float"><label>Figure 15.</label><caption><p><italic>Machimosaurus mosae</italic>, neotype. Skull (<italic>a</italic>) photograph in dorsal view and (<italic>b</italic>) line drawing in dorsal view; mandible (<italic>c</italic>) photograph in dorsal view and (<italic>d</italic>) line drawing in dorsal view. ar, articular; den, dentary; fr, frontal; mx, maxilla; na, nasals; oc, occipital condyle; pmx, premaxilla; qu, quadrate; sp, splenial.</p></caption><graphic xlink:href="rsos140222-g15"></graphic></fig><fig id="RSOS140222F16" orientation="portrait" position="float"><label>Figure 16.</label><caption><p><italic>Machimosaurus mosae</italic>, neotype. Skull (orbital and temporal region) in (<italic>a</italic>) dorsal view and (<italic>b</italic>) ventral view.</p></caption><graphic xlink:href="rsos140222-g16"></graphic></fig><fig id="RSOS140222F17" orientation="portrait" position="float"><label>Figure 17.</label><caption><p><italic>Machimosaurus mosae</italic>, neotype. Skull (rostrum) in (<italic>a</italic>) dorsal view and (<italic>b</italic>) ventral view.</p></caption><graphic xlink:href="rsos140222-g17"></graphic></fig><fig id="RSOS140222F18" orientation="portrait" position="float"><label>Figure 18.</label><caption><p><italic>Machimosaurus mosae</italic>, neotype. Skull close-ups, (<italic>a</italic>) pterygoid, internal choana, basisphenoid and basioccipital in palatal view, (<italic>b</italic>) occipit in occipital/posterior view, (<italic>c</italic>) orbital region in left lateral view and (<italic>d</italic>) left quadrate, squamosal and paroccipital process region in lateral view.</p></caption><graphic xlink:href="rsos140222-g18"></graphic></fig><fig id="RSOS140222F19" orientation="portrait" position="float"><label>Figure 19.</label><caption><p><italic>Machimosaurus mosae</italic>, neotype. Lower jaw (lacking the posterior ends of the rami) in (<italic>a</italic>) dorsal view and (<italic>b</italic>) ventral view.</p></caption><graphic xlink:href="rsos140222-g19"></graphic></fig><fig id="RSOS140222F20" orientation="portrait" position="float"><label>Figure 20.</label><caption><p><italic>Machimosaurus mosae</italic>, neotype. Postcrania, (<italic>a</italic>) keeled ventral osteoderm, (<italic>b</italic>) ventral osteoderm, (<italic>c</italic>) dorsal osteoderm, (<italic>d</italic>) dorsal osteoderm, (<italic>e</italic>) right coracoid in medial view, (<italic>f</italic>) left pubis in medial view and (<italic>g</italic>) left ischium in lateral view.</p></caption><graphic xlink:href="rsos140222-g20"></graphic></fig></p><p>From Germany:
<list list-type="simple"><list-item><p>— von Meyer [<xref rid="RSOS140222C31" ref-type="bibr">31</xref>] referred a tooth from Kahlenberg, near Hannover, in Lower Saxony to <italic>Machimosaurus</italic>. This locality is now within the urban area of Hannover [<xref rid="RSOS140222C32" ref-type="bibr">32</xref>].</p></list-item><list-item><p>— Numerous isolated teeth have been discovered at the Oker quarry, Langenberg area (Langenberg Formation) of Lower Saxony [<xref rid="RSOS140222C8" ref-type="bibr">8</xref>,<xref rid="RSOS140222C32" ref-type="bibr">32</xref>] (<xref ref-type="fig" rid="RSOS140222F21">figure 21</xref><italic>c</italic>–<italic>h</italic>). Sediments from this area were deposited in a shallow-water basin, either a bay or a lagoon, and, along with <italic>Machimosaurus</italic>, numerous other crocodylomorphs are known from this region: <italic>Goniopholis simus</italic>, <italic>Theriosuchus pusillus</italic>, <italic>Steneosaurus</italic> sp. and two gen. et sp. nov. [<xref rid="RSOS140222C32" ref-type="bibr">32</xref>].</p></list-item><list-item><p>— A skull, lower jaw and partial postcranial skeleton of <italic>M. hugii</italic> was discovered in a quarry at Neuffen, Baden-Württemberg (<italic>Ataxioceras hypselocyclum</italic> Sub-Mediterranean ammonite zone, Weißer Jura gamma 2, Lower Kimmeridgian) [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] (<xref ref-type="fig" rid="RSOS140222F22">figures 22</xref>–<xref ref-type="fig" rid="RSOS140222F27">27</xref>).</p></list-item><list-item><p>— <italic>Machimosaurus</italic> has also been reported from Fritzow, Mecklenburg-Vorpommern, Germany [<xref rid="RSOS140222C33" ref-type="bibr">33</xref>].</p></list-item></list><fig id="RSOS140222F21" orientation="portrait" position="float"><label>Figure 21.</label><caption><p>Isolated tooth crowns either referable, or likely to be referable to <italic>M. buffetauti</italic> sp. nov. NHMUK PV R1774 in (<italic>a</italic>) lingual view and (<italic>b</italic>) left lateral view; DFMMh FV 330 in (<italic>c</italic>) right lateral view, (<italic>d</italic>) lingual view, (<italic>e</italic>) left lateral view, (<italic>f</italic>) labial view, (<italic>g</italic>) apical view and (<italic>h</italic>) basal view.</p></caption><graphic xlink:href="rsos140222-g21"></graphic></fig><fig id="RSOS140222F22" orientation="portrait" position="float"><label>Figure 22.</label><caption><p><italic>Machimosaurus buffetauti</italic> sp. nov., SMNS 91415, holotype. Skull (with associated postcrania) in dorsal view, (<italic>a</italic>) photograph and (<italic>b</italic>) line drawing. fr, frontal; mx, maxilla; na, nasals; pre, prefrontal; pmx, premaxilla.</p></caption><graphic xlink:href="rsos140222-g22"></graphic></fig><fig id="RSOS140222F23" orientation="portrait" position="float"><label>Figure 23.</label><caption><p><italic>Machimosaurus buffetauti</italic> sp. nov., SMNS 91415, holotype. Skull (with lower jaw and associated postcrania) in right lateral view, (<italic>a</italic>) photograph and (<italic>b</italic>) line drawing. den, dentary; fr, frontal; lac, lacrimal; mx, maxilla; pmx, premaxilla.</p></caption><graphic xlink:href="rsos140222-g23"></graphic></fig><fig id="RSOS140222F24" orientation="portrait" position="float"><label>Figure 24.</label><caption><p><italic>Machimosaurus buffetauti</italic> sp. nov., SMNS 91415, holotype. Skull in occipital view, (<italic>a</italic>) photograph and (<italic>b</italic>) line drawing. bo, basioccipital; bt, basioccipital tuberosities; eo, exoccipital; oc, occipital condyle; qu, quadrate; so, supraoccipital.</p></caption><graphic xlink:href="rsos140222-g24"></graphic></fig><fig id="RSOS140222F25" orientation="portrait" position="float"><label>Figure 25.</label><caption><p><italic>Machimosaurus buffetauti</italic> sp. nov., SMNS 91415, holotype. Lower jaw (with associated postcrania) in dorsal view, (<italic>a</italic>) photograph and (<italic>b</italic>) line drawing. ar, articular; den, dentary; pra, prearticular; sp, splenial.</p></caption><graphic xlink:href="rsos140222-g25"></graphic></fig><fig id="RSOS140222F26" orientation="portrait" position="float"><label>Figure 26.</label><caption><p><italic>Machimosaurus buffetauti</italic> sp. nov., SMNS 91415, holotype. Lower jaw (with skull and associated postcrania) in right lateral view, (<italic>a</italic>) photograph and (<italic>b</italic>) line drawing. den, dentary.</p></caption><graphic xlink:href="rsos140222-g26"></graphic></fig><fig id="RSOS140222F27" orientation="portrait" position="float"><label>Figure 27.</label><caption><p><italic>Machimosaurus buffetauti</italic> sp. nov., SMNS 91415, holotype. (<italic>a</italic>) Close-up on the dentary dentition, (<italic>b</italic>) the atlas–axis in lateral view and (<italic>c</italic>) the dorsal osteoderms in dorsal view.</p></caption><graphic xlink:href="rsos140222-g27"></graphic></fig></p><p>From Switzerland, <italic>Machimosaurus</italic> material is known from:
<list list-type="simple"><list-item><p>— A dorsal vertebra attributed to <italic>M</italic>. <italic>hugii</italic> from Moutier, Canton Bern (possibly Early Kimmeridgian) [<xref rid="RSOS140222C34" ref-type="bibr">34</xref>]. In addition, a broken sauropod dinosaur (<italic>Cetiosauriscus greppini</italic>) femur has bite marks matching <italic>Machimosaurus</italic> teeth [<xref rid="RSOS140222C34" ref-type="bibr">34</xref>]. Interestingly, it has been suggested that these specimens were buried in freshwater sediments due to the greenish marl [<xref rid="RSOS140222C34" ref-type="bibr">34</xref>].</p></list-item><list-item><p>— Isolated teeth are known from the ‘Solothurn Turtle Limestone’ (uppermost part of the Reuchenette Formation, Late Kimmeridgian, <italic>Hybonoticeras</italic><italic>beckeri</italic> Sub-Mediterranean ammonite zone [<xref rid="RSOS140222C35" ref-type="bibr">35</xref>]) of Solothurn, Canton Solothurn (<xref ref-type="fig" rid="RSOS140222F28">figures 28</xref> and <xref ref-type="fig" rid="RSOS140222F29">29</xref>). Marine turtle shells (Plesiochelyidae) discovered from this limestone are known to have bite marks matching <italic>Machimosaurus</italic> teeth, and in some instances still have <italic>Machimosaurus</italic> teeth imbedded within them [<xref rid="RSOS140222C36" ref-type="bibr">36</xref>] (<xref ref-type="fig" rid="RSOS140222F30">figure 30</xref>). The ‘Solothurn Turtle Limestone’ is interpreted as being a shallow protected lagoon [<xref rid="RSOS140222C37" ref-type="bibr">37</xref>]. The type series (isolated tooth crowns) of <italic>M. hugii</italic> were found in this limestone [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>,<xref rid="RSOS140222C31" ref-type="bibr">31</xref>].</p></list-item><list-item><p>— A second sea turtle assemblage found in the Virgula Marls near Porrentruy, Canton Jura (Late Kimmeridgian, <italic>A</italic>. <italic>eudoxus</italic> Sub-Mediterranean ammonite zone [<xref rid="RSOS140222C35" ref-type="bibr">35</xref>]) also has yielded isolated <italic>Machimosaurus</italic> teeth [<xref rid="RSOS140222C38" ref-type="bibr">38</xref>].</p></list-item></list><fig id="RSOS140222F28" orientation="portrait" position="float"><label>Figure 28.</label><caption><p><italic>Machimosaurus hugii</italic>, NMS 8342, lectotype. Isolated tooth crown in (<italic>a</italic>) labial view, (<italic>b</italic>) lingual view and (<italic>c</italic>) apical view.</p></caption><graphic xlink:href="rsos140222-g28"></graphic></fig><fig id="RSOS140222F29" orientation="portrait" position="float"><label>Figure 29.</label><caption><p><italic>Machimosaurus hugii</italic> referred specimens. NHMUK PV OR33239 in (<italic>a</italic>) labial view and (<italic>b</italic>) right lateral view; NHMUK PV R5 in (<italic>c</italic>) labial view and (<italic>d</italic>) right lateral view.</p></caption><graphic xlink:href="rsos140222-g29"></graphic></fig><fig id="RSOS140222F30" orientation="portrait" position="float"><label>Figure 30.</label><caption><p>Marine turtle specimens from the Solothurn Turtle Limestone of Switzerland with <italic>Machimosaurus</italic> bite marks. All are from Steinbruch, Solothurn. Plastron fragments of Testudinata indet. NMS 23828 in (<italic>a</italic>) dorsal view and (<italic>b</italic>) ventral view. Plastron fragments of NMS 23829 in (<italic>c</italic>) dorsal view and (<italic>d</italic>) ventral view. (<italic>e</italic>) Carapace of <italic>Plesiochelys</italic> sp. NMS 21499.</p></caption><graphic xlink:href="rsos140222-g30"></graphic></fig></p><p>From Poland, isolated <italic>Machimosaurus</italic> teeth are known from Lower Kimmeridgian deposits of the Czarnogłowy quarry, West Pomerania (note that prior to 1945 Czarnogłowy was in Germany and was called Zarnglaff, and that this name is used in pre-1945 literature) [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C39" ref-type="bibr">39</xref>,<xref rid="RSOS140222C40" ref-type="bibr">40</xref>] (<xref ref-type="fig" rid="RSOS140222F31">figures 31</xref> and <xref ref-type="fig" rid="RSOS140222F32">32</xref>). Dzik [<xref rid="RSOS140222C40" ref-type="bibr">40</xref>, fig. 9.20C] figured a mandibular fragment (symphyseal region lacking the anterior-most half/third) from Czarnogłowy as <italic>Machimosaurus</italic>. However, based on comparisons between <italic>Steneosaurus</italic> and <italic>Machimosaurus</italic> specimens, this partial mandible is in fact <italic>Steneosaurus</italic>, as: (i) it has a proportionally narrow mandible with a high tooth count; (ii) the anterior-most preserved alveoli have inter-alveolar spaces which are too long, being greater than the length of the adjacent alveoli; (iii) the splenial is very elongated and has at least 16 pairs of symphyseal alveoli adjacent; and (iv) the Meckelian groove is very deeply excavated, especially at the mandibular midline [<xref rid="RSOS140222C1" ref-type="bibr">1</xref>,<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C10" ref-type="bibr">10</xref>,<xref rid="RSOS140222C14" ref-type="bibr">14</xref>].
<fig id="RSOS140222F31" orientation="portrait" position="float"><label>Figure 31.</label><caption><p><italic>Machimosaurus</italic> cf.<italic>buffetauti</italic>, GPIT/RE/9280. Isolated tooth crown in (<italic>a</italic>) right lateral view, (<italic>b</italic>) labial view, (<italic>c</italic>) left lateral view, (<italic>d</italic>) lingual view and (<italic>e</italic>) apical view.</p></caption><graphic xlink:href="rsos140222-g31"></graphic></fig><fig id="RSOS140222F32" orientation="portrait" position="float"><label>Figure 32.</label><caption><p><italic>Machimosaurus</italic> cf. <italic>buffetauti</italic>. Isolated tooth crown GPIT/RE/328 in (<italic>a</italic>) lingual view, (<italic>b</italic>) labial view and (<italic>c</italic>) apical view. Isolated tooth crown GPIT/RE/9281 in (<italic>d</italic>) right lateral view, (<italic>e</italic>) labial view and (<italic>f</italic>) apical view.</p></caption><graphic xlink:href="rsos140222-g32"></graphic></fig></p><p>From England,<italic>Machimosaurus</italic> is solely known from a single incomplete isolated tooth crown discovered at Smallmouth Sands, Dorset (Lower Kimmeridge Clay Formation) [<xref rid="RSOS140222C5" ref-type="bibr">5</xref>, fig. 215A,B]. Until recently, <italic>M. mosae</italic> was considered to be present in the Upper Kimmeridge Clay Formation of England (Early Tithonian). This was based on a very large skull and mandible from Kimmeridge in Dorset, which was recently shown to pertain to the metriorhynchid crocodylomorph <italic>Plesiosuchus manselii</italic> [<xref rid="RSOS140222C41" ref-type="bibr">41</xref>]. This means that the single tooth crown from Smallmouth Sands is the only <italic>Machimosaurus</italic> specimen known from England. Although the Dorset succession of the Kimmeridge Clay Formation is considered to have formed at an outer-shelf water depth of between 150 and 200 m [<xref rid="RSOS140222C42" ref-type="bibr">42</xref>], during the Early Kimmeridgian the water depth was very shallow in the Dorset succession, between 10 and 30 m [<xref rid="RSOS140222C43" ref-type="bibr">43</xref>].</p></sec><sec id="s2f"><label>3.6</label><title>The Kimmeridgian–Tithonian boundary</title><p>All <italic>Machimosaurus</italic> specimens from the Lourinhã Formation in Portugal are from the Praia Azul Member, which was a brackish to coastal platform that comprised the Kimmeridgian–Tithonian boundary at 152.1 Ma [<xref rid="RSOS140222C44" ref-type="bibr">44</xref>,<xref rid="RSOS140222C45" ref-type="bibr">45</xref>]. However this can be better dated as <italic>ca</italic> the Upper Kimmeridgian–Lower Tithonian transition. The rest of the Lourinhã Formation extends to the Jurassic–Cretaceous boundary, but the sediments are strictly continental (flood-plain mudstones and fluvial sandstone bodies [<xref rid="RSOS140222C46" ref-type="bibr">46</xref>]). Thus, the absence of <italic>Machimosaurus</italic> in the Tithonian of Portugal is solely due to a shift in palaeoenvironment, rather than a true disappearance of the genus in the Tithonian. Sauvage [<xref rid="RSOS140222C18" ref-type="bibr">18</xref>] reported an isolated tooth from the Upper Kimmeridgian–Lower Tithonian of Santa-Cruz (Praia Azul Member, Lourinhã Formation [<xref rid="RSOS140222C44" ref-type="bibr">44</xref>]). New discoveries in the Lourinhã area include: several isolated teeth, all from the Praia Azul Member (<italic>sensu</italic> [<xref rid="RSOS140222C44" ref-type="bibr">44</xref>]), from the following localities: Porto das Barcas (ML491, ML495, ML959 and ML1955), Peralta (ML647), Zimbral (ML657 and ML658) and around coastline (ML647, ML733 and ML902) (<xref ref-type="fig" rid="RSOS140222F33">figures 33</xref> and <xref ref-type="fig" rid="RSOS140222F34">34</xref>). Some of these tooth crowns are very large in size: the tooth ML495 from Porto das Barcas (N39°13.943′; 9°20.349′) has an apicobasal length of 41.4 mm (crown alone is 27.12 mm) and maximal diameter of 24.2 mm (<xref ref-type="fig" rid="RSOS140222F35">figure 35</xref>).
<fig id="RSOS140222F33" orientation="portrait" position="float"><label>Figure 33.</label><caption><p><italic>Machimosaurus hugii</italic>, referred specimens. Isolated tooth crown (<italic>a</italic>) ML 491 in labial view, (<italic>b</italic>) ML 657 in labial view, (<italic>c</italic>) ML 658 in labial view and (<italic>d</italic>) ML 657 in lingual/lateral view.</p></caption><graphic xlink:href="rsos140222-g33"></graphic></fig><fig id="RSOS140222F34" orientation="portrait" position="float"><label>Figure 34.</label><caption><p><italic>Machimosaurus hugii</italic>, ML 647, referred specimens. First isolated tooth crown in (<italic>a</italic>) left lateral view. The second tooth crown in (<italic>b</italic>) labial view, (<italic>c</italic>) left lateral view, (<italic>d</italic>) right lateral view, (<italic>e</italic>) lingual view and (<italic>f</italic>) basal view.</p></caption><graphic xlink:href="rsos140222-g34"></graphic></fig><fig id="RSOS140222F35" orientation="portrait" position="float"><label>Figure 35.</label><caption><p><italic>Machimosaurus hugii</italic>, ML 495, referred specimen. Isolated tooth crown in (<italic>a</italic>) labial view, (<italic>b</italic>) right lateral view, (<italic>c</italic>) lingual view, (<italic>d</italic>) left lateral view, (<italic>e</italic>) apical view and (<italic>f</italic>) basal view.</p></caption><graphic xlink:href="rsos140222-g35"></graphic></fig></p><p>A large vertebra previously referred to cf. <italic>Machimosaurus</italic> [<xref rid="RSOS140222C47" ref-type="bibr">47</xref>] is here no longer regarded as a crocodylomorph. The Lourinhã Formation has a rich and diverse dinosaur fauna, including eggs and tracks, but crocodylomorph remains are also common, mostly <italic>Goniopholis</italic> and <italic>Theriosuchus</italic>.</p></sec><sec id="s2g"><label>3.7</label><title>Tithonian</title><p>An isolated <italic>Machimosaurus</italic> tooth is known from marine deposits of the Higueruelas Formation at Buñol, Valencia Province, Eastern Spain [<xref rid="RSOS140222C48" ref-type="bibr">48</xref>].</p><p>Sauvage [<xref rid="RSOS140222C11" ref-type="bibr">11</xref>, planche 3] stated that the holotype (‘type de l'espèce’) of <italic>M. interruptus</italic> is from the ‘Portlandien à <italic>Ammonites portlandicus</italic> de Mont-Lambert (near Boulogne-sur-Mer, France)’. As <italic>Ammonities portlandicus</italic> is a synonym of <italic>G. gigas</italic> [<xref rid="RSOS140222C11" ref-type="bibr">11</xref>], it places this tooth in the Early Tithonian <italic>G. gigas</italic>/<italic>P. elegans</italic> Sub-Boreal ammonite zone.</p></sec><sec id="s2h"><label>3.8</label><title>Berriasian</title><p>Isolated teeth from Spain, with a possible Berriasian age, have previously been attributed to <italic>Machimosaurus</italic>. In 1916–1918, JM Catalá discovered a series of fossil vertebrates from Benagéber, Valencia province, Spain. The description of Beltrán [<xref rid="RSOS140222C49" ref-type="bibr">49</xref>] stated that the fossils were from the ‘Wealdense’ (i.e. Wealden facies) and that <italic>Gonophilus</italic> (sic) teeth were among the collection. Royo y Gómez [<xref rid="RSOS140222C50" ref-type="bibr">50</xref>] reviewed the Catalá collection when it was temporarily loaned to the Museo Nacional de Ciencias Naturales in Madrid. Two species of crocodylomorphs were reported as being present: <italic>Goniopholis</italic> and <italic>Machimosaurus</italic>. One year later, Royo y Gómez assigned the Benagéber crocodiles to<italic>Steneosaurus</italic> cf. <italic>obtusidens</italic> and <italic>Machimosaurus</italic> sp. nov. [<xref rid="RSOS140222C51" ref-type="bibr">51</xref>], specifying its age as ‘Purbequiense’ (i.e. Purbeck facies) [<xref rid="RSOS140222C52" ref-type="bibr">52</xref>]. Therefore, it is possible that the teeth were Tithonian or Berriasian in age.</p><p>Unfortunately, these teeth were never described or figured and are currently missing. It is possible that they were destroyed in the fire that ruined the Museo de Historia Natural de la Universidad de Valencia in 1932, were Beltrán was a professor [<xref rid="RSOS140222C53" ref-type="bibr">53</xref>]. As such, the presence of <italic>Machimosaurus</italic> in the earliest Cretaceous of Spain cannot currently be confirmed.</p></sec><sec id="s2i"><label>3.9</label><title>Valanginian</title><p>A partial dentary with <italic>in situ</italic> tooth crowns from the Valanginian of southern France (Département des Bouches-du-Rhône) was described as <italic>Steneosaurus</italic> sp. due to a superficial similarity in dental morphology between it and ‘<italic>S</italic>.’ <italic>obtusidens</italic> [<xref rid="RSOS140222C54" ref-type="bibr">54</xref>]. However, recent re-examination of the Valanginian specimen demonstrated it to be a metriorhynchid closely related to <italic>P. manselii</italic> [<xref rid="RSOS140222C55" ref-type="bibr">55</xref>].</p></sec><sec id="s2j"><label>3.10</label><title>Upper Hauterivian–Lower Barremian</title><p>Sanz <italic>et al.</italic> [<xref rid="RSOS140222C56" ref-type="bibr">56</xref>] referred isolated tooth crowns from the Lower Cretaceous of Galve (Teruel province, Spain) to cf. <italic>Machimosaurus</italic> sp. These teeth come from the sediments on the top of El Castellar Formation (uppermost Hauterivian–lowermost Barremian), which is lacustrine in origin [<xref rid="RSOS140222C57" ref-type="bibr">57</xref>]. Intriguingly, <italic>Machimosaurus</italic> was not cited in the subsequent papers on the Galve crocodylomorphs made by the same authors (e.g. [<xref rid="RSOS140222C58" ref-type="bibr">58</xref>,<xref rid="RSOS140222C59" ref-type="bibr">59</xref>]), indicating that they may have been hesitant in this taxonomic assignment.</p><p>The teeth were described as being: ‘distinguished by blunt crowns with a very characteristic enamel ornamentation of anastomosed and braided ridges’ [<xref rid="RSOS140222C52" ref-type="bibr">52</xref>, p. 207]. One of the teeth was figured [<xref rid="RSOS140222C56" ref-type="bibr">56</xref>, p. 208, fig. 3<italic>a</italic>,<italic>b</italic>], and based on that line drawing we do not consider it referable to <italic>Machimosaurus</italic>. This is due to the tooth's strong lingual curvature, concave lingual surface and strongly pronounced mesial and distal carinae, all characteristics not seen among <italic>Machimosaurus</italic> teeth.</p><p>Gasca <italic>et al.</italic> [<xref rid="RSOS140222C60" ref-type="bibr">60</xref>] mentioned aff. <italic>Machimosaurus</italic> teeth in the Lower Cretaceous of Allepuz (Teruel province, Spain). These teeth came from a microvertebrate site in the Camarillas Formation (Lower Barremian) that originated in an avulsion deposit, namely an ephemeral fluviatile pond, and presents a mixture of terrestrial (theropod dinosaurs and crocodylomorphs: Bernissartiidae, Atoposauridae, aff. <italic>Machimosaurus</italic>) and freshwater vertebrates (hybodontid sharks, bony fishes and amphibians). Unfortunately, these teeth have never been described or figured, thus we cannot determine the reliability of this taxonomic assessment. Therefore, there is no evidence that <italic>Machimosaurus</italic>, or any other teleosaurid, survived into the Cretaceous.</p></sec></sec><sec id="s3"><label>4.</label><title>Recent taxonomic changes to <italic>Machimosaurus hugii</italic></title><sec id="s3a"><label>4.1</label><title>The diverse and long-lived <italic>Machimosaurus hugii</italic></title><p>Recently, Pierce <italic>et al</italic>. [<xref rid="RSOS140222C9" ref-type="bibr">9</xref>] proposed that <italic>M. hugii</italic> was the senior subjective synonym of various Callovian teleosaurids: ‘<italic>S</italic>.’ <italic>obtusidens</italic>, <italic>Steneosaurus durobrivensis</italic> and <italic>Steneosaurus hulkei</italic>. No evidence for this taxonomic revision was given. However, they considered <italic>M. mosae</italic> to be distinct from this long-lived (more than 10 Ma) <italic>M. hugii</italic> species. Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>, p. 194] criticized the content of their species diagnoses, as: ‘most of the content of these diagnoses reveal to be either diagnostic at the genus level or to characterize all Teleosauridae’. Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>, pp. 194–195] went on to show that the very high variation in maxillary and dentary tooth counts among the various Callovian teleosaurids is: ‘sufficient difference to discard such an interpretation (the synonymy)’. We concur with this assessment. Below, we propose a revised diagnosis for <italic>Machimosaurus</italic>, which has numerous autapomorphies absent in these Callovian species. Moreover, multiple phylogenetic analyses falsify the synonymy of <italic>M. hugii</italic> with <italic>S. durobrivensis</italic> and/or ‘<italic>S</italic>.’ <italic>obtusidens</italic> [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>,<xref rid="RSOS140222C41" ref-type="bibr">41</xref>,<xref rid="RSOS140222C61" ref-type="bibr">61</xref>].</p></sec><sec id="s3b"><label>4.2</label><title>The diverse Kimmeridgian <italic>Machimosaurus hugii</italic></title><p>Recently, Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] described an incomplete skeleton of <italic>Machimosaurus</italic> from the Lower Kimmeridgian of Germany. They referred this specimen to <italic>M. hugii</italic>, along with all other Kimmeridgian <italic>Machimosaurus</italic> specimens from Europe. This included synonymizing <italic>M. mosae</italic> with <italic>M. hugii</italic>. However, they did not discount that a second taxon could be determined based upon further investigation of relevant specimens [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>, p. 193].</p><p>This paper reopened an old debate about whether <italic>Machimosaurus</italic> is a monotypic genus, and whether the differences between <italic>M. hugii</italic> and <italic>M. mosae</italic> are due to ontogeny. This issue has been examined in detail by Hua [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>] and Vignaud (1995, unpublished PhD thesis), who studied teleosaurid ontogeny using the numerous European skulls available. Both of these authors considered <italic>M. mosae</italic> to be taxonomically distinct from <italic>M. hugii</italic>. We reject the hypothesis that the <italic>M. mosae</italic> neotype is a juvenile of <italic>M. hugii</italic> below, based on four fundamental flaws in this ‘juvenile hypothesis’: (i) the <italic>M. mosae</italic> neotype is comparable in size to the French [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>] and German [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] skulls referred to <italic>M. hugii</italic>, all three of which differ in length by only 7 cm (93–100 cm; <xref ref-type="table" rid="RSOS140222TB1">table 1</xref>); (ii) the lack of juvenile characteristics in any of the French [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>,<xref rid="RSOS140222C4" ref-type="bibr">4</xref>] and German [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] skulls [<xref rid="RSOS140222C58" ref-type="bibr">58</xref>]; (iii) the <italic>M. mosae</italic> neotype has exostoses (formation of new bone on the surface of bones, usually seen in mature individuals) in the femur, right pubis and on the transverse processes of some caudal vertebrae [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>]; and (iv) the <italic>M. mosae</italic> neotype is from the uppermost Kimmeridgian, whereas the two skulls referred to <italic>M. hugii</italic> are from the Lower Kimmeridgian, a temporal gap of some 3–5 million years.
<table-wrap id="RSOS140222TB1" orientation="portrait" position="float"><label>Table 1.</label><caption><p>Comparison of biometric variation among well-preserved <italic>Machimosaurus</italic> specimens.</p></caption><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col></colgroup><thead valign="bottom"><tr><th align="left" rowspan="1" colspan="1">species</th><th align="left" colspan="2" rowspan="1"><italic>M. buffetauti</italic><hr></hr></th><th align="left" colspan="2" rowspan="1"><italic>M. mosae</italic><hr></hr></th><th align="left" rowspan="1" colspan="1"><italic>M. hugii</italic></th></tr><tr><th align="left" rowspan="1" colspan="1">reference</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C10" ref-type="bibr">10</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C3" ref-type="bibr">3</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C62" ref-type="bibr">62</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C4" ref-type="bibr">4</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C6" ref-type="bibr">6</xref>]</th></tr><tr><th align="left" rowspan="1" colspan="1">basicranial length (cm)</th><th align="left" rowspan="1" colspan="1">93.5</th><th align="left" rowspan="1" colspan="1">100</th><th align="left" rowspan="1" colspan="1">approx. 130</th><th align="left" rowspan="1" colspan="1">96.5</th><th align="left" rowspan="1" colspan="1">approx. 149</th></tr></thead><tbody><tr><td rowspan="1" colspan="1">rostrum length (cm)</td><td rowspan="1" colspan="1">54.7</td><td rowspan="1" colspan="1">58</td><td rowspan="1" colspan="1">72<sup><italic>b</italic></sup></td><td rowspan="1" colspan="1">56.2</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">ratio of rostrum length to basicranial length (%)</td><td rowspan="1" colspan="1">58.5</td><td rowspan="1" colspan="1">58</td><td rowspan="1" colspan="1">55<sup><italic>c</italic></sup></td><td rowspan="1" colspan="1">58.2</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">maximum width of the skull (cm)</td><td rowspan="1" colspan="1">39.7</td><td rowspan="1" colspan="1">33<sup><italic>a</italic></sup></td><td rowspan="1" colspan="1">58</td><td rowspan="1" colspan="1">43</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">ratio of maximum skull width to basicranial length (%)</td><td rowspan="1" colspan="1">42.5</td><td rowspan="1" colspan="1">33<sup><italic>a</italic></sup></td><td rowspan="1" colspan="1">44.6</td><td rowspan="1" colspan="1">44.6</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">maximum supratemporal fenestra length (cm)</td><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1">27.5</td><td rowspan="1" colspan="1">∼40</td><td rowspan="1" colspan="1">32.2</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">ratio of maximum supratemporal fossa length to basicranial length (%)</td><td rowspan="1" colspan="1">27.8</td><td rowspan="1" colspan="1">27.5</td><td rowspan="1" colspan="1">30.8</td><td rowspan="1" colspan="1">33.4</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">length of mandible (cm)</td><td rowspan="1" colspan="1">95.4</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">132.5<sup><italic>b</italic></sup></td><td rowspan="1" colspan="1">112</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">length of mandibular symphysis (cm)</td><td rowspan="1" colspan="1">48.6</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">62<sup><italic>b</italic></sup></td><td rowspan="1" colspan="1">47.5</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">ratio of symphysis length to mandible length (%)</td><td rowspan="1" colspan="1">50.9</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">46.8<sup><italic>b</italic></sup></td><td rowspan="1" colspan="1">42.4</td><td rowspan="1" colspan="1">?</td></tr></tbody></table><table-wrap-foot><fn><p><sup><italic>a</italic></sup>Much of the skull has been reconstructed with plaster, making it difficult to discern what is bone and what is plaster ([<xref rid="RSOS140222C3" ref-type="bibr">3</xref>] and E. Buffetaut 2014, personal communication).</p></fn><fn><p><sup><italic>b</italic></sup>Estimate [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>].</p></fn><fn><p><sup><italic>c</italic></sup>The basicranial and rostrum length estimates of Sauvage & Liénard [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>] are most likely slight underestimates. From the skull line drawing in plate 1 fig. 1, it looks like the rostrum is too close to the orbital region of the skull. As such, the skull was probably mesorostrine (ratio of rostrum length to basicranial length would have been slightly more than 55%).
</p></fn></table-wrap-foot></table-wrap></p><p>Moreover, there is another character that shows that <italic>M. mosae</italic> is a distinct taxon from <italic>M</italic>. <italic>hugii</italic>, the presence of the prearticular (<xref ref-type="fig" rid="RSOS140222F25">figure 25</xref>). Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] described the first prearticular ever mentioned for a teleosaurid (although they did not highlight its significance). Prearticular bones were previously only known in Metriorhynchidae among thalattosuchians [<xref rid="RSOS140222C1" ref-type="bibr">1</xref>]. Interestingly, and most importantly for <italic>Machimosaurus</italic> systematics, the prearticulars are not found in the mandible of <italic>M. mosae</italic> [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>]. Therefore, the loss of the prearticulars is a specific character for <italic>M. mosae</italic> (as the prearticulars are also present in <italic>S. larteti</italic>; <xref ref-type="fig" rid="RSOS140222F1">figure 1</xref><italic>a</italic>,<italic>b</italic>). (Owing to the status of the <italic>M. mosae</italic> neotype, we cannot currently test the hypothesis whether the absence of the prearticulars is a preservational artefact.)</p><p>Postcranial characteristics supporting the distinction of <italic>M</italic>. <italic>hugii</italic> and <italic>M</italic>. <italic>mosae</italic>—not examined by Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], who unfortunately mainly focused on the skull—include:
<list list-type="simple"><list-item><p>— Coracoids are highly variable among teleosaurid species [<xref rid="RSOS140222C1" ref-type="bibr">1</xref>]. The same is true between the new specimen of <italic>M</italic>. <italic>hugii</italic> and <italic>M. mosae</italic>, especially in the shape and size of the postglenoid and glenoid processes (<xref ref-type="fig" rid="RSOS140222F36">figure 36</xref><italic>a</italic>,<italic>b</italic>). In the German <italic>M</italic>. <italic>hugii</italic> skull [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], the coracoid glenoid process (process near the glenoid fossa that projects posterodorsally) is elongate, extending considerably from the head of the coracoid, and is a sub-isosceles triangle in shape when seen in lateral view; the coracoid postglenoid process anterior margin is very slightly concave and terminates approximately in the samef frontal plane as the glenoid; and the postglenoid process posterior margin is strongly concave and terminates approximately in the same frontal plane as the posterior-end of the glenoid process. However in the <italic>M. mosae</italic> neotype [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>], the coracoid glenoid process is very short, not extending far from the head of the coracoid, and is a right-angled triangle in shape when seen in lateral view; the coracoid postglenoid process anterior margin is strongly concave and terminates in a frontal plane anterior to the glenoid; and the postglenoid process posterior margin is strongly concave distally but shifts to being somewhat convex proximally and terminates in a frontal plane posterior to the posterior-end of the glenoid process.</p></list-item><list-item><p>— The difference in axis neural arch shape between their new specimen of <italic>M</italic>. <italic>hugii</italic> and <italic>M. mosae</italic> (<xref ref-type="fig" rid="RSOS140222F36">figure 36</xref><italic>g</italic>,<italic>h</italic>). In the German <italic>M. hugii</italic> specimen [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], the axis neural arch has a strongly concave dorsal margin and the postzygapophyses terminate notably posterior to the posterior surface of the centrum (<xref ref-type="fig" rid="RSOS140222F36">figure 36</xref><italic>h</italic>), whereas in the <italic>M. mosae</italic> neotype [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>] the dorsal margin is only weakly concave and the postzygapophyses are not as long posteriorly (<xref ref-type="fig" rid="RSOS140222F36">figure 36</xref><italic>g</italic>). Compare Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>, p. 191, fig. 9<italic>a</italic>–<italic>c</italic>] with Hua [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>, plate 3 fig. 1–3].</p></list-item></list><fig id="RSOS140222F36" orientation="portrait" position="float"><label>Figure 36.</label><caption><p>Postcranial element comparison between the holotype of <italic>M. buffetauti</italic> and the neotype of <italic>M. mosae</italic> (based on the figures in [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C10" ref-type="bibr">10</xref>]). Coracoids of (<italic>a</italic>) <italic>M. mosae</italic> and (<italic>b</italic>) <italic>M</italic>. <italic>buffetauti</italic>; the dorsal osteoderms of (<italic>c</italic>,<italic>e</italic>) <italic>M</italic>. <italic>mosae</italic> and (<italic>d</italic>,<italic>f</italic>) <italic>M</italic>. <italic>buffetauti</italic>; the atlas–axis of (<italic>g</italic>) <italic>M. mosae</italic> and (<italic>h</italic>) <italic>M. buffetauti</italic>.</p></caption><graphic xlink:href="rsos140222-g36"></graphic></fig></p><p>Thanks to the new specimen described by Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] there are now numerous postcranial characteristics, along with the absence of the prearticular (assuming that it is not a preservational artefact), that allow easy identification of <italic>M. mosae</italic> and differentiate it from <italic>M</italic>. <italic>hugii</italic>. Furthermore, they described a depression on the dorsal surface of the quadrates near the hemicondyles on the German <italic>M. hugii</italic> skull. These depressions are not seen in <italic>M. mosae</italic> and comprise another feature differentiating these two species [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>].</p></sec></sec><sec id="s4"><label>5.</label><title>Description of <italic>Machimosaurus hugii</italic> by von Huene [<xref rid="RSOS140222C63" ref-type="bibr">63</xref>] and Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>]</title><p><italic>Machimosaurus</italic> teeth have long been known, particularly from the Kimmeridgian of Solothurn, Switzerland. One such tooth was figured by Cuvier in 1824 [<xref rid="RSOS140222C64" ref-type="bibr">64</xref>, plate 6 fig. 7]. In 1836, Römer [<xref rid="RSOS140222C65" ref-type="bibr">65</xref>] figured a <italic>Machimosaurus</italic> tooth from Kahlenberg, Germany, although he considered it to be <italic>Ichthyosaurus</italic> [<xref rid="RSOS140222C65" ref-type="bibr">65</xref>, p. 12, plate 12 fig. 19]. It was not until 1837 that the binomial <italic>M. hugii</italic> was specifically established for the Solothurn and Kahlenberg teeth [<xref rid="RSOS140222C31" ref-type="bibr">31</xref>]. Unfortunately, the name was misspelt as <italic>Madrimosaurus hugii</italic> in that publication [<xref rid="RSOS140222C31" ref-type="bibr">31</xref>], something von Meyer attributed to: ‘Die Undeutlichkeit meiner Handschrift’—‘the indistinctness of my handwriting’ [<xref rid="RSOS140222C66" ref-type="bibr">66</xref>, p. 415]. As such, von Meyer corrected the spelling in an 1838 publication [<xref rid="RSOS140222C66" ref-type="bibr">66</xref>].</p><p>Curiously, throughout all the various competing arguments over specimen synonymies regarding <italic>M. hugii</italic>, the description and figures of von Huene [<xref rid="RSOS140222C63" ref-type="bibr">63</xref>] and Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>] (as well as the Solothurn teeth) depict a taxon which is distinct from the <italic>M. mosae</italic> and the ‘<italic>M. hugii</italic>’ specimens described during the 1980s—2010s from France and Germany. This taxon is known from Swiss and Portuguese material. The distinctiveness of this taxon relative to other <italic>Machimosaurus</italic> specimens has not been clearly recognized until now. As we show below, this taxon is <italic>M</italic>. <italic>hugii</italic>.</p><p>von Huene [<xref rid="RSOS140222C63" ref-type="bibr">63</xref>] described and figured a number of fragmentary skull and mandibular fragments from Switzerland that belong to this taxon (NMS 7012, NMS 7015 and NMS 7021; <xref ref-type="fig" rid="RSOS140222F37">figures 37</xref>–<xref ref-type="fig" rid="RSOS140222F40">40</xref>). His figures demonstrate five autapomorphies among <italic>Machimosaurus</italic> (and other teleosaurids): (i) the external surfaces of the snout bones are poorly ornamented with low relief ridges mostly orientated anteroposterly; (ii) sub-globidont dentition (blunt apices, low apicobasal height to basal width ratio, but the teeth lack the pronounced ‘globular’/bulbous morphology of true globidonty); (iii) apicobasally aligned enamel ridges immediately adjacent to the apical anastomosed region that are closely packed on both the labial and lingual tooth surfaces; (iv) uniform inter-alveolar spaces in the posterior–mid region of the maxillae, with the inter-alveolar spaces becoming slightly larger anteriorly but still being largely uniform in size; and (v) uniformly narrow inter-alveolar spaces in the dentaries. These characteristics are distinct from the morphologies seen in the <italic>M. mosae</italic> neotype [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C21" ref-type="bibr">21</xref>] and the French and German ‘<italic>M</italic>. <italic>hugii</italic>’ specimens [<xref rid="RSOS140222C2" ref-type="bibr">2</xref>,<xref rid="RSOS140222C3" ref-type="bibr">3</xref>,<xref rid="RSOS140222C10" ref-type="bibr">10</xref>,<xref rid="RSOS140222C17" ref-type="bibr">17</xref>] (<xref ref-type="fig" rid="RSOS140222F12">figures 12</xref>–<xref ref-type="fig" rid="RSOS140222F14">14</xref> and <xref ref-type="fig" rid="RSOS140222F21">21</xref>–<xref ref-type="fig" rid="RSOS140222F27">27</xref>), in which: (i) the external surfaces of the snout bones are more strongly ornamented, with higher relief ridges and sub-circular/oval pits; (ii) no tooth crowns are sub-globidont; (iii) apicobasally aligned enamel ridges immediately adjacent to the apical anastomosed region are closely packed only on the lingual tooth surface, but on the labial surface these ridges are more widely spaced; (iv) the maxillary and (v) dentary inter-alveolar spaces are variable in size, some of which can be quite large proportionaly to the adjacent alveoli.
<fig id="RSOS140222F37" orientation="portrait" position="float"><label>Figure 37.</label><caption><p><italic>Machimosaurus hugii</italic>, NMS 7012, referred specimen. Incomplete snout (fragment consisting of the lacrimals, nasals and maxilla), (<italic>a</italic>) photograph in dorsal view, (<italic>b</italic>) line drawing in dorsal view, (<italic>c</italic>) photograph in ventral view and (<italic>d</italic>) line drawing in ventral view. lac, lacrimal; mx, maxilla; na, nasals.</p></caption><graphic xlink:href="rsos140222-g37"></graphic></fig><fig id="RSOS140222F38" orientation="portrait" position="float"><label>Figure 38.</label><caption><p><italic>Machimosaurus hugii</italic>, NMS 7012, referred specimen. Incomplete snout (fragment consisting of the maxilla and premaxilla), (<italic>a</italic>) photograph in dorsal view, (<italic>b</italic>) line drawing in dorsal view, (<italic>c</italic>) photograph in ventral view and (<italic>d</italic>) line drawing in ventral view. mx, maxilla; pmx, premaxilla.</p></caption><graphic xlink:href="rsos140222-g38"></graphic></fig><fig id="RSOS140222F39" orientation="portrait" position="float"><label>Figure 39.</label><caption><p><italic>Machimosaurus hugii</italic>, NMS 7015, referred specimen. Incomplete snout (fragment consisting of the maxilla and palatines, damage makes determining other bones difficult), (<italic>a</italic>) photograph in dorsal view, (<italic>b</italic>) line drawing in dorsal view, (<italic>c</italic>) photograph in ventral view and (<italic>d</italic>) line drawing in ventral view. mx, maxilla; pal, palatines.</p></caption><graphic xlink:href="rsos140222-g39"></graphic></fig><fig id="RSOS140222F40" orientation="portrait" position="float"><label>Figure 40.</label><caption><p><italic>Machimosaurus hugii</italic>, NMS 7021, referred specimen. Incomplete lower jaw, (<italic>a</italic>) photograph in dorsal view, (<italic>b</italic>) line drawing in dorsal view, (<italic>c</italic>) photograph in ventral view and (<italic>d</italic>) line drawing in ventral view. den, dentary; sp, splenial.</p></caption><graphic xlink:href="rsos140222-g40"></graphic></fig></p><p>Krebs' [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>] description and figures of the large, but fragmentary, Leiria skull from Portugal (MG-8730-1 and MG-8730-2; <xref ref-type="fig" rid="RSOS140222F4">figures 4</xref>–<xref ref-type="fig" rid="RSOS140222F9">9</xref>) reveal seven autapomorphies: (i) the external surfaces of the snout bones are poorly ornamented with low relief ridges mostly orientated anteroposterly; (ii) sub-globidont dentition; (iii) apicobasally aligned enamel ridges immediately adjacent to the apical anastomosed region that are closely packed on both the labial and lingual tooth surfaces; (iv) paroccipital processes that are greatly enlarged, both elongated mediolaterally and with lateral ends that are expanded; (v) basioccipital tuberosities (basal tubera) that are very large in size and are sub-rectangular in shape when seen in occipital view; (vi) the inter-basioccipital tubera notch is a large inverse ‘U’-shape when seen in occipital view; and (vii) uniformly spaced inter-alveolar spaces in the mid region of the maxillae. Moreover, the apicobasal ridge characteristic is also seen in the lectotype of <italic>M. hugii</italic> [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>]. Once again, these characteristics are distinct from the morphologies seen in the <italic>M. mosae</italic> neotype [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C21" ref-type="bibr">21</xref>] and the French and German ‘<italic>M</italic>. <italic>hugii</italic>’ specimens [<xref rid="RSOS140222C2" ref-type="bibr">2</xref>,<xref rid="RSOS140222C3" ref-type="bibr">3</xref>,<xref rid="RSOS140222C10" ref-type="bibr">10</xref>,<xref rid="RSOS140222C17" ref-type="bibr">17</xref>].</p><p>Even though these Swiss and Portuguese specimens are fragmentary, they share four autapomorphies that are not seen in any other teleosaurid: (i) the external surfaces of the snout bones are poorly ornamented with low relief ridges mostly orientated anteroposterly; (ii) sub-globidont dentition; (iii) apicobasally aligned enamel ridges immediately adjacent to the apical anastomosed region that are closely packed on both the labial and lingual tooth surfaces; and (iv) uniform inter-alveolar spaces in the maxillae that are proportionally narrow relative to alveoli. None of these characteristics are found in the French or German specimens (<xref ref-type="table" rid="RSOS140222TB2">table 2</xref>). They indicate that the Swiss and Portuguese material belongs to the same diagnostic taxon.
<table-wrap id="RSOS140222TB2" orientation="portrait" position="float"><label>Table 2.</label><caption><p>Comparison of dental morphologies and alveolar counts among <italic>Machimosaurus</italic> specimens.</p></caption><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col></colgroup><thead valign="bottom"><tr><th align="left" rowspan="1" colspan="1">species</th><th align="left" colspan="3" rowspan="1"><italic>M. buffetauti</italic><hr></hr></th><th align="left" colspan="2" rowspan="1"><italic>M. mosae</italic><hr></hr></th><th align="left" colspan="2" rowspan="1"><italic>M. hugii</italic><hr></hr></th></tr><tr><th align="left" rowspan="1" colspan="1">reference</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C17" ref-type="bibr">17</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C10" ref-type="bibr">10</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C3" ref-type="bibr">3</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C62" ref-type="bibr">62</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C4" ref-type="bibr">4</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C6" ref-type="bibr">6</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C63" ref-type="bibr">63</xref>]</th></tr><tr><th align="left" rowspan="1" colspan="1">premaxillary alveoli count</th><th align="left" rowspan="1" colspan="1">3</th><th align="left" rowspan="1" colspan="1">3</th><th align="left" rowspan="1" colspan="1">3</th><th align="left" rowspan="1" colspan="1">3</th><th align="left" rowspan="1" colspan="1">3</th><th align="left" rowspan="1" colspan="1">?</th><th align="left" rowspan="1" colspan="1">?</th></tr></thead><tbody><tr><td rowspan="1" colspan="1">maxillary alveoli count</td><td rowspan="1" colspan="1">>18</td><td rowspan="1" colspan="1">22<sup><italic>a</italic></sup></td><td rowspan="1" colspan="1">21<sup><italic>a</italic></sup></td><td rowspan="1" colspan="1">probably 17 or 18<sup><italic>b</italic></sup></td><td rowspan="1" colspan="1">probably 18 or 19<sup><italic>d</italic></sup></td><td rowspan="1" colspan="1">at least 18</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">maxillary alveoli anterior to palatines</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?16–17</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?7<sup><italic>c</italic></sup></td><td rowspan="1" colspan="1">14?</td><td rowspan="1" colspan="1">at least 12</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">inter-alveolar spaces between the maxillary alveoli</td><td rowspan="1" colspan="1">variable in size, some large</td><td rowspan="1" colspan="1">variable in size, some large</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">variable in size, some large</td><td rowspan="1" colspan="1">variable in size, some large</td><td rowspan="1" colspan="1">posterior–mid inter-alveolar spaces uniform in size</td><td rowspan="1" colspan="1">posterior–mid inter-alveolar spaces uniform in size. Anterior spaces larger but still uniform in size</td></tr><tr><td rowspan="1" colspan="1">dentary alveoli count</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">21/22</td><td rowspan="1" colspan="1">approx. 24/25<sup><italic>a</italic></sup></td><td rowspan="1" colspan="1">probably 18<sup><italic>b</italic></sup></td><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">>15</td></tr><tr><td rowspan="1" colspan="1">dentary alveoli anterior to the splenial</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">>8</td></tr><tr><td rowspan="1" colspan="1">dentary alveoli adjacent to mandibular symphysis</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">19–20</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">15–16</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">>13</td></tr><tr><td rowspan="1" colspan="1">diastema between fourth and fifth dentary alveoli</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">yes</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">yes</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">inter-alveolar spaces between the dentary alveoli</td><td rowspan="1" colspan="1">variable in size, some large</td><td rowspan="1" colspan="1">variable in size, some large</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">variable in size, some large</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">mostly uniform in size, and small</td></tr><tr><td rowspan="1" colspan="1">sub-globidont dentition</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">yes</td><td rowspan="1" colspan="1">yes</td></tr><tr><td rowspan="1" colspan="1">apicobasal ridges immediately adjacent to the apical anastomosed region: closely packed on both the labial and lingual surfaces</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">yes</td><td rowspan="1" colspan="1">yes</td></tr></tbody></table><table-wrap-foot><fn><p><sup><italic>a</italic></sup>Estimate [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>].</p></fn><fn><p><sup><italic>b</italic></sup>We estimate there to be 17 or 18 maxillary alveoli, and most likely a similar number of dentary alveoli. This is higher than the 16 maxillary and dentary alveoli estimate of Sauvage & Liénard [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>].</p></fn><fn><p><sup><italic>c</italic></sup>The seven pre-palatine maxillary count is based on the figure in Sauvage & Liénard [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>]. However, this count assumes that the reconstruction is accurate, which it may not be. This is due to the unusual shape of the anterior process of the palatines and the breaks in the specimen.</p></fn><fn><p><sup><italic>d</italic></sup>Hua [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>] estimated there to be 17 maxillary alveoli. As the posterior maxillary alveoli are small in <italic>Machimosaurus</italic> [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], a maxillary tooth count of 18 or 19 is likely.</p></fn></table-wrap-foot></table-wrap></p></sec><sec id="s5"><label>6.</label><title><italic>Steneosaurus bouchardi</italic> skull of von Huene [<xref rid="RSOS140222C63" ref-type="bibr">63</xref>]</title><p>von Huene [<xref rid="RSOS140222C63" ref-type="bibr">63</xref>] also described an incomplete skull and mandible from Solothurn and referred them to <italic>Steneosaurus bouchardi</italic>, an Upper Kimmeridgian species of longirostrine teleosaurid. The skull (NMS 7049; <xref ref-type="fig" rid="RSOS140222F41">figures 41</xref>–<xref ref-type="fig" rid="RSOS140222F43">43</xref>) lacks most of the snout, right supratemporal arch, both quadrates and the palatal surface is poorly preserved. Its specimen labels show that ‘Zangerl, Chicago’ referred this skull to <italic>M. hugii</italic> in 1947.
<fig id="RSOS140222F41" orientation="portrait" position="float"><label>Figure 41.</label><caption><p><italic>Machimosaurus hugii</italic>, NMS 7029, referred specimen. Incomplete skull (orbital and temporal region) in dorsal view, (<italic>a</italic>) photograph and (<italic>b</italic>) line drawing. bo, basioccipital; fr, frontal; or, orbit; p, parietal; po, postorbital.</p></caption><graphic xlink:href="rsos140222-g41"></graphic></fig><fig id="RSOS140222F42" orientation="portrait" position="float"><label>Figure 42.</label><caption><p><italic>Machimosaurus hugii</italic>, NMS 7029, referred specimen. Incomplete skull (orbital and temporal region) in ventral view, (<italic>a</italic>) photograph and (<italic>b</italic>) line drawing. bo, basioccipital; bs, basisphenoid; max, maxilla; pal, palatine; pt, pterygoid; sof, suborbital fenestra.</p></caption><graphic xlink:href="rsos140222-g42"></graphic></fig><fig id="RSOS140222F43" orientation="portrait" position="float"><label>Figure 43.</label><caption><p><italic>Machimosaurus hugii</italic>, NMS 7029, referred specimen. Incomplete skull (orbital and temporal region) in occipital/posterior view, (<italic>a</italic>) photograph and (<italic>b</italic>) line drawing. bo, basioccipital; bt, basioccipital tuberosities; fr, frontal; po, postorbital; sq, squamosal.</p></caption><graphic xlink:href="rsos140222-g43"></graphic></fig></p><p>The Solothurn skull (NMS 7049) shares three autapomorphies with the braincase of the Leiria skull (MG-8730-2): (i) paroccipital processes are greatly enlarged, both elongated mediolaterally and with lateral ends that are expanded; (ii) basioccipital tuberosities (basal tubera) are very large in size and are sub-rectangular in shape when seen in occipital view; and (iii) the inter-basioccipital tubera notch is a large inverse ‘U’-shape when seen in occipital view. Moreover, like the other Swiss specimens and the Portuguese specimen, the external surfaces of the skull bones are poorly ornamented. These features support the referral of NMS 7049 to the same taxon as the Swiss and Portuguese material described above. As we show below that this taxon is <italic>M</italic>. <italic>hugii</italic>, we therefore conclude that Zangerl's referral of NMS 7049 to <italic>M</italic>. <italic>hugii</italic> is correct.</p></sec><sec id="s6"><label>7.</label><title>Systematic palaeontology</title><p>Crocodylomorpha Hay, 1930 [<xref rid="RSOS140222C67" ref-type="bibr">67</xref>]</p><p>Thalattosuchia Fraas, 1901 [<xref rid="RSOS140222C68" ref-type="bibr">68</xref>]</p><p>Teleosauridae Geoffroy, 1831 [<xref rid="RSOS140222C69" ref-type="bibr">69</xref>]</p><p><italic>Machimosaurus</italic> von Meyer, 1837 [<xref rid="RSOS140222C31" ref-type="bibr">31</xref>] emend. von Meyer, 1838 [<xref rid="RSOS140222C66" ref-type="bibr">66</xref>].</p><sec id="s6a"><label>7.1</label><title>Type species</title><p><italic>Machimosaurus hugii</italic> von Meyer, 1837 [<xref rid="RSOS140222C31" ref-type="bibr">31</xref>] emend. von Meyer, 1838 [<xref rid="RSOS140222C66" ref-type="bibr">66</xref>].</p></sec><sec id="s6b"><label>7.2</label><title>Referred species</title><p><italic>Machimosaurus buffetauti</italic> sp. nov., <italic>M. mosae</italic> Sauvage & Liénard, 1879 [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>] and <italic>Machimosaurus nowackianus</italic> (von Huene, 1938 [<xref rid="RSOS140222C19" ref-type="bibr">19</xref>]) comb. nov.</p></sec><sec id="s6c"><label>7.3</label><title>Etymology</title><p>‘Pugnacious lizard’. <italic>Machimo</italic> is derived from the Greek word <italic>machimoi</italic> (<inline-formula><mml:math id="IM1"><mml:mi>μ</mml:mi><mml:mrow><mml:mover><mml:mi>α</mml:mi><mml:mo>´</mml:mo></mml:mover></mml:mrow><mml:mi>χ</mml:mi><mml:mi>ι</mml:mi><mml:mi>μ</mml:mi><mml:mi>o</mml:mi><mml:mi>ι</mml:mi></mml:math></inline-formula>), meaning pugnacious (Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>] translated it into German as streitbar). In the Hellenic world, <italic>machimoi</italic> was used to describe non-Greek armies, especially native Egyptian troops during the Ptolemaic Dynasties.<italic>Saurus</italic> is the Latinized form of <italic>sauros</italic> (<italic>σαυρoς</italic>), the Ancient Greek for lizard.</p></sec><sec id="s6d"><label>7.4</label><title>Geological range</title><p>Middle Oxfordian to Lower Tithonian. (As noted above, we cannot confirm the presence of <italic>Machimosaurus</italic> in deposits younger than the Lower Tithonian.)</p></sec><sec id="s6e"><label>7.5</label><title>Geographical range</title><p>Africa (Ethiopia) and Europe (England, France, Germany, Portugal, Spain and Switzerland).</p></sec><sec id="s6f"><label>7.6</label><title>Generic diagnosis</title><p>Teleosaurid crocodylomorphs with the following unique combination of characters (autapomorphic characters among teleosaurids are indicated by an asterisk *): large body size (basicranial length typically 90–110 cm, but can exceed 140 cm)*; three alveoli per premaxilla (shared with <italic>Peipehsuchus teleorhinus</italic>); the first premaxillary alveoli are orientated strongly anteroventrally*; 18–22 alveoli per maxilla*; 19–25 alveoli per dentary*; vertically orientated, interlocking dentition, with pronounced reception pits at the premaxillary, maxillary and dentary inter-alveolar spaces*; conical teeth with blunt/rounded apices (shared with ‘<italic>S</italic>.’ <italic>obtusidens</italic>); tooth enamel ornamentation varies along the crown, in the basal region enamel ornamentation is composed of numerous apicobasally aligned ridges of high relief, which become an anastomosed pattern in the apical region (shared with ‘<italic>S</italic>.’ <italic>obtusidens</italic>); presence of carinae is variable, in anterior teeth they can be prominent but in shorter blunter teeth carinae are either very poorly developed or absent*; ratio of crown apicobasal height to basal transverse width can be as low as 1.5 in the posterior teeth; rostrum is broad and mesorostrine, constituting less than 60% of basicranial length*; antorbital fenestrae are absent (possibly shared with <italic>Steneosaurus heberti</italic>); supratemporal fossae are parallelogram in shape (shared with ‘<italic>S</italic>.’ <italic>obtusidens</italic>); ratio of maximum supratemporal fossa length to basicranial length is greater than 27%*; three sacral vertebrae (possibly shared with ‘<italic>S</italic>.’ <italic>obtusidens</italic>)*; medial quadrate hemicondyle is considerably smaller than the lateral hemicondyle*; exoccipital is excluded from the occipital condyle (composed solely of the basioccipital) (shared with <italic>S. heberti</italic>); axis neural spine is tall in lateral view, terminating in a transverse plane that is notably dorsal to the plane of the pre- and postzygapophyses*; axis neural spine posteriorly expanded when seen in lateral view, with the posterior margin terminating approximately in the same frontal plane as the posterior margin of the postzygapophyses*.</p></sec><sec id="s6g"><label>7.7</label><title><italic>Machimosaurus</italic> sp.</title><sec id="s6g1"><label>7.7.1</label><title>Specimens</title><p>MG23—partial maxilla (Malhão, Algarve, south Portugal; Oxfordian) [<xref rid="RSOS140222C18" ref-type="bibr">18</xref>].</p><p>ML1208—isolated tooth (Middle Oxfordian of Cesaredas, central west Portugal).</p><p>Musée de la Princerie (Verdun, France) 2007.0.14—incomplete lower jaw and isolated tooth crowns (Upper Oxfordian, <italic>Perisphinctes</italic> variocostatus subzone of the <italic>P. cautisnigrae</italic> N–W European ammonite zone. From Haudainville, near Verdun, Département de la Meuse, Lorraine, France) (<xref ref-type="fig" rid="RSOS140222F2">figure 2</xref>) [<xref rid="RSOS140222C12" ref-type="bibr">12</xref>].</p><p>NHMUK PV R36793—isolated tooth (Upper Oxfordian of Villerville, Département du Calvados, Basse-Normandie, France; Calcaire gréseux d'Hennequeville Formation) [<xref rid="RSOS140222C17" ref-type="bibr">17</xref>].</p></sec><sec id="s6g2"><label>7.7.2</label><title>Geological range</title><p>Middle—Upper Oxfordian.</p></sec><sec id="s6g3"><label>7.7.3</label><title>Geographical range</title><p>Europe (France and Portugal).</p><p><italic>Note.</italic> The Oxfordian <italic>Machimosaurus</italic> material is taxonomically indeterminate.</p></sec></sec><sec id="s6h"><label>7.8</label><title><italic>Machimosaurus hugii</italic> von Meyer, 1837 [<xref rid="RSOS140222C31" ref-type="bibr">31</xref>] emend von Meyer, 1838 [<xref rid="RSOS140222C66" ref-type="bibr">66</xref>]</title><p><table-wrap orientation="portrait" id="d35e3770" position="anchor"><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col></colgroup><tbody><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1824</td><td rowspan="1" colspan="1">Dent obtuse d'un crocodile du Jura, peut-être d'une espèce différence de la précédente [crocodile de Caen]—Cuvier [<xref rid="RSOS140222C64" ref-type="bibr">64</xref>, planche 6 fig. 7]</td></tr><tr><td rowspan="1" colspan="1">v*</td><td rowspan="1" colspan="1"><italic>1837</italic></td><td rowspan="1" colspan="1"><italic>Madrimosaurus hugii</italic> sp. nov.—von Meyer [<xref rid="RSOS140222C31" ref-type="bibr">31</xref>, p. 560]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1"><italic>1838</italic></td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—von Meyer [<xref rid="RSOS140222C66" ref-type="bibr">66</xref>, p. 415]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1"><italic>1888</italic></td><td rowspan="1" colspan="1"><italic>Machimosaurus hughi</italic> von Meyer—Lydekker [<xref rid="RSOS140222C70" ref-type="bibr">70</xref>, p. 103] (sic)</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1897–97</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Sauvage [<xref rid="RSOS140222C18" ref-type="bibr">18</xref>, p. 27, plate 3 figs 9–10 and plate 5 figs 6–7]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1925</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—von Huene [<xref rid="RSOS140222C63" ref-type="bibr">63</xref>, pp. 584–588, plate 25 all figures]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1925</td><td rowspan="1" colspan="1"><italic>Steneosaurus bouchardi</italic> Sauvage—von Huene [<xref rid="RSOS140222C63" ref-type="bibr">63</xref>, p. 589, plate 26 fig. 1<italic>a</italic>–<italic>c</italic>]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1943</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Teixeira [<xref rid="RSOS140222C22" ref-type="bibr">22</xref>, p. 109, fig. 1]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1967</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>, pp. 46–58, figs 1–4]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1968</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Krebs [<xref rid="RSOS140222C7" ref-type="bibr">7</xref>, pp. 21–53, figs 1–18]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1973</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Steel [<xref rid="RSOS140222C71" ref-type="bibr">71</xref>, pp. 25, 30, fig. 14 (8) (<italic>partim</italic>)]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1"><italic>2008</italic></td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Pierce <italic>et al</italic>. [<xref rid="RSOS140222C9" ref-type="bibr">9</xref>, p. 1085 (<italic>partim</italic>)]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">2010</td><td rowspan="1" colspan="1">cf. <italic>Machimosaurus</italic> sp.—Ruiz-Omeñaca <italic>et al</italic>. [<xref rid="RSOS140222C48" ref-type="bibr">48</xref>, pp. 81–81, fig. 1<italic>d</italic>]</td></tr></tbody></table></table-wrap></p><sec id="s6h1"><label>7.8.1</label><title>Lectotype</title><p>NMS 8342: isolated tooth crown (<xref ref-type="fig" rid="RSOS140222F28">figure 28</xref>). Previously catalogued as specimen number 96 [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>].</p></sec><sec id="s6h2"><label>7.8.2</label><title>‘Holotype’/syntypes</title><p>von Meyer [<xref rid="RSOS140222C31" ref-type="bibr">31</xref>,<xref rid="RSOS140222C66" ref-type="bibr">66</xref>] never designated a holotype for <italic>M. hugii</italic>. When establishing <italic>M. hugii</italic>, he referred isolated tooth crowns from Solothurn, Switzerland and Kahlenberg, Germany to the species (i.e. a type series or syntypes). Note that Steel [<xref rid="RSOS140222C71" ref-type="bibr">71</xref>] mistook Kahlenberg, Hannover as being Kahlenberg in Austria. There are, to our knowledge, no <italic>Machimosaurus</italic> specimens known from Austria.</p><p>Therefore, Pierce <italic>et al</italic>. [<xref rid="RSOS140222C9" ref-type="bibr">9</xref>] and Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] were incorrect in referring to a holotype for this taxon. They appear to have considered the lectotype as being the holotype, as the same specimen number is given (the old 96 number which Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>] used). Also, Pierce <italic>et al</italic>. [<xref rid="RSOS140222C9" ref-type="bibr">9</xref>] stated that the ‘holotype’ is from the Palaeontologische Sammlung im Museum der Stadt Solothurn, the former name for the Naturmuseum Solothurn (again, the old name which Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>] used). Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], however, listed the ‘holotype’ as being from the Staatliche Naturhistorische Sammlungen Dresden, which is the former name for the Senckenberg Naturhistorische Sammlungen Dresden. No reason is stated why they believed that the ‘holotype’ was moved to a different museum in a different country. Regardless, we can confirm that the lectotype tooth is still in Solothurn.</p></sec><sec id="s6h3"><label>7.8.3</label><title>‘Neotype’</title><p>More confusingly, Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>, p. 192] claimed that Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>] made the Portuguese specimen (MG-8730-1, MG-8730-2) the neotype of <italic>M. hugii</italic>. However, Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>]: (i) never refers to a neotype and (ii) clearly referred to the Swiss tooth (NMS 8342, then catalogued as specimen number 96) as the ‘lectotypus’ of <italic>M. hugii</italic>. Moreover, this is the earliest mention of a lectotype we can find for <italic>M. hugii</italic>, and Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>] may have designated it acting as first reviser. Moreover, in Krebs' later paper, he still refers to the isolated tooth crown as being the lectotype [<xref rid="RSOS140222C7" ref-type="bibr">7</xref>, p. 35 and figured on p. 37]. Other than in Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], we can find no reference to a <italic>M. hugii</italic> ‘neotype’.</p></sec><sec id="s6h4"><label>7.8.4</label><title>Lectotype locality</title><p>Kreuzen Quarry at St. Verena, near Solothurn, Canton Solothurn, Switzerland [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>]; 47° N, 7° E.</p></sec><sec id="s6h5"><label>7.8.5</label><title>Lectotype horizon</title><p>‘Rätschenbank’ der Schildkrötenschichten [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>] (= Solothurn Turtle Limestone, Reuchenette Formation). Uppermost Kimmeridgian, Upper Jurassic.</p></sec><sec id="s6h6"><label>7.8.6</label><title>Etymology</title><p>‘Hugi's pugnacious lizard’. Named in honour of Franz Joseph Hugi (1791–1855), the Swiss geologist and naturalist.</p></sec><sec id="s6h7"><label>7.8.7</label><title>Referred specimens</title><p>MG-8730-1, MG-8730-2 and unnumbered elements—incomplete skull and postcranial elements (Lower or Upper Kimmeridgian of Guimarota near Leiria, Portugal [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>]; <xref ref-type="fig" rid="RSOS140222F4">figures 4</xref>–<xref ref-type="fig" rid="RSOS140222F10">10</xref>).</p><p>ML491, ML495, ML959 and ML1955—isolated teeth (Upper Kimmeridgian of Porto das Barcas, Lourinhã, Portugal; Praia Azul Member of the Lourinhã Formation; <xref ref-type="fig" rid="RSOS140222F33">figures 33</xref><italic>a</italic> and <xref ref-type="fig" rid="RSOS140222F35">35</xref>).</p><p>ML647—isolated tooth crown (Upper Kimmeridgian of Peralta, Lourinhã, Portugal; Praia Azul Member of the Lourinhã Formation; <xref ref-type="fig" rid="RSOS140222F34">figure 34</xref>).</p><p>ML657 and ML658—isolated teeth (Upper Kimmeridgian of Zimbral, Lourinhã, Portugal; Praia Azul Member of the Lourinhã Formation; <xref ref-type="fig" rid="RSOS140222F33">figure 33</xref><italic>b</italic>–<italic>d</italic>).</p><p>ML647, ML733 and ML902—isolated teeth (Upper Kimmeridgian of beach near Lourinhã, Portugal; Praia Azul Member of the Lourinhã Formation).</p><p>MUJA-1008 and MUJA-1922—isolated teeth (Kimmeridgian of Playa de La Griega, Colunga, Asturias, Spain; Tereñes Formation; <xref ref-type="fig" rid="RSOS140222F11">figure 11</xref><italic>c</italic>,<italic>d</italic>) [<xref rid="RSOS140222C25" ref-type="bibr">25</xref>].</p><p>MUJA-1298—isolated tooth crown (Kimmeridgian of La Escalera, Villaviciosa, Asturias, Spain; Lastres Formation; <xref ref-type="fig" rid="RSOS140222F11">figure 11</xref><italic>a</italic>–<italic>d</italic>) [<xref rid="RSOS140222C25" ref-type="bibr">25</xref>].</p><p>MCNV-CC-4—isolated tooth crown (Tithonian of Cantera Carcalín near Buñol, Valencia Province, Spain [<xref rid="RSOS140222C48" ref-type="bibr">48</xref>]).</p><p>From the Kimmeridgian of Solothurn, Switzerland: NHMUK PV OR33239, NHMUK PV OR43638, NHMUK PV R5, NHMUK PV R232—isolated tooth crowns (<xref ref-type="fig" rid="RSOS140222F29">figure 29</xref>). NMS 7012—incomplete snout (<xref ref-type="fig" rid="RSOS140222F37">figures 37</xref> and <xref ref-type="fig" rid="RSOS140222F38">38</xref>). NMS 7015—incomplete snout (maxilla-palatine fragment; <xref ref-type="fig" rid="RSOS140222F39">figure 39</xref>). NMS 7021—incomplete mandible (<xref ref-type="fig" rid="RSOS140222F40">figure 40</xref>). NMS 7029—temporal and orbital region of a skull (<xref ref-type="fig" rid="RSOS140222F41">figures 41</xref>–<xref ref-type="fig" rid="RSOS140222F43">43</xref>).</p></sec><sec id="s6h8"><label>7.8.8</label><title>Geological range</title><p>(Lower Kimmeridgian?) Upper Kimmeridgian—Lower Tithonian.</p></sec><sec id="s6h9"><label>7.8.9</label><title>Geographical range</title><p>Europe (Portugal, Spain and Switzerland).</p></sec><sec id="s6h10"><label>7.8.10</label><title>Species diagnosis</title><p>Teleosaurid crocodylomorph within the genus <italic>Machimosaurus</italic> with the following unique combination of characters (autapomorphic characters are indicated by an asterisk *): the external surfaces of the skull bones are poorly ornamented, in particular those of the rostrum and around the orbits*; sub-globidont dentition*; apicobasally aligned enamel ridges immediately adjacent to the apical anastomosed region that are closely packed on both the labial and lingual tooth surfaces*; moderate post-symphyseal dentary tooth count (three to four pairs); inter-alveolar spaces between the maxillary and dentary alveoli are very small (closely packed alveoli)*; the premaxilla is notably wide at the level of the external nares, much wider than the width of the anterior end of the maxilla*; orbits are sub-rectangular in shape*; paroccipital processes are greatly enlarged, both elongated mediolaterally and with lateral ends that are expanded*; basioccipital apophysis has a ‘U-shaped’ cross section (teleosaurid symplesiomorphy); basioccipital tuberosities (basal tubera) are very large in size and are a sub-rectangular shape in occipital view*; the inter-basioccipital tubera notch is a large inverse ‘U’-shape when seen in occipital view*; dorsal osteoderm ornamentation is composed of small-to-large irregularly shaped pits arranged in a random manner, that are well separated from one another (somewhat similar to <italic>Steneosaurus leedsi</italic>).</p></sec></sec><sec id="s6i"><label>7.9</label><title><italic>Machimosaurus buffetauti</italic> sp. nov.</title><p><table-wrap orientation="portrait" id="d35e4231" position="anchor"><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col></colgroup><tbody><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1"><italic>1873</italic></td><td rowspan="1" colspan="1"><italic>Steneosaurus burgensis</italic> nomen nudum—Jarrin ([<xref rid="RSOS140222C72" ref-type="bibr">72</xref>], pp. 103–104)</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1"><italic>1876</italic></td><td rowspan="1" colspan="1"><italic>Steneosaurus burgensis</italic> nomen nudum—Jarrin ([<xref rid="RSOS140222C73" ref-type="bibr">73</xref>], pp. 94–96)</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1905</td><td rowspan="1" colspan="1"><italic>Steneosaurus burgensis Chanti</italic> nomen nudum—Chanel ([<xref rid="RSOS140222C74" ref-type="bibr">74</xref>], pp. 17–39), figs 1–3</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1982</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Buffetaut ([<xref rid="RSOS140222C2" ref-type="bibr">2</xref>], pp. 19–22), plate 1 figs A–D</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1982</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Buffetaut ([<xref rid="RSOS140222C3" ref-type="bibr">3</xref>], pp. 17–24), plate 1</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">2004</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Karl & Tichy [<xref rid="RSOS140222C8" ref-type="bibr">8</xref>], figs 1, 2</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">2006</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Karl <italic>et al</italic>. ([<xref rid="RSOS140222C32" ref-type="bibr">32</xref>], pp. 67–69), fig. 8</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">2008</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Lepage <italic>et al</italic>. ([<xref rid="RSOS140222C17" ref-type="bibr">17</xref>], pp. 116–118), figs 1–7</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1"><italic>2008</italic></td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Pierce <italic>et al</italic>. ([<xref rid="RSOS140222C9" ref-type="bibr">9</xref>], p. 1085) (<italic>partim</italic>)</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">2013</td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Martin & Vincent ([<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], pp. 179–196), figs 1–9</td></tr></tbody></table></table-wrap></p><sec id="s6i1"><label>7.9.1</label><title>Holotype</title><p>SMNS 91415: complete skull and mandible, with partial postcranial skeleton (<xref ref-type="fig" rid="RSOS140222F22">figures 22</xref>–<xref ref-type="fig" rid="RSOS140222F27">27</xref>).</p></sec><sec id="s6i2"><label>7.9.2</label><title>Holotype locality</title><p>Am Hörnle Quarry, Neuffen, Baden-Württemberg, Germany [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>].</p></sec><sec id="s6i3"><label>7.9.3</label><title>Holotype horizon</title><p>Lacunosamergel Formation, <italic>A. hypselocyclum</italic> Sub-Mediterranean ammonite Zone (Weißer Jura gamma 2), Lower Kimmeridgian, Upper Jurassic [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>].</p></sec><sec id="s6i4"><label>7.9.4</label><title>Etymology</title><p>‘Buffetaut's pugnacious lizard’. Named in honour of Eric Buffetaut (b. 1950), whose research has greatly elucidated thalattosuchian and crocodylomorph evolution.</p></sec><sec id="s6i5"><label>7.9.5</label><title>Referred specimens</title><p>MPV V.1600.Bo and V.1601.Bo—anterior half of rostrum (premaxilla, maxilla and dentary) in occlusion and a maxilla-nasal fragment (Calcaires Coquilliers Formation; <italic>P</italic>. <italic>baylei</italic> Sub-Boreal ammonite Zone, lowermost Kimmeridgian of Cricqueboeuf, Normandy, Northern France; <xref ref-type="fig" rid="RSOS140222F12">figures 12</xref>–<xref ref-type="fig" rid="RSOS140222F14">14</xref>) [<xref rid="RSOS140222C2" ref-type="bibr">2</xref>,<xref rid="RSOS140222C17" ref-type="bibr">17</xref>].</p><p>DFMMh FV 330, DFMMh FV 541: isolated tooth crowns (Langenberg Formation; Langenberg near Oker, Lower Saxony, Germany; Kimmeridgian; <xref ref-type="fig" rid="RSOS140222F21">figure 21</xref><italic>c</italic>–<italic>h</italic>) [<xref rid="RSOS140222C8" ref-type="bibr">8</xref>,<xref rid="RSOS140222C32" ref-type="bibr">32</xref>].</p><p>Musée de Brou (Bourg-en-Bresse, France), specimen number unknown—a complete skull and mandible in articulation (Calcaires à ptérocères Formation, Lower Kimmeridgian; Montmerle, Bourg-en-Bresse, département de l'Ain, France [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>]).</p></sec><sec id="s6i6"><label>7.9.6</label><title>Geological range</title><p>Lower Kimmeridgian.</p></sec><sec id="s6i7"><label>7.9.7</label><title>Geographical range</title><p>Europe (France and Germany). An isolated tooth from Smallmouth Sands, England (NHMUK PV R1774; <xref ref-type="fig" rid="RSOS140222F21">figure 21</xref><italic>a</italic>,<italic>b</italic>) is possibly referable to this taxon, as are isolated teeth from Czarnogłowy, Poland (GPIT/RE/328, GPIT/RE/9280 and GPIT/RE/9281; <xref ref-type="fig" rid="RSOS140222F31">figures 31</xref> and <xref ref-type="fig" rid="RSOS140222F32">32</xref>).</p></sec><sec id="s6i8"><label>7.9.8</label><title>Species diagnosis</title><p>Teleosaurid crocodylomorph within the genus <italic>Machimosaurus</italic> with the following unique combination of characters (autapomorphic characters are indicated by an asterisk *): 21–22 alveoli per maxilla (approx. 16–17 of which are anterior to the palatines); 24/25 alveoli per dentary (19–20 of which are adjacent to the mandibular symphysis); low post-symphyseal dentary tooth count (two pairs)*; inter-alveolar spaces between the maxillary and dentary alveoli are variable in size (thalattosuchian symplesiomorphy); orbits are sub-circular in shape (transverse and anteroposterior axes are sub-equal; the <italic>Steneosaurus brevior</italic> holotype also has circular orbits)*; the quadrates have a single large circular depression on the dorsal surface near the hemicondyles*; basioccipital apophysis has a ‘U-shaped’ cross section (teleosaurid symplesiomorphy); the inter-basioccipital tubera notch is a wide and gentle inverse semicircle when seen in occipital view (teleosaurid symplesiomorphy); basioccipital tuberosities (basal tubera) are reduced in size when seen in occipital view (apomorphy shared with <italic>M. mosae</italic>); axis neural arch dorsal margin is strongly concave when seen in lateral view*; axis postzygapophyses terminate significantly posterior to posterior surface of the centrum view (somewhat similar to that seen in <italic>S. durobrivensis</italic>); coracoid glenoid process (process near the glenoid fossa that projects posterodorsally) is elongate, extending considerably from the head of the coracoid, and is a sub-isosceles triangle in shape when seen in lateral view*; coracoid postglenoid process anterior margin is very slightly concave and terminates approximately in the same frontal plane as the glenoid*; coracoid postglenoid process posterior margin is strongly concave and terminates approximately in the same frontal plane as the posterior end of the glenoid process*; dorsal osteoderm ornamentation is composed of small-to-large irregularly shaped pits arranged in a random manner, that are well separated from one another (somewhat similar to <italic>S. leedsi</italic>).</p><p><italic>Steneosaurus burgensis</italic>. The names <italic>S. burgensis</italic> and <italic>S. burgensis chanti</italic> have been applied to the <italic>Machimosaurus</italic> skull from Ain, France [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>,<xref rid="RSOS140222C72" ref-type="bibr">72</xref>–<xref rid="RSOS140222C74" ref-type="bibr">74</xref>]. These specific and sub-specific names are however <italic>nomina nuda</italic>. Both Jarrin [<xref rid="RSOS140222C72" ref-type="bibr">72</xref>] and Chanel [<xref rid="RSOS140222C74" ref-type="bibr">74</xref>] stated that the Ain skull was sent to Caen for preparation and study by Eugène Eudes-Deslongchamps, who proposed the name <italic>S. burgensis</italic> for the specimen, in consultation with the Société d'émulation de l'Ain. Neither Jarrin [<xref rid="RSOS140222C72" ref-type="bibr">72</xref>,<xref rid="RSOS140222C73" ref-type="bibr">73</xref>] nor Chanel [<xref rid="RSOS140222C74" ref-type="bibr">74</xref>] established the name under Article 12 of the International Commission on Zoological Nomenclature (ICZN) Code as: they did not describe the specimen, nor did they provide a definition of the species; they simply reported that the name was proposed by Eudes-Deslongchamps. Unfortunately, Eudes-Deslongchamps never published his description of the Ain skull [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>,<xref rid="RSOS140222C74" ref-type="bibr">74</xref>]. The specimen was not formally described until 1982, and then it was referred to <italic>M. hugii</italic> [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>]. The sub-specific epithet <italic>chanti</italic> was apparently established by those who did not fully understand zoological nomenclature [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>], as it was added to the Ain skull's specimen plaque solely to honour the discoverer [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>,<xref rid="RSOS140222C74" ref-type="bibr">74</xref>].</p><p>Our decision to establish a new taxon based on SMNS 91415, and not formally establish <italic>S. burgensis</italic> for the Ain skull, was for several reasons: (i) the Ain skull is partially reconstructed, and it is unclear how much is plaster and how much is real bone [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>]; (ii) the cranium and lower jaw of the Ain skull are in articulation, meaning that the palatal and dorsal mandibular morphologies cannot be seen [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>,<xref rid="RSOS140222C74" ref-type="bibr">74</xref>]; (iii) the German skull SMNS 91415 has the cranium and lower jaw disarticulated, allowing these morphologies to be observed [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>]; and (iv) SMNS 91415 has associated postcranial material, greatly aiding in comparisons with other <italic>Machimosaurus</italic> taxa, in particular <italic>M. mosae</italic>.</p></sec></sec><sec id="s6j"><label>7.10</label><title><italic>Machimosaurus mosae</italic> Sauvage & Liénard, 1879 [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>]</title><p><table-wrap orientation="portrait" id="d35e4620" position="anchor"><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col></colgroup><tbody><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">1876</td><td rowspan="1" colspan="1"><italic>Teleosaurusmosae</italic> sp. nov.—Liénard (manuscript name)</td></tr><tr><td rowspan="1" colspan="1">v*</td><td rowspan="1" colspan="1">1879</td><td rowspan="1" colspan="1"><italic>Machimosaurus mosae</italic> comb. nov.—Sauvage & Liénard [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>, pp. 1–31, plates 1–4]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1"><italic>1973</italic></td><td rowspan="1" colspan="1"><italic>Machimosaurus hugii</italic> von Meyer—Steel [<xref rid="RSOS140222C71" ref-type="bibr">71</xref>, p. 25 (<italic>partim</italic>)]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1993</td><td rowspan="1" colspan="1"><italic>Machimosaurus mosae</italic> Sauvage & Liénard—Hua <italic>et al</italic>. [<xref rid="RSOS140222C21" ref-type="bibr">21</xref>, pp. 851–856, texte-fig. 1]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1999</td><td rowspan="1" colspan="1"><italic>Machimosaurus mosae</italic> Sauvage & Liénard—Hua [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>, pp. 141–170, figs 1 and 2, plates 1–6]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1"><italic>2009</italic></td><td rowspan="1" colspan="1"><italic>Machimosaurus mosae</italic> Sauvage & Liénard—Pierce <italic>et al</italic>. [<xref rid="RSOS140222C9" ref-type="bibr">9</xref>, p. 1085]</td></tr></tbody></table></table-wrap></p><sec id="s6j1"><label>7.10.1</label><title>Holotype</title><p>Much of the skeleton: incomplete skull, mandible, 22 vertebrae, part of the pelvis, numerous ribs, limb bones and 22 osteoderms. The specimen disappeared during the First World War [<xref rid="RSOS140222C20" ref-type="bibr">20</xref>] and is presumed to have been destroyed.</p></sec><sec id="s6j2"><label>7.10.2</label><title>Holotype locality and horizon</title><p>Issoncourt, near Verdun, Département de la Meuse, Lorraine, France. The specimen most likely comes from the <italic>A. autissiodorensis</italic> Sub-Boreal ammonite zone, ‘Marnes supérieures de la Meuse’ [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>].</p></sec><sec id="s6j3"><label>7.10.3</label><title>Neotype</title><p>An almost complete skeleton: nearly complete skull, mandible, half of the cervical vertebrae, all the dorsal and sacral vertebrae, approximately a third of the caudal vertebrae, two chevrons, cervical and dorsal ribs, left scapula, right coracoid, right fibula, both pubes, both ilia, left ischium, right femur, left tibia, and dorsal and ventral osteoderms [<xref rid="RSOS140222C21" ref-type="bibr">21</xref>] (<xref ref-type="fig" rid="RSOS140222F15">figures 15</xref>–<xref ref-type="fig" rid="RSOS140222F20">20</xref>).</p></sec><sec id="s6j4"><label>7.10.4</label><title>Neotype locality</title><p>A beach near Ambleteuse, Boulonnais, Département du Pas-de-Calais, Nord Pas-de-Calais, France.</p></sec><sec id="s6j5"><label>7.10.5</label><title>Neotype horizon</title><p>Argiles de Châtillon Formation [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C21" ref-type="bibr">21</xref>]. From either the <italic>A. autissiodorensis</italic> Sub-Boreal ammonite zone, uppermost Kimmeridgian, or the <italic>G. gigas</italic>/<italic>P. elegans</italic> Sub-Boreal ammonite zone, lowermost Tithonian.</p></sec><sec id="s6j6"><label>7.10.6</label><title>Neotype note</title><p>The neotype was originally catalogued as BHN2R 1100. While the BHN2R closed in 2003, the neotype was removed from the museum prior to this. It is assumed that the neotype is now in a private collection, but this cannot be confirmed. A cast of the neotype is on display in RBINS. It was purchased from Eldonia Paléontologie, and it is unclear how much is based on the original specimen (P. Godefroit 2014, personal communication). The original cast was made by the University of Paris 6—Université Pierre-et-Marie-Curie (E. Buffetaut 2014, personal communication), and we are unsure how Eldonia Paléontologie obtained a copy.</p></sec><sec id="s6j7"><label>7.10.7</label><title>Etymology</title><p>‘Pugnacious lizard of the Meuse’. Named after the French river Meuse, near which the holotype was discovered.</p></sec><sec id="s6j8"><label>7.10.8</label><title>Previously referred specimen</title><p>Lydekker [<xref rid="RSOS140222C70" ref-type="bibr">70</xref>] referred an incomplete skull and mandible (NHMUKPV R1089) from Upper Kimmeridge Clay Formation (Early Tithonian) of England to <italic>M. mosae</italic>. This specimen, however, was recently shown to belong to the metriorhynchid <italic>P. manselii</italic> [<xref rid="RSOS140222C41" ref-type="bibr">41</xref>].</p></sec><sec id="s6j9"><label>7.10.9</label><title>Geological range</title><p>Uppermost Kimmeridgian and/or lowermost Tithonian.</p></sec><sec id="s6j10"><label>7.10.10</label><title>Geographical range</title><p>Europe (northeastern France).</p></sec><sec id="s6j11"><label>7.10.11</label><title>Species diagnosis</title><p>Teleosaurid crocodylomorph within the genus <italic>Machimosaurus</italic> with the following unique combination of characters (autapomorphic characters are indicated by an asterisk *): 17–18 alveoli per maxilla (approx. 14 of which are anterior to the palatines); 19 alveoli per dentary (15–16 of which are adjacent to the mandibular symphysis); moderate post-symphyseal dentary tooth count (three to four pairs); inter-alveolar spaces between the maxillary and dentary alveoli are variable in size (thalattosuchian symplesiomorphy); orbits are transverse ellipsoids in shape (anteroposterior axis is 79.7% the length of the transverse axis)*; prearticular is absent*; basioccipital apophysis has a ‘V-shaped’ cross section*; the inter-basioccipital tubera notch is a wide and gentle inverse semicircle when seen in occipital view (teleosaurid symplesiomorphy); basioccipital tuberosities (basal tubera) are reduced in size when seen in occipital view (apomorphy shared with <italic>M. buffetauti</italic>); axis neural arch dorsal margin is subtly concave when seen in lateral view (somewhat similar to that seen in <italic>S. leedsi</italic>); axis postzygapophyses terminate only slightly posterior to posterior surface of the centrum*; coracoid glenoid process is very short, not extending far from the head of the coracoid and is a right-angled triangle in shape when seen in lateral view*; coracoid postglenoid process anterior margin is strongly concave and terminates in a frontal plane anterior to the glenoid*; coracoid postglenoid process posterior margin is strongly concave distally but shifts to being somewhat convex proximally and terminates in a frontal plane posterior to the posterior end of the glenoid process*; dorsal osteoderm ornamentation is composed of numerous small, irregularly shaped pits arranged in an anastomosed pattern, these pits can fuse and become elongate grooves that radiate from the keel (similar to ‘<italic>S</italic>.’ <italic>obtusidens</italic>)*; ventral osteoderms have a longitudinal keel*.</p></sec><sec id="s6j12"><label>7.10.12</label><title>Validity of <italic>Machimosaurus mosae</italic></title><p>Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>, p. 193] claimed that <italic>M. mosae</italic> was an invalid name, based upon Articles 8 and 9 of the Code of the ICZN. Their contention was based on the 1876 manuscript of Liénard entitled ‘Le <italic>Teleosaurus Mosae</italic>, fossile des marnes kimméridgiennes de la Meuse’ [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>, p. 7], which was the first to use the name <italic>T. Mosae</italic>. Sauvage & Liénard [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>, p. 7] stated that Liénard's description remained in manuscript form and was sent to the Ministry of Public Education at the end of 1876, ‘ont été indiqués par lui dans un travail resté manuscrit et adressé à la fin de l'année 1876 au Ministère de l'Instruction publique’.</p><p>Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] are correct that Sauvage & Liénard [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>, p. 11] used the following headings:</p><p>Deuxieme Partie.—Description du Machimosaurus mosæ, <italic>F. Liénard</italic> sp. (1).</p><p>However, this is key to why <italic>T. mosae</italic> is an available name under the ICZN Code. Immediately below those headings is a detailed species diagnosis, followed by an eight-page description of the skeleton, a long discussion on the affinities of <italic>Machimosaurus</italic>, and four plates with line drawings of the specimen. As such, along with the paper being published in a scientific journal, this description clearly fulfils the criteria set out in Articles 8, 11 and 12 of the ICZN Code.</p><p>Article 11.6 of the ICZN Code, publication as synonymy, states: ‘A name which when first published in an available work was treated as a junior synonym of a name then used as valid is not thereby made available’. Article 11.6.1 states: ‘However, if such a name published as a junior synonym had been treated before 1961 as an available name and either adopted as the name of a taxon or treated as a senior homonym, it is made available thereby but dates from its first publication as a synonym’. Clearly, Sauvage & Liénard [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>] treated <italic>T. mosae</italic> as an available name, and they adopted its specific name for their validly described taxon. Consequently, under Article 11.6.1 <italic>M. mosae</italic> is an available name.</p><p>However, the authorship is Sauvage & Liénard, 1879 not Liénard, 1876. This is due to Article 50.7 of the ICZN Code, which states: ‘If a scientific name (taken, for example, from a label or manuscript) was first published in the synonymy of an available name and became available before 1961 through the provisions of Article 11.6, its author is the person who published it as a synonym, even if some other originator is cited, and is not the person who subsequently adopted it as a valid name.’ Therefore, <italic>contra</italic> Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], <italic>M. mosae</italic> is not invalid, and the nominal authors of the specific name are indeed Sauvage & Liénard [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>].</p></sec><sec id="s6j13"><label>7.10.13</label><title>Tooth taxa of Sauvage [<xref rid="RSOS140222C11" ref-type="bibr">11</xref>]</title><p>Sauvage [<xref rid="RSOS140222C11" ref-type="bibr">11</xref>] listed three <italic>Machimosaurus</italic> species living in the Late Kimmeridgian–Early Tithonian of Northern France (around Boulogne-sur-Mer): <italic>M. hugii</italic>, <italic>M. interruptus</italic> and <italic>M. ferox</italic>. The latter two were established by Sauvage [<xref rid="RSOS140222C11" ref-type="bibr">11</xref>] for isolated tooth crowns from the area. Krebs [<xref rid="RSOS140222C8" ref-type="bibr">8</xref>, p. 48] stated that Sauvage invoked insignificant differences between the crowns when establishing his species and that Sauvage himself later withdrew the name <italic>M. ferox</italic>. Owing to the geological age and location of these isolated teeth, it is possible they are referable to <italic>M. mosae</italic>. If so, <italic>M. interruptus</italic> would have priority. As the holotype of <italic>M. interruptus</italic> cannot be found, and the neotype of <italic>M. mosae</italic> is currently unavailable for study, this possibility cannot be explored.</p></sec></sec><sec id="s6k"><label>7.11</label><title><italic>Machimosaurus nowackianus</italic> (von Huene, 1938 [<xref rid="RSOS140222C18" ref-type="bibr">18</xref>]) comb. nov.</title><p><table-wrap orientation="portrait" id="d35e4979" position="anchor"><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col></colgroup><tbody><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1938</td><td rowspan="1" colspan="1">cf. <italic>Simolestes nowackianus</italic> sp. nov. —von Huene [<xref rid="RSOS140222C19" ref-type="bibr">19</xref>, pp. 370–376, figs 1–4]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1960</td><td rowspan="1" colspan="1"><italic>Simolestes nowackianus</italic> von Huene—Tarlo [<xref rid="RSOS140222C75" ref-type="bibr">75</xref>, pp. 173, 183, fig. 3<italic>c</italic>]</td></tr><tr><td rowspan="1" colspan="1">v</td><td rowspan="1" colspan="1">1996</td><td rowspan="1" colspan="1"><italic>Machimosaurus</italic> sp. —Bardet & Hua [<xref rid="RSOS140222C20" ref-type="bibr">20</xref>, pp. 65–71, figs 1–2]</td></tr></tbody></table></table-wrap></p><sec id="s6k1"><label>7.11.1</label><title>Holotype</title><p>GPIT Orig. Huene 1938 figs <xref ref-type="fig" rid="RSOS140222F1">1</xref>–<xref ref-type="fig" rid="RSOS140222F4">4</xref>: anterior region of the dentary (<xref ref-type="fig" rid="RSOS140222F3">figure 3</xref>). Note that the specimen currently cannot be located in GPIT.</p></sec><sec id="s6k2"><label>7.11.2</label><title>Holotype locality</title><p>Near Feyambiro, east of Harrar, Harrar Province, Ethiopia [<xref rid="RSOS140222C19" ref-type="bibr">19</xref>].</p></sec><sec id="s6k3"><label>7.11.3</label><title>Holoype horizon</title><p>It is not clear what formation the specimen was found in [<xref rid="RSOS140222C19" ref-type="bibr">19</xref>,<xref rid="RSOS140222C20" ref-type="bibr">20</xref>]. It was described as being found in sandy clays, 8 m above a crystalline basement.</p></sec><sec id="s6k4"><label>7.11.4</label><title>Etymology</title><p>‘Nowack's pugnacious lizard’. Named in honour of the holotype's discoverer, Mr Nowack.</p></sec><sec id="s6k5"><label>7.11.5</label><title>Geological range</title><p>Oxfordian or Kimmeridgian.</p></sec><sec id="s6k6"><label>7.11.6</label><title>Geographical range</title><p>Africa (Ethiopia).</p></sec><sec id="s6k7"><label>7.11.7</label><title>Species diagnosis</title><p>Teleosaurid crocodylomorph within the genus <italic>Machimosaurus</italic> with the following unique combination of characters (autapomorphic characters are indicated by an asterisk *): the anterior dentary inter-alveolar spaces are reduced, being notably smaller in size than in <italic>M. buffetauti</italic> and <italic>M. mosae</italic>, in particular the D1–D2 and D2–D3 inter-alveolar spaces which are both less than half the length of the D2 alveoli*; the interdentary distance between the D3 and D4 alveolar couplets is notably smaller than in <italic>M. buffetatuti</italic> and <italic>M. mosae</italic>, such that the transverse width of the left and right D4 alveoli are as large as, or greater than, the immediately adjacent flat dentary region*.</p><p><italic>Note.</italic> While the two anterior dentary characteristics readily differentiate <italic>M. nowackianus</italic> from both <italic>M. buffetauti</italic> and <italic>M. mosae</italic>, no known <italic>M. hugii</italic> specimen preserves the anterior region of the dentary.</p></sec></sec></sec><sec sec-type="discussion" id="s7"><label>8.</label><title>Discussion</title><sec id="s7a"><label>8.1</label><title>Ontogeny</title><p>The differences in tooth count, symphyseal length and length of the supratemporal fossae enable easy identification for two of the four <italic>Machimosaurus</italic> species (<italic>M. buffetauti</italic> and <italic>M. mosae</italic>; <xref ref-type="table" rid="RSOS140222TB1">table 1</xref>). Interestingly, there are three skulls of <italic>M. buffetauti</italic> and <italic>M. mosae</italic> between 93 and 100 cm in basicranial length. All of these have a snout (=preorbital) length to basicranial length ratio of approximately 58%, even though there is variation in maxillary tooth count, dentary tooth count, dentary symphyseal tooth count and mandibular symphysis length. This shows that proportional snout length does not vary between these two species (even though these dental and mandibular characteristics do vary). This indicates that the differences between these two species are not due to ontogeny, because it would be unusual for juveniles and adults of the same taxon to have nearly identical skull lengths and nearly identically proportioned snouts. Moreover, when we look beyond tooth counts and biometric ratios and examine cranial, mandibular and postcranial morphologies, we find numerous characteristics that differentiate these two species (see diagnoses and <xref ref-type="table" rid="RSOS140222TB2">tables 2</xref> and <xref ref-type="table" rid="RSOS140222TB3">3</xref>).
<table-wrap id="RSOS140222TB3" orientation="portrait" position="float"><label>Table 3.</label><caption><p>Comparison of craniomandibular and postcranial morphologies among <italic>Machimosaurus</italic> specimens.</p></caption><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col></colgroup><thead valign="bottom"><tr><th align="left" rowspan="1" colspan="1">species</th><th align="left" colspan="2" rowspan="1"><italic>M. buffetauti</italic><hr></hr></th><th align="left" colspan="2" rowspan="1"><italic>M. mosae</italic><hr></hr></th><th align="left" colspan="2" rowspan="1"><italic>M. hugii</italic><hr></hr></th></tr><tr><th align="left" rowspan="1" colspan="1">reference</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C10" ref-type="bibr">10</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C3" ref-type="bibr">3</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C62" ref-type="bibr">62</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C4" ref-type="bibr">4</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C6" ref-type="bibr">6</xref>]</th><th align="left" rowspan="1" colspan="1">[<xref rid="RSOS140222C63" ref-type="bibr">63</xref>]</th></tr></thead><tbody><tr><td rowspan="1" colspan="1">orbit shape</td><td rowspan="1" colspan="1">circular</td><td rowspan="1" colspan="1">circular</td><td rowspan="1" colspan="1">transverse ellipsoid</td><td rowspan="1" colspan="1">transverse ellipsoid</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">sub-rectangular</td></tr><tr><td rowspan="1" colspan="1">quadrate dorsal surfaces have a large circular depression</td><td rowspan="1" colspan="1">yes</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">no</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">paroccipital processes</td><td rowspan="1" colspan="1">similar in size and shape to other teleosaurids</td><td rowspan="1" colspan="1">similar in size and shape to other teleosaurids</td><td rowspan="1" colspan="1">similar in size and shape to other teleosaurids</td><td rowspan="1" colspan="1">similar in size and shape to other teleosaurids</td><td rowspan="1" colspan="1">mediolaterally elongate and the lateral ends are greatly expanded</td><td rowspan="1" colspan="1">mediolaterally elongate and the lateral ends are greatly expanded</td></tr><tr><td rowspan="1" colspan="1">basioccipital condyle apophysis cross section</td><td rowspan="1" colspan="1">‘U’-shaped</td><td rowspan="1" colspan="1">‘U’-shaped</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">‘V’-shaped</td><td rowspan="1" colspan="1">‘U’-shaped</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">basioccipital tuberosity size</td><td rowspan="1" colspan="1">small</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">small</td><td rowspan="1" colspan="1">very large</td><td rowspan="1" colspan="1">very large</td></tr><tr><td rowspan="1" colspan="1">inter-tuber notch (basioccipital tuberosity) shape in occipital view</td><td rowspan="1" colspan="1">semicircular, mediolaterally wide</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">semicircular, mediolaterally wide</td><td rowspan="1" colspan="1">inverse ‘U’-shape, mediolaterally narrow</td><td rowspan="1" colspan="1">inverse ‘U’-shape, mediolaterally narrow</td></tr><tr><td rowspan="1" colspan="1">prearticular</td><td rowspan="1" colspan="1">present</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">absent</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">axis neural arch dorsal margin in lateral view</td><td rowspan="1" colspan="1">strongly concave</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">subtly concave</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">coracoid glenoid process, shape in lateral view</td><td rowspan="1" colspan="1">sub-isosceles triangle</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">right-angled triangle</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">coracoid postglenoid process, anterior margin shape</td><td rowspan="1" colspan="1">very slightly concave, in the same frontal plane as the glenoid</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">strongly concave, in a frontal plane that is anterior to the glenoid</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">coracoid postglenoid process, posterior margin shape</td><td rowspan="1" colspan="1">strongly concave, in the same frontal plane as the posterior end of the glenoid process</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">strongly concave distally and becomes convex proximally, in a frontal plane posterior to the posterior end of the glenoid process</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1">dorsal osteoderm ornamentation pattern</td><td rowspan="1" colspan="1">small-to-large sub-circular pits that are well separated</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">tightly packed, small, sub-circular pits arranged in an anastomosed pattern</td><td rowspan="1" colspan="1">small-to-large sub-circular pits that are well separated</td><td rowspan="1" colspan="1">?</td></tr></tbody></table></table-wrap></p><p>This leaves the type species <italic>M. hugii</italic> as a remaining point of discussion for the European species. Unfortunately, the postcranial characteristics that differentiate <italic>M. buffetauti</italic> and <italic>M. mosae</italic> are unknown in <italic>M. hugii</italic> (<xref ref-type="table" rid="RSOS140222TB3">table 3</xref>), as are the differences in cranial biometrics and tooth counts (<xref ref-type="table" rid="RSOS140222TB1">tables 1</xref> and <xref ref-type="table" rid="RSOS140222TB2">2</xref>). <italic>Machimosaurus hugii</italic> does have three braincase apomorphies: (i) paroccipital processes are greatly enlarged, both elongated mediolaterally and with lateral ends that are expanded; (ii) basioccipital tuberosities (basal tubera) are very large in size; and (iii) inter-basioccipital tubera notch is a large inverse ‘U’-shape when seen in occipital view. <italic>Machimosaurus hugii</italic> also has two dental apomorphies: (i) sub-globidont dentition and (ii) apicobasally aligned enamel ridges immediately adjacent to the apical anastomosed region that are closely packed on both the labial and lingual tooth surfaces. These five characteristics readily allow <italic>M. hugii</italic> to be identified, and seem unlikely to be under ontogenetic control (for example, a drastic change between non-globidont and sub-globidont dentition is not known in any other crocodylomorph and would be greatly unexpected). Therefore, it is unlikely that <italic>M. hugii</italic> is the adult version of either <italic>M. buffetauti</italic> or <italic>M. mosae</italic>. As further support of this conclusion, we note that the smaller <italic>M. buffetauti</italic> and <italic>M. mosae</italic> individuals do not show any juvenile characteristics (such as having proportionally large orbits to basicranial length or paired frontals (P. Vignaud 1995, unpublished PhD thesis)). Moreover, there is variation in orbit shape, frontal shape and premaxilla width (<xref ref-type="fig" rid="RSOS140222F44">figure 44</xref>).
<fig id="RSOS140222F44" orientation="portrait" position="float"><label>Figure 44.</label><caption><p>Reconstructions of the skulls of the European <italic>Machimosaurus</italic> species in dorsal view. (<italic>a</italic>) <italic>Machimosaurus mosae</italic> (the neotype), (<italic>b</italic>) <italic>M. buffetauti</italic> (the holotype SMNS 91415) and (<italic>c</italic>) <italic>M. hugii</italic> (based on MG-8730-1, MG-8730-2, NMS 7012 and NMS 7029).</p></caption><graphic xlink:href="rsos140222-g44"></graphic></fig></p></sec><sec id="s7b"><label>8.2</label><title>Age and locality</title><p>The four <italic>Machimosaurus</italic> species were not spatio-temporally contemporaneous. All of the specimens we refer to <italic>M. buffetauti</italic> are from the Lower Kimmeridgian of France and Germany (and possibly also England and Poland, although we cannot confirm this at present). <italic>Machimosaurus mosae</italic> is known from the final Sub-Boreal ammonite zone of the Kimmeridgian and/or the first Sub-Boreal ammonite zone of the Tithonian, and only known from northeastern France. As such, <italic>M. buffetauti</italic> and <italic>M. mosae</italic> were separated in time by 3–5 million years (<xref ref-type="fig" rid="RSOS140222F45">figure 45</xref>).
<fig id="RSOS140222F45" orientation="portrait" position="float"><label>Figure 45.</label><caption><p>Map of <italic>Machimosaurus</italic> specimens from across Europe, (<italic>a</italic>) in the Oxfordian, (<italic>b</italic>) the Early Kimmeridgian and (<italic>c</italic>) during the Late Kimmeridgian–Early Tithonian. Localities listed below with an asterisk (*) denote where we are not certain that specimens from that locality can be referred to the listed species. <italic>Machimosaurus</italic> sp.: (1) Haudainville near Verdun, Lorraine, France; (2) Villerville, Normandy, France; (3) Cesaredas, central west Portugal; (4) Malhão, Algarve, Portugal; (5) Cesareda, Portugal. <italic>Machimosaurus buffetauti</italic>: (6) Smallmouth Sands, Dorset, England*; (7) Cricqueboeuf, Normandy, France; (8) Montmerle, Bourg-en-Bresse, France; (9) Neuffen, Baden-Württemberg, Germany; (10) Langenberg near Oker, Germany; (11) Czarnogłowy, West Pomerania, Poland. <italic>Machimosaurus hugii</italic>: (12) Guimarota, Leiria, Portugal; (13) La Escalera, Villaviciosa, Asturias, Spain; (14) Playa de La Griega, Colunga, Asturias, Spain; (15) Cantera Carcalín near Buñol, Valencia Province, Spain; (16) Solothurn, Canton Solothurn, Switzerland; (17) Moutier, Canton Bern, Switzerland*; (18) Porrentruy, Canton Jura, Switzerland*; (19–22) Santa-Cruz, Porto das Barcas, Peralta and Zimbral, Lourinhã area, Portugal. <italic>Machimosaurus mosae</italic>: (23) Issoncourt, near Verdun, Lorraine, France; (24) Mont-Lambert near Boulogne-sur-Mer, France*; (25) beach near Ambleteuse, Boulonnais, France. Palaeomaps were modified version of high-resolution versions kindly provided by Ron Blakey (<uri xlink:href="http://cpgeosystems.com/">http://cpgeosystems.com/</uri>).</p></caption><graphic xlink:href="rsos140222-g45"></graphic></fig></p><p>The lectotype of <italic>M. hugii</italic> is known from the Solothurn turtle limestone, a member of the Reuchenette Formation (Switzerland), which was deposited during the final Sub-Mediterranean ammonite zone of the Kimmeridgian. This suggests that <italic>M. hugii</italic> and <italic>M. mosae</italic>were probably contemporaneous in age, but lived in different European provinces, with <italic>M. hugii</italic> in the Sub-Mediterranean realm and <italic>M. mosae</italic> in the Sub-Boreal realm (<xref ref-type="fig" rid="RSOS140222F45">figure 45</xref>).</p><p>The other <italic>M. hugii</italic> specimens are known from Portugal and Spain. As noted above, the exact age of the Leiria specimen is unknown, with it being either Early or Late Kimmeridgian in age. The Lourinhã specimens are close to the Kimmeridgian–Tithonian boundary. Isolated Spanish teeth are Kimmeridgian and Early Tithonian in age [<xref rid="RSOS140222C25" ref-type="bibr">25</xref>,<xref rid="RSOS140222C48" ref-type="bibr">48</xref>]. All the Iberian specimens have the same dental morphologies as the uppermost Kimmeridgian Swiss specimens (apicobasally aligned enamel ridges immediately adjacent to the apical anastomosed region that are closely packed on both the labial and lingual tooth surfaces [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C48" ref-type="bibr">48</xref>]).</p><p><italic>Machimosaurus nowackianus</italic> is the only known non-European taxon, and thus was widely separated from the three European taxa. It is currently difficult to assess whether it may have been temporally separated from the European taxa as well, as the age of the Ethiopian deposits is only coarsely constrained to the Oxfordian–Kimmeridgian. In summary, all four <italic>Machimosaurus</italic> species were either separated by several million years in time or by hundreds or thousands of kilometres.</p></sec><sec id="s7c"><label>8.3</label><title>Body size</title><p>The neotype of <italic>M. mosae</italic> gives the first definitive evidence of body length in this genus. The skeleton was approximately 6 m in length and had a basicranial length of 96.5 cm [<xref rid="RSOS140222C21" ref-type="bibr">21</xref>]. This therefore allows us to make an estimate of total body length for any <italic>Machimosaurus</italic> specimen with a basicranial length (assuming skull-to-body length scaling remains consistent). These estimates reveal something quite interesting about three of the four species (<xref ref-type="table" rid="RSOS140222TB4">table 4</xref>). The largest known <italic>M. buffetauti</italic> skull is from Ain, France [<xref rid="RSOS140222C3" ref-type="bibr">3</xref>]. At 100 cm long, it gives a body length estimate of 6.22 m. The holotype of <italic>M. buffetauti</italic> has a basicranial length of 93.5 cm [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>]. This results in a slightly smaller body length of 5.81 m.
<table-wrap id="RSOS140222TB4" orientation="portrait" position="float"><label>Table 4.</label><caption><p>Estimated total body lengths for <italic>Machimosaurus</italic> specimens. Estimate based on the ratio of the basicranial length (96.5 cm) to total body length (approx. 600 cm) of the <italic>M. mosae</italic> neotype [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C21" ref-type="bibr">21</xref>].</p></caption><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col><col align="left" span="1"></col></colgroup><thead valign="bottom"><tr><th align="left" rowspan="1" colspan="1">species</th><th align="left" rowspan="1" colspan="1">references</th><th align="left" rowspan="1" colspan="1">basicranial length (cm)</th><th align="left" rowspan="1" colspan="1">body length estimate (m)</th></tr></thead><tbody><tr><td rowspan="1" colspan="1"><italic>M. hugii</italic></td><td rowspan="1" colspan="1">[<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>]</td><td rowspan="1" colspan="1">149</td><td rowspan="1" colspan="1">9.26</td></tr><tr><td rowspan="1" colspan="1"><italic>M. mosae</italic></td><td rowspan="1" colspan="1">[<xref rid="RSOS140222C62" ref-type="bibr">62</xref>]</td><td rowspan="1" colspan="1">130</td><td rowspan="1" colspan="1">8.08</td></tr><tr><td rowspan="1" colspan="1"><italic>M. buffetauti</italic></td><td rowspan="1" colspan="1">[<xref rid="RSOS140222C3" ref-type="bibr">3</xref>]</td><td rowspan="1" colspan="1">100</td><td rowspan="1" colspan="1">6.22</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1">[<xref rid="RSOS140222C10" ref-type="bibr">10</xref>]</td><td rowspan="1" colspan="1">93.5</td><td rowspan="1" colspan="1">5.81</td></tr></tbody></table></table-wrap></p><p>While the neotype of <italic>M. mosae</italic> is approximately 6 m in body length, the lost holotype was far larger. It had an estimated basicranial length of 130 cm [<xref rid="RSOS140222C62" ref-type="bibr">62</xref>]. This results in a body length estimate of 8.08 m, making it substantially larger than the older <italic>M. buffetauti</italic>. The type species <italic>M. hugii</italic> was larger still. The incomplete Leira skull was estimated to have a basicranial length of 149 cm [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>]. This gives a body length estimate of 9.26 m. As such, Krebs' [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>] estimate of the Leira specimen exceeding 9 m long in length was reasonable.</p><p>Therefore, there were three European species of <italic>Machimosaurus</italic>, which appear to have differed in total body length (<xref ref-type="fig" rid="RSOS140222F46">figure 46</xref>). The Lower Kimmeridgian species <italic>M. buffetauti</italic> is both the geologically oldest and smallest of the three <italic>Machimosaurus</italic> species, with a body length in the range of 5.8–6.2 m. The uppermost Kimmeridgian–lowermost Tithonian species <italic>M. mosae</italic> was larger, with a body length in the range of approximately 6–8 m. The Upper Kimmeridgian–Lower Tithonian species <italic>M. hugii</italic> was the largest of all these species, with a body length exceeding 9 m. Unfortunately, <italic>M. nowackianus</italic> is only known from an anterior dentary, so its body length cannot be reliably estimated. However, this hypothesis requires further testing as new specimens are discovered, because the sample size used here is small (although it includes all known complete skulls).
<fig id="RSOS140222F46" orientation="portrait" position="float"><label>Figure 46.</label><caption><p>Life reconstructions showing the maximum body lengths of the three <italic>Machimosaurus</italic> species present in the Kimmeridgian–Tithonian of Europe. The human diver is 1.8 m in height. The life reconstructions were made by Dmitry Bogdanov.</p></caption><graphic xlink:href="rsos140222-g46"></graphic></fig></p><p>Therefore, at over 9 m in length, <italic>M. hugii</italic> is the largest known crocodylomorph of the Triassic and Jurassic Periods, and until the Cretaceous it was the largest crocodylomorph that had ever existed in Europe.</p></sec><sec id="s7d"><label>8.4</label><title>Hypothetical lifestyles</title><p>Two very different lifestyles have been hypothesized for this genus. Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>] presented evidence that the Leiria <italic>M. hugii</italic> specimen was adapted for living in open seas, whereas Hua [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>] put forward evidence that the <italic>M. mosae</italic> neotype was adapted for living in coastal, high-energy environments. These two hypothetical lifestyles are very different and contradictory. If Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] were correct and there is only one <italic>Machimosaurus</italic> taxon, this would present a major anomaly. However, as we have already shown, <italic>Machimosaurus</italic> taxa differ in craniomandibular, dental and postcranial morphologies. They also differed in geological age, locality and total body length. As such, there is overwhelming evidence that there were three non-sympatric <italic>Machimosaurus</italic> species in Europe during the Kimmeridgian, and there is no reason why these distinct species could not have had distinct lifestyles.</p><p>Krebs [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>] postulated a convincing argument that the Leiria <italic>M. hugii</italic> specimen was well suited to an open sea lifestyle: based on the vertebral zygapophyseal articulations, <italic>M. hugii</italic> would have been well suited to swimming by lateral undulations of the tail, perhaps using the limbs for steering and balancing, and was a fast swimmer. Also, the cervicocranial depressor musculature would have been well developed, as their attachment sites on the skull were enlarged (the basioccipital tubera and the paroccipital processes), which would have greatly assisted <italic>Machimosaurus</italic> in diving [<xref rid="RSOS140222C7" ref-type="bibr">7</xref>]. Krebs' [<xref rid="RSOS140222C6" ref-type="bibr">6</xref>,<xref rid="RSOS140222C7" ref-type="bibr">7</xref>] hypothesis that <italic>M. hugii</italic> was well suited to an open sea lifestyle helps explain this species’ initially odd geographical distribution: the northern Tethyian and eastern Proto-Atlantic margins (Switzerland, Spain and Portugal).</p><p>An additional line of evidence that <italic>M. hugii</italic> was better suited to a more aquatic/pelagic lifestyle comes from dermal bone and osteoderm ornamentation. The ornamentation of dermal bones and osteoderm pits in extant crocodylians are known to be vascularized [<xref rid="RSOS140222C76" ref-type="bibr">76</xref>–<xref rid="RSOS140222C78" ref-type="bibr">78</xref>]. Infrared thermal imaging of basking broad-snouted caimans (<italic>Caiman latirostris</italic>) demonstrates heat exchange between osteoderms and the environment. Moreover, at low temperatures (16°C) the dorsal surface of the snout is one of the warmest regions of the body, whereas at higher temperatures (25°C) it no longer is [<xref rid="RSOS140222C78" ref-type="bibr">78</xref>]. Furthermore, the distribution of dermal ornamentation on the skull of Pseudosuchia in general is consistent with the hypothesis that the most exposed (dorsal) parts of the skull are the most ornamented (M. Fau, M. Laurin and V. de Buffrénil 2014, personal communication). Therefore, vascularized dermal bone/osteoderm dorsal surfaces, heat exchange and blood flow alternation appear to have a role in thermoregulatory terrestrial basking behaviours [<xref rid="RSOS140222C76" ref-type="bibr">76</xref>–<xref rid="RSOS140222C78" ref-type="bibr">78</xref>]. However, as noted above, skull and osteoderm ornamentation of <italic>M. hugii</italic> was reduced (also see [<xref rid="RSOS140222C7" ref-type="bibr">7</xref>, fig. 17]) suggesting it was less vascularized than other <italic>Machimosaurus</italic> species. This supports current research on skull ornamentation in crocodylomorphs, which found an inverse relationship between regional dermal ornamentation levels and aquatic specialization [<xref rid="RSOS140222C79" ref-type="bibr">79</xref>].</p><p>This relationship certainly exists within the thalattosuchian clade Metriorhynchoidea, in which basal members have highly ornamented crania [<xref rid="RSOS140222C80" ref-type="bibr">80</xref>], while within the pelagic clade Metriorhynchidae there is repeated evolution of a ‘smooth’ skull [<xref rid="RSOS140222C81" ref-type="bibr">81</xref>]. Metriorhynchoids have a more extreme shift in thermoregulatory behaviour, as basal metriorhynchoids have osteoderms and retain external mandibular fenestrae (which enables the musculus intramandibularis of extant crocodylians to fix the jaws in a gaping position during mouth-gaping basking behaviour), whereas metriorhynchids lack osteoderms and the external mandibular fenestrae [<xref rid="RSOS140222C80" ref-type="bibr">80</xref>].</p><p>Hua [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>] postulated a convincing lifestyle for the <italic>M. mosae</italic> neotype. The robust ribs, thick and keeled ventral osteoderms, thick gastralia and three sacral vertebrae would have helped <italic>M. mosae</italic> remain in place in a high-energy/turbulent environment [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>]. The small paroccipital processes and basioccipital tubera show that this species was not well suited for diving [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>,<xref rid="RSOS140222C21" ref-type="bibr">21</xref>]. Moreover, the high number of tightly packed dorsal osteoderm pits and the well-ornamented skull suggest that <italic>M. mosae</italic> had a highly vascularized dorsal surface, well suited for terrestrial basking behaviours.</p><p>No hypothetical lifestyle has been postulated for <italic>M. buffetauti</italic>. However, this taxon is similar to <italic>M. mosae</italic> (<xref ref-type="table" rid="RSOS140222TB1">tables 1</xref>–<xref ref-type="table" rid="RSOS140222TB3">3</xref>). However, comparing the figures of the ventral osteoderms and ribs in Hua [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>] to those in Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>] reveals that <italic>M. mosae</italic> was a more robust taxon than <italic>M. buffetauti</italic>. Interestingly, the dermal bone and osteoderm ornamentation of <italic>M. buffetauti</italic> is more similar to <italic>M. hugii</italic> than <italic>M. mosae</italic>, suggesting that this Lower Kimmeridgian species was perhaps intermediate between the two Upper Kimmeridgian extremes. As <italic>M. nowackianus</italic> is only known from an anterior dentary, no hypothetical lifestyle can be postulated.</p></sec></sec><sec sec-type="conclusions" id="s8"><label>9.</label><title>Conclusion</title><p>We here review and clarify the systematics of <italic>Machimosaurus</italic>, one of the most distinctive teleosaurid crocodylomorphs from the Jurassic. We show that <italic>M. mosae</italic> is indeed an available name. We also show that the type specimen (the lectotype) of <italic>M. hugii</italic> is an isolated tooth crown from Switzerland, which is still curated at Solothurn, and that the Portuguese specimen was not considered to be the neotype of this species. We also established <italic>M</italic>. <italic>buffetauti</italic> sp. nov. for the Lower Kimmeridgian specimens from France and Germany, based on a previously described specimen. Most importantly, we demonstrate that there were three species of <italic>Machimosaurus</italic> in the Kimmeridgian (Upper Jurassic) of Europe, and another taxon in Ethiopia. This conclusion is not solely based on tooth counts and cranial biometric ratios, unlike recent revisions of the genus. Our revision uses these characteristics, but expands upon them to include comparative anatomy (craniomandibular, dental and postcranial), body size, hypothetical lifestyle, geological age and geographical range. This holistic approach readily identifies three non-sympatric European species and reveals that potentially contemporaneous taxa were adapted for very different ecosystems. What is surprising, however, is that much of the reasoning we outline here has been long known and thoroughly described, but had not been synthesized.</p><p>One new aspect that helps differentiate <italic>Machimosaurus</italic> species in this paper is postcranial morphology. Prior to description of the <italic>M. buffetauti</italic> holotype by Martin & Vincent [<xref rid="RSOS140222C10" ref-type="bibr">10</xref>], the only postcranial skeleton that was well described and figured was the <italic>M. mosae</italic> neotype [<xref rid="RSOS140222C4" ref-type="bibr">4</xref>]. We have attempted to begin to rectify the long neglected study of teleosaurid postcranial morphology here, as we show for the first time that the postcranial skeletons of <italic>M. buffetauti</italic> and <italic>M. mosae</italic> are distinct (<xref ref-type="table" rid="RSOS140222TB3">table 3</xref>). We recommend that future studies on thalattosuchians do not solely focus on snout length, tooth counts and biometric ratios of skull measurements, but thoroughly investigate craniomandibular and postcranial morphologies.</p><p>As <italic>Machimosaurus</italic> specimens are becoming increasingly abundant, one potentially interesting hypothesis could be investigated with future discoveries. Was <italic>Machimosaurus</italic> in the Kimmeridgian–Lower Tithonian equivalent to the Pliocene–Holocene Asian–Australasian and American subclades of <italic>Crocodylus</italic>? The extant Asian–Australasian <italic>Crocodylus</italic> subclade has one large-bodied taxon well suited to traversing marine barriers (<italic>C. porosus</italic>) and five geographically limited taxa across its range (<italic>C. johnsoni</italic>, <italic>C. mindorensis</italic>, <italic>C. novaeguineae</italic>, <italic>C. palustris</italic> and <italic>C. siamensis</italic>), while the extant American <italic>Crocodylus</italic> subclade has one large-bodied taxon well suited to traversing marine barriers (<italic>C. acutus</italic>) and three geographically limited taxa across its range (<italic>C. intermedius</italic>, <italic>C. moreletii</italic> and <italic>C. rhombifer</italic>) [<xref rid="RSOS140222C82" ref-type="bibr">82</xref>]. In <italic>Machimosaurus</italic>, there is one large-bodied taxon well suited to traversing marine barriers (<italic>M. hugii</italic>), with geographically limited but temporally distinct taxa living in Europe and Ethiopia. Could there be more geographically limited <italic>Machimosaurus</italic> taxa along the margins of Tethys and/or palaeo-islands? It is a potentially interesting hypothesis that will require future study.</p><p>Recent phylogenetic analyses place <italic>Machimosaurus</italic> and ‘<italic>S</italic>.’ <italic>obtusidens</italic> as sister taxa, forming a clade of durophagous/generalist teleosaurids that lived during the Callovian–Tithonian. Their large size, robust craniomandibular and dental morphologies, and broad dietary range suggest ‘<italic>S</italic>.’ <italic>obtusidens</italic> and <italic>Machimosaurus</italic> were an important component of Jurassic shallow marine/brackish ecosystems. We anticipate that future discoveries, especially from outside of Europe, will further elucidate the evolution of this remarkable clade of marine crocodylomorphs.</p></sec></body><back><ack><title>Acknowledgements</title><p>We thank Nathalie Bardet, Eric Buffetaut and Rainer Schoch for discussion on <italic>Machimosaurus</italic> and sharing specimen photographs, Marine Fau and Michel Laurin for discussion on crocodylomorph dermatocranium ornamentation, Nils Knötschke (DFMMh), Eliza Howlett (OUMNH) and Sylvia Humphrey (TWCMS) for collections access, Simão Mateus who photographed the Portuguese isolated teeth, Ron Blakey (<uri xlink:href="http://cpgeosystems.com/">http://cpgeosystems.com/</uri>) for providing the palaeomaps, Sarah Dylewski (Musée de la Princerie) for arranging specimen photography and Dmitry Bogdanov for creating the life reconstructions. We also thank NHMUK image resources for digitizing photograph negatives, photographing and photoshopping the NHMUK specimens. Finally, we thank Andrea Cau and an anonymous reviewer for their constructive comments to an earlier version of this paper. M.T.Y. conceived and drafted the manuscript; S.H. helped conceive and draft the manuscript, and provided papers and the original negatives of the <italic>M. mosae</italic> photographs; L.S. provided collections access and specimen information, arranged specimen photography and digital conversion of photograph negatives and helped draft the manuscript; D.F. prepared and created the figures and helped draft the manuscript; S.L.B. and M.B.A. helped draft the manuscript; S.T., O.M., J.I.R-O. and P.H. provided specimen information, arranged and/or carried out specimen photography and helped draft the manuscript. N.K. and Y.L. provided specimens for loan to M.T.Y. and L.S. for study and photography and helped draft the manuscript. 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