{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T01:07:25Z","timestamp":1790730445267,"version":"4.1.0"},"reference-count":75,"publisher":"Wiley","issue":"8","license":[{"start":{"date-parts":[[2023,12,20]],"date-time":"2023-12-20T00:00:00Z","timestamp":1703030400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","award":["ANR19\u2010CE31\u20100006\u201001"],"award-info":[{"award-number":["ANR19\u2010CE31\u20100006\u201001"]}],"id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["anatomypubs.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Anatomical Record"],"published-print":{"date-parts":[[2024,8]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p><jats:italic>Voay robustus<\/jats:italic>, the extinct Malagasy \u201chorned\u201d crocodile, was originally considered to be the only crocodylian representative in Madagascar during most part of the Holocene. However, Malagasy crocodylian remains have had confused taxonomic attributions and recent studies have underlined that <jats:italic>Crocodylus<\/jats:italic> and <jats:italic>Voay<\/jats:italic> populations coexisted on the island for at least 7500\u2009years. Here, we describe the inner braincase anatomy of <jats:italic>Voay robustus<\/jats:italic> using x\u2010ray computed tomography on four specimens, to provide new anatomical information that distinguishes <jats:italic>Voay<\/jats:italic> from <jats:italic>Crocodylus<\/jats:italic>, especially features of the brain endocast and the paratympanic sinuses. Geometric morphometric analyses are performed on 3D models of the internal organs to compare statistically <jats:italic>Voay<\/jats:italic> with a subset of extant Crocodylidae. Following these comparisons, we build an endocranial morphological matrix to discuss the proposed phylogenetic affinities of <jats:italic>Voay<\/jats:italic> with Osteolaeminae from an endocranial point of view. Additionally, we discuss the use of internal characters in systematic studies and find that they can have a major impact on morphological analyses. Finally, new radiocarbon data on <jats:italic>Voay<\/jats:italic> and subfossil <jats:italic>Crocodylus<\/jats:italic> specimens are recovered between 2010 and 2750\u2009cal BP, which confirm the cohabitation of the two species in the same area for a long period of time. We thus assess several extinction scenarios, and propose a slightly different ecology of <jats:italic>Voay<\/jats:italic> compared to <jats:italic>Crocodylus<\/jats:italic>, which could have allowed habitat partitioning on the island. Our approach complements information obtained from previous molecular and morphological phylogenies, as well as previous radiocarbon dating, together revealing past diversity and faunal turnovers in Madagascar.<\/jats:p>","DOI":"10.1002\/ar.25367","type":"journal-article","created":{"date-parts":[[2023,12,20]],"date-time":"2023-12-20T06:22:56Z","timestamp":1703053376000},"page":"2749-2786","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":12,"title":["Neuroanatomy and pneumaticity of the extinct Malagasy \u201chorned\u201d crocodile <i>Voay robustus<\/i> and its implications for crocodylid phylogeny and palaeoecology"],"prefix":"10.1002","volume":"307","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5831-764X","authenticated-orcid":false,"given":"Gwendal","family":"Perrichon","sequence":"first","affiliation":[{"name":"CNRS UMR 5276, Universit\u00e9 Claude Bernard Lyon 1, ENS de Lyon, Laboratoire de G\u00e9ologie de Lyon\u2010Terre, Plan\u00e8tes, Environnement  Villeurbanne France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6216-2721","authenticated-orcid":false,"given":"Yohan","family":"Pochat\u2010Cottilloux","sequence":"additional","affiliation":[{"name":"CNRS UMR 5276, Universit\u00e9 Claude Bernard Lyon 1, ENS de Lyon, Laboratoire de G\u00e9ologie de Lyon\u2010Terre, Plan\u00e8tes, Environnement  Villeurbanne France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Davide","family":"Conedera","sequence":"additional","affiliation":[{"name":"CNRS UMR 5276, Universit\u00e9 Claude Bernard Lyon 1, ENS de Lyon, Laboratoire de G\u00e9ologie de Lyon\u2010Terre, Plan\u00e8tes, Environnement  Villeurbanne France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6741-0352","authenticated-orcid":false,"given":"Pascale","family":"Richardin","sequence":"additional","affiliation":[{"name":"Centre de Recherche et de Restauration des Mus\u00e9es de France (C2RMF), Palais du Louvre, Porte des Lions  Paris France"},{"name":"CNRS\u2010UMR 8068, Technologie Ethnologie des Mondes Pr\u00e9historiques (TEMPS), Universit\u00e9 Paris Nanterre  Nanterre Cedex France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8315-1458","authenticated-orcid":false,"given":"Vincent","family":"Fernandez","sequence":"additional","affiliation":[{"name":"Imaging and Analysis Centre, The Natural History Museum  London UK"},{"name":"European Synchrotron Radiation Facility  Grenoble France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8701-5421","authenticated-orcid":false,"given":"Lionel","family":"Hautier","sequence":"additional","affiliation":[{"name":"Institut des Sciences de l'\u00c9volution, Universit\u00e9 Montpellier, CNRS, IRD, EPHE  Montpellier France"},{"name":"Mammal Section, Life Sciences, Vertebrate Division, The Natural History Museum  London UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9159-645X","authenticated-orcid":false,"given":"Jeremy E.","family":"Martin","sequence":"additional","affiliation":[{"name":"CNRS UMR 5276, Universit\u00e9 Claude Bernard Lyon 1, ENS de Lyon, Laboratoire de G\u00e9ologie de Lyon\u2010Terre, Plan\u00e8tes, Environnement  Villeurbanne France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2023,12,20]]},"reference":[{"key":"e_1_2_14_2_1","article-title":"Geometric morphometric analyses of 2D\/3D landmark data","author":"Adams D.","year":"2022","journal-title":"R Package Version 3.2"},{"key":"e_1_2_14_3_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1442-9993.2001.01070.pp.x"},{"issue":"3","key":"e_1_2_14_4_1","first-page":"186","article-title":"Sur une t\u00eate osseuse de Crocodile de Madagascar (Crocodilus robustus Grand. et Vaill.)","volume":"1","author":"Angel F.","year":"1929","journal-title":"Bulletin du Museum National d'histoire Naturelle"},{"issue":"14","key":"e_1_2_14_5_1","first-page":"488","article-title":"Vertebrata from Madagascar: Amphibia and Reptilia","volume":"16","author":"Barbour T.","year":"1918","journal-title":"Bulletin of the Museum of Comparative Zoology"},{"key":"e_1_2_14_6_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-46959-2"},{"key":"e_1_2_14_7_1","doi-asserted-by":"publisher","DOI":"10.1002\/mmng.200800007"},{"key":"e_1_2_14_8_1","doi-asserted-by":"publisher","DOI":"10.1093\/sysbio\/46.3.479"},{"issue":"3","key":"e_1_2_14_9_1","first-page":"564","article-title":"Crocodylian snouts in space and time: Phylogenetic approaches toward adaptive radiation","volume":"41","author":"Brochu C. A.","year":"2001","journal-title":"American Zoologist"},{"key":"e_1_2_14_10_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.earth.31.100901.141308"},{"key":"e_1_2_14_11_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.2007.00315.x"},{"key":"e_1_2_14_12_1","doi-asserted-by":"publisher","DOI":"10.1017\/S1755691013000200"},{"key":"e_1_2_14_13_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.25005"},{"key":"e_1_2_14_14_1","doi-asserted-by":"publisher","DOI":"10.1080\/02724634.2012.633585"},{"key":"e_1_2_14_15_1","doi-asserted-by":"publisher","DOI":"10.1080\/02724634.2012.652324"},{"key":"e_1_2_14_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2021.03.086"},{"key":"e_1_2_14_17_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jhevol.2004.05.005"},{"key":"e_1_2_14_18_1","doi-asserted-by":"publisher","DOI":"10.1006\/qres.1993.1060"},{"key":"e_1_2_14_19_1","doi-asserted-by":"publisher","DOI":"10.18814\/epiiugs\/2013\/v36i3\/002"},{"key":"e_1_2_14_20_1","doi-asserted-by":"publisher","DOI":"10.2307\/1438813"},{"key":"e_1_2_14_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.quascirev.2010.06.030"},{"key":"e_1_2_14_22_1","doi-asserted-by":"publisher","DOI":"10.1177\/0959683613484675"},{"key":"e_1_2_14_23_1","first-page":"8","article-title":"Sur les diff\u00e9rentes esp\u00e8ces de crocodiles vivants et sur leurs caract\u00e8res distinctifs","volume":"10","author":"Cuvier","year":"1807","journal-title":"Annales du Mus\u00e9um National d'Histoire Naturelle"},{"key":"e_1_2_14_24_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-020-68482-5"},{"key":"e_1_2_14_25_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.quascirev.2019.105878"},{"key":"e_1_2_14_26_1","first-page":"127","volume-title":"Crocodiles","author":"Eaton M. J.","year":"2010"},{"key":"e_1_2_14_27_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.10201"},{"key":"e_1_2_14_28_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.22666"},{"key":"e_1_2_14_29_1","doi-asserted-by":"publisher","DOI":"10.26879\/527"},{"key":"e_1_2_14_30_1","doi-asserted-by":"publisher","DOI":"10.3389\/fevo.2021.742203"},{"key":"e_1_2_14_31_1","doi-asserted-by":"publisher","DOI":"10.1111\/cla.12524"},{"key":"e_1_2_14_32_1","first-page":"150","article-title":"Sur le crocodile fossile d'Amboulintsatre (Madagascar)","volume":"75","author":"Grandidier A.","year":"1872","journal-title":"Comptes Rendus Hebdomadaires des Seances de L'Academie des Sciences"},{"key":"e_1_2_14_33_1","doi-asserted-by":"publisher","DOI":"10.1093\/zoolinnean\/zlz117"},{"key":"e_1_2_14_34_1","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.aat6925"},{"key":"e_1_2_14_35_1","doi-asserted-by":"publisher","DOI":"10.1080\/10635150390197028"},{"key":"e_1_2_14_36_1","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-021-02017-0"},{"key":"e_1_2_14_37_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-294X.2011.05245.x"},{"key":"e_1_2_14_38_1","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.5686"},{"key":"e_1_2_14_39_1","doi-asserted-by":"publisher","DOI":"10.1017\/RDC.2020.59"},{"key":"e_1_2_14_40_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.24579"},{"key":"e_1_2_14_41_1","first-page":"e.T45433088A301","volume-title":"Crocodylus niloticus","author":"Isberg S.","year":"2019"},{"key":"e_1_2_14_42_1","doi-asserted-by":"publisher","DOI":"10.1145\/2487228.2487237"},{"key":"e_1_2_14_43_1","doi-asserted-by":"publisher","DOI":"10.1017\/S0266467400006490"},{"key":"e_1_2_14_44_1","volume-title":"anatomy, development and evolution of braincase in Crocodylomorpha (Diapsida: Archosauria)","author":"Kuzmin I. T.","year":"2022"},{"key":"e_1_2_14_45_1","doi-asserted-by":"publisher","DOI":"10.1111\/joa.13490"},{"key":"e_1_2_14_46_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.24550"},{"key":"e_1_2_14_47_1","doi-asserted-by":"publisher","DOI":"10.1080\/02724634.2021.2063058"},{"key":"e_1_2_14_48_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2011.03.026"},{"key":"e_1_2_14_49_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2020.106924"},{"key":"e_1_2_14_50_1","first-page":"25","article-title":"Description of a skull of the extinct Madagascar crocodile, Crocodilus robustus Vaillant and Grandidier","volume":"44","author":"Mook C. C.","year":"1921","journal-title":"Bulletin of the AMNH"},{"key":"e_1_2_14_51_1","doi-asserted-by":"publisher","DOI":"10.1643\/CG-19-240"},{"key":"e_1_2_14_52_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1558-5646.2011.01373.x"},{"key":"e_1_2_14_53_1","first-page":"60","article-title":"Report on British fossil reptiles, part II","volume":"1841","author":"Owen R.","year":"1842","journal-title":"Report for the British Association for the Advancement of Science"},{"key":"e_1_2_14_54_1","doi-asserted-by":"publisher","DOI":"10.1093\/zoolinnean\/zlaa074"},{"key":"e_1_2_14_55_1","doi-asserted-by":"publisher","DOI":"10.1111\/joa.13830"},{"key":"e_1_2_14_56_1","doi-asserted-by":"publisher","DOI":"10.18563\/journal.m3.205"},{"key":"e_1_2_14_57_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.24826"},{"key":"e_1_2_14_58_1","doi-asserted-by":"publisher","DOI":"10.1111\/joa.13887"},{"key":"e_1_2_14_59_1","doi-asserted-by":"publisher","DOI":"10.1111\/joa.12539"},{"key":"e_1_2_14_60_1","volume-title":"R: A language and environment for statistical computing","author":"R Core Team","year":"2021"},{"key":"e_1_2_14_61_1","first-page":"94","article-title":"Description of the subfossil crocodylians from a new late Pleistocene subfossil site (Tsaramody, Sambaina Basin) in Central Madagascar","volume":"15","author":"Rakotozandry R.","year":"2021","journal-title":"Malagasy Nature"},{"key":"e_1_2_14_62_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ecolsys.27.1.83"},{"key":"e_1_2_14_63_1","doi-asserted-by":"publisher","DOI":"10.1017\/RDC.2016.102"},{"key":"e_1_2_14_64_1","doi-asserted-by":"publisher","DOI":"10.2307\/1447477"},{"key":"e_1_2_14_65_1","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.12094"},{"key":"e_1_2_14_66_1","doi-asserted-by":"publisher","DOI":"10.1002\/jqs.3096"},{"key":"e_1_2_14_67_1","doi-asserted-by":"publisher","DOI":"10.1038\/nmeth.2089"},{"key":"e_1_2_14_68_1","doi-asserted-by":"publisher","DOI":"10.1111\/aje.12365"},{"key":"e_1_2_14_69_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.24932"},{"key":"e_1_2_14_70_1","volume-title":"Histoire physique, naturelle, et politique de Madagascar. Volume XVII, Histoire naturelle des reptiles; premi\u00e8re partie: crocodiles et tortues","author":"Vaillant L.","year":"1910"},{"key":"e_1_2_14_71_1","first-page":"260","volume-title":"Present knowledge on the west African slender\u2010snouted crocodile, Crocodylus cataphractus Cuvier 1824 and the west African dwarf crocodile Osteolaemus tetraspis cope 1861, Crocodiles. Their ecology, management, and conservation","author":"Waitkuwait W. E.","year":"1986"},{"key":"e_1_2_14_72_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2008.1390"},{"key":"e_1_2_14_73_1","doi-asserted-by":"publisher","DOI":"10.1111\/joa.12918"},{"key":"e_1_2_14_74_1","volume-title":"Fossil hunting in Madagascar","author":"White E. I.","year":"1930"},{"key":"e_1_2_14_75_1","doi-asserted-by":"publisher","DOI":"10.1002\/wics.147"},{"key":"e_1_2_14_76_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.24704"}],"container-title":["The Anatomical Record"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ar.25367","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T14:51:42Z","timestamp":1759935102000},"score":1,"resource":{"primary":{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/10.1002\/ar.25367"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,12,20]]},"references-count":75,"journal-issue":{"issue":"8","published-print":{"date-parts":[[2024,8]]}},"alternative-id":["10.1002\/ar.25367"],"URL":"https:\/\/doi.org\/10.1002\/ar.25367","archive":["Portico"],"relation":{},"ISSN":["1932-8486","1932-8494"],"issn-type":[{"value":"1932-8486","type":"print"},{"value":"1932-8494","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,12,20]]},"assertion":[{"value":"2023-06-22","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2023-11-29","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2023-12-20","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}