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However, Gavialoidea displays a rich and near-cosmopolitan diversity in the fossil record, with over 40 species currently recognized. Several of these gavialoids are recovered from Cenozoic deposits in the Mediterranean region, with many assigned historically to the genus Tomistoma. However, recent phylogenetic analyses do not typically recover these species as closely related to the false gharial, with several fossil species in need of taxonomic revision. Here, we present a detailed re-evaluation of one of these species, Tomistoma cairense from the Middle Eocene Mokattam Formation of Egypt, based on the holotype skull and several referred specimens. Supplemented by endocranial anatomy revealed via computed tomography (CT), we identify numerous anatomical differences with T. schlegelii and other gavialoids, including the approximately contemporaneous Egyptian species, Eogavialis africanum and Paratomistoma courti. These differences are reinforced through phylogenetic analysis under maximum parsimony, in which T. cairense is recovered as a gavialoid that does not cluster with T. schlegelii or Mediterranean taxa. We erect a new genus name for T. cairense, with the resultant new combination of Herugavialis cairensis diagnosed by several characteristics, including telescoped orbits. An apparent decline in Mediterranean gavialoid diversity during the Late Eocene\u2013Oligocene is potentially driven by a reduction in habitat, as a result of global cooling and declining eustatic sea level. Warming in the Miocene might have facilitated a subsequent rebound in gavialoid diversity, prior to the extirpation of crocodylians in the Mediterranean region in the Late Miocene, which might have been caused by the Messinian Salinity Crisis. The taxonomic revision of additional Mediterranean gavialoid species should facilitate greater insight into their evolutionary history and demise.<\/jats:p>","DOI":"10.1098\/rsos.261173","type":"journal-article","created":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T00:11:38Z","timestamp":1791331898000},"source":"Crossref","is-referenced-by-count":0,"title":["Cranial anatomy of the Middle Eocene Egyptian crocodylian\n                    <i>Herugavialis cairensis<\/i>\n                    provides new insights into gavialoid diversity in the Mediterranean region"],"prefix":"10.1098","volume":"13","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0328-7003","authenticated-orcid":false,"given":"Paul M. J.","family":"Burke","sequence":"first","affiliation":[{"name":"Department of Earth Sciences, University College London 1 , London, UK"},{"name":"Department of Palaeobiology, Naturhistoriska riksmuseet 2 , Stockholm, Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3197-8752","authenticated-orcid":false,"given":"Stephan N. F.","family":"Spiekman","sequence":"additional","affiliation":[{"name":"Staatliches Museum f\u00fcr Naturkunde Stuttgart 3 , Stuttgart, Germany"},{"name":"Department of Paleontology, University of Hohenheim 4 , Stuttgart, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9403-8792","authenticated-orcid":false,"given":"Belal S.","family":"Salem","sequence":"additional","affiliation":[{"name":"Mansoura University Vertebrate Paleontology Center (MUVP), Faculty of Science, Mansoura University 5 , Mansoura 35516, Egypt"},{"name":"Department of Biological Sciences, Ohio University 6 , Athens, Ohio 45701, USA"},{"name":"Department of Geology, Faculty of Science, Benha University 7 , Benha 13518, Egypt"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-4036-2785","authenticated-orcid":false,"given":"Sara","family":"Saber","sequence":"additional","affiliation":[{"name":"Mansoura University Vertebrate Paleontology Center (MUVP), Faculty of Science, Mansoura University 5 , Mansoura 35516, Egypt"},{"name":"Department of Geology, Faculty of Science, Assiut University 8 , Assuit 7515, Egypt"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6584-6312","authenticated-orcid":false,"given":"Hesham M.","family":"Sallam","sequence":"additional","affiliation":[{"name":"Mansoura University Vertebrate Paleontology Center (MUVP), Faculty of Science, Mansoura University 5 , Mansoura 35516, Egypt"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9361-6941","authenticated-orcid":false,"given":"Philip D.","family":"Mannion","sequence":"additional","affiliation":[{"name":"Department of Earth Sciences, University College London 1 , London, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2026,10,7]]},"reference":[{"key":"2026101310265733800_RSOS261173C1","first-page":"133","volume-title":"Crocodiles: status survey and conservation action plan","author":"Bezuijen","year":"2010","edition":"3rd edn."},{"key":"2026101310265733800_RSOS261173C2","first-page":"e.T21981A214287051","volume-title":"Tomistoma schlegelii","author":"Shaney","year":"2023"},{"key":"2026101310265733800_RSOS261173C3","first-page":"315","article-title":"Phylogenetic position of the crocodylian Megadontosuchus arduini and tomistomine palaeobiogeography","volume":"52","author":"Piras","year":"2007","journal-title":"Acta Palaeontol. 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Linnean Soc."},{"key":"2026101310265733800_RSOS261173C6","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1080\/14772019.2014.913078","article-title":"Maroccosuchus zennaroi (Crocodylia: Tomistominae) from the Eocene of Morocco: phylogenetic and palaeobiogeographical implications of the basalmost tomistomine","volume":"13","author":"Jouve","year":"2015","journal-title":"J. Syst. Paleontol."},{"key":"2026101310265733800_RSOS261173C7","doi-asserted-by":"crossref","first-page":"1853","DOI":"10.1080\/14772019.2020.1855603","article-title":"A re-examination of the anatomy and systematics of the tomistomine crocodylians from the Miocene of Italy and Malta","volume":"18","author":"Nicholl","year":"2020","journal-title":"J. Syst. Paleontol."},{"key":"2026101310265733800_RSOS261173C8","doi-asserted-by":"crossref","first-page":"623","DOI":"10.1111\/cla.12372","article-title":"Mosaic nature in the skeleton of East Asian crocodylians fills the morphological gap between \u201cTomistominae\u201d and Gavialinae","volume":"35","author":"Iijima","year":"2019","journal-title":"Cladistics"},{"key":"2026101310265733800_RSOS261173C9","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1002\/ar.25569","article-title":"Endocranial anatomy and phylogenetic position of the crocodylian Eosuchus lerichei from the Late Paleocene of northwestern Europe and potential adaptations for transoceanic dispersal in gavialoids","volume":"308","author":"Burke","year":"2025","journal-title":"Anat. Rec."},{"key":"2026101310265733800_RSOS261173C10","doi-asserted-by":"crossref","first-page":"e12094","DOI":"10.7717\/peerj.12094","article-title":"Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem","volume":"9","author":"Rio","year":"2021","journal-title":"PeerJ"},{"key":"2026101310265733800_RSOS261173C11","first-page":"20 220 380","article-title":"Miocene fossils from the southeastern Pacific shed light on the last radiation of marine crocodylians","volume-title":"Proc. R. Soc. 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Paleontol."},{"key":"2026101310265733800_RSOS261173C18","doi-asserted-by":"crossref","first-page":"927","DOI":"10.1017\/jpa.2023.36","article-title":"Crocodylian princess in Taiwan: revising the taxonomic status of Tomistoma taiwanicus from the Pleistocene of Taiwan and its paleobiogeographic implications","volume":"97","author":"Cho","year":"2023","journal-title":"J. Paleontol."},{"key":"2026101310265733800_RSOS261173C19","doi-asserted-by":"crossref","first-page":"672","DOI":"10.1016\/j.palwor.2017.03.006","article-title":"Maomingosuchus petrolica, a restudy of \u2018Tomistoma\u2019 petrolica Yeh, 1958","volume":"26","author":"Shan","year":"2017","journal-title":"Palaeoworld"},{"key":"2026101310265733800_RSOS261173C20","doi-asserted-by":"crossref","first-page":"67","DOI":"10.1016\/j.palaeo.2004.03.013","article-title":"Italian Cenozoic crocodilians: taxa, timing and palaeobiogeographic implications","volume":"210","author":"Kotsakis","year":"2004","journal-title":"Palaeogeogr. 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Rec."},{"key":"2026101310265733800_RSOS261173C24","first-page":"251","article-title":"New tomistomine crocodylian from the Middle Eocene (Bartonian) of Wadi Hitan, Fayum Province, Egypt","volume":"30","author":"Brochu","year":"2000","journal-title":"Contrib. Mus. Paleontol."},{"key":"2026101310265733800_RSOS261173C25","doi-asserted-by":"crossref","first-page":"e2604600","DOI":"10.1080\/02724634.2025.2604600","article-title":"Revision of the longirostrine crocodylian Thoracosaurus isorhynchus from the Maastrichtian\u2013Danian of northwestern Europe","volume":"45","author":"Boerman","year":"2026","journal-title":"J. Vertebr. Paleontol."},{"key":"2026101310265733800_RSOS261173C26","first-page":"137","article-title":"Late Cenozoic crocodiles (Reptilia: Crocodylidae) from the Western Rift, Uganda","volume":"2","author":"Pickford","year":"1994","journal-title":"Geol. Palaeobiol. Albert. Rift Valley, Uganda-Zaire"},{"key":"2026101310265733800_RSOS261173C27","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1111\/cla.12524","article-title":"TNT version 1.6, with a graphical interface for MacOS and Linux, including new routines in parallel","volume":"39","author":"Goloboff","year":"2023","journal-title":"Cladistics"},{"key":"2026101310265733800_RSOS261173C28","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1111\/cla.12047","article-title":"Extended implied weighting","volume":"30","author":"Goloboff","year":"2014","journal-title":"Cladistics"},{"key":"2026101310265733800_RSOS261173C29","doi-asserted-by":"crossref","first-page":"379","DOI":"10.1111\/pala.12142","article-title":"strap: an R package for plotting phylogenies against stratigraphy and assessing their stratigraphic congruence","volume":"58","author":"Bell","year":"2015","journal-title":"Palaeontology"},{"key":"2026101310265733800_RSOS261173C30","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1144\/GSL.JGS.1875.031.01-04.29","article-title":"On Stagonolepis robertsoni, and on the evolution of the Crocodilia","volume":"31","author":"Huxley","year":"1875","journal-title":"Q. J. Geol. Soc."},{"key":"2026101310265733800_RSOS261173C31","first-page":"1033","author":"Gmelin","year":"1789"},{"key":"2026101310265733800_RSOS261173C32","volume-title":"Second bibliography and catalogue of the fossil vertebrata of North America","author":"Hay","year":"1930"},{"key":"2026101310265733800_RSOS261173C33","volume-title":"Sphinx: history of a monument","author":"Zivie-Coche","year":"2004"},{"key":"2026101310265733800_RSOS261173C34","first-page":"19","article-title":"Geologic study of the Sphinx","volume":"127","author":"Gauri","year":"1984","journal-title":"Am. Res. Center Egypt Newslett."},{"key":"2026101310265733800_RSOS261173C35","first-page":"1","article-title":"Marine mammals (Cetacea and Sirenia) from the Eocene of Gebel Mokattam and Fayum, Egypt: stratigraphy, age and paleoenvironments","volume":"30","author":"Gingerich","year":"1992","journal-title":"Papers Paleontol."},{"key":"2026101310265733800_RSOS261173C36","first-page":"3","article-title":"Tomistoma lusitanica, crocodilien du Mioc\u00e9ne du Portugal","volume":"9","author":"Antunes","year":"1961","journal-title":"Rev. Facult. Ci\u00eanc. Univ. Lisb\u00f5a"},{"key":"2026101310265733800_RSOS261173C37","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1666\/0022-3360(2006)080[0162:OAPSOE]2.0.CO;2","article-title":"Osteology and phylogenetic significance of Eosuchus minor (Marsh, 1870) new combination, a longirostrine crocodylian from the late Paleocene of North America","volume":"80","author":"Brochu","year":"2006","journal-title":"J. Paleontol."},{"key":"2026101310265733800_RSOS261173C38","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1093\/zoolinnean\/zly064","article-title":"A new eusuchian crocodylomorph from the Cenomanian (Late Cretaceous) of Portugal reveals novel implications on the origin of Crocodylia","volume":"186","author":"Mateus","year":"2019","journal-title":"Zool. J. Linnean Soc."},{"key":"2026101310265733800_RSOS261173C39","doi-asserted-by":"crossref","first-page":"12 158","DOI":"10.1038\/s41598-021-91717-y","article-title":"First record of a tomistomine crocodylian from Australia","volume":"11","author":"Ristevski","year":"2021","journal-title":"Sci. Rep."},{"key":"2026101310265733800_RSOS261173C40","doi-asserted-by":"crossref","first-page":"1049","DOI":"10.1080\/14772019.2018.1495275","article-title":"New Miocene Caribbean gavialoids and patterns of longirostry in crocodylians","volume":"17","author":"Salas-Gismondi","year":"2019","journal-title":"J. Syst. 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Z."},{"key":"2026101310265733800_RSOS261173C47","volume-title":"A descriptive catalogue of the tertiary vertebrata of the Fayum, Egypt","author":"Andrews","year":"1906"},{"key":"2026101310265733800_RSOS261173C48","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1671\/0272-4634(2004)024[0610:ANLCGC]2.0.CO;2","article-title":"A new Late Cretaceous gavialoid crocodylian from eastern North America and the phylogenetic relationships of thoracosaurs","volume":"24","author":"Brochu","year":"2004","journal-title":"J. Vertebr. Paleontol."},{"key":"2026101310265733800_RSOS261173C49","first-page":"61","article-title":"A gavialoid crocodylian from the Lower Miocene of Venezuela","volume":"71","author":"Brochu","year":"2004","journal-title":"Spec. Pap. Palaeontol."},{"key":"2026101310265733800_RSOS261173C50","first-page":"565","article-title":"Anatomy and phylogeny of the gavialoid crocodylian Eosuchus lerichei from the Paleogene of Europe","volume":"50","author":"Delfino","year":"2005","journal-title":"Acta Palaeontol. Pol."},{"key":"2026101310265733800_RSOS261173C51","doi-asserted-by":"crossref","first-page":"e0152453","DOI":"10.1371\/journal.pone.0152453","article-title":"A new 13 million year old gavialoid crocodylian from proto-Amazonian mega-wetlands reveals parallel evolutionary trends in skull shape linked to longirostry","volume":"11","author":"Salas-Gismondi","year":"2016","journal-title":"PloS one"},{"key":"2026101310265733800_RSOS261173C52","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1007\/BF02988368","article-title":"The cranium of Piscogavialis jugaliperforatus n. gen., n. sp. (Gavialidae, Crocodylia) from the Miocene of Peru","volume":"72","author":"Kraus","year":"1998","journal-title":"Pal\u00e4ontol. Z."},{"key":"2026101310265733800_RSOS261173C53","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1111\/zoj.12357","article-title":"A new basal tomistomine (Crocodylia, Crocodyloidea) from Issel (Middle Eocene; France): palaeobiogeography of basal tomistomines and palaeogeographic consequences","volume":"177","author":"Jouve","year":"2016","journal-title":"Zool. J. Linnean Soc."},{"key":"2026101310265733800_RSOS261173C54","first-page":"20 220 085","article-title":"An intermediate crocodylian linking two extant gharials from the Bronze Age of China and its human-induced extinction","volume-title":"Proc. R. Soc. B","author":"Iijima","year":"2022"},{"key":"2026101310265733800_RSOS261173C55","doi-asserted-by":"crossref","first-page":"1146","DOI":"10.1111\/joa.13836","article-title":"Neuroanatomy of the crocodylomorph Portugalosuchus azenhae from the late Cretaceous of Portugal","volume":"242","author":"Pu\u00e9rtolas-Pascual","year":"2023","journal-title":"J. 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