{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T23:09:57Z","timestamp":1790809797198,"version":"4.1.0"},"reference-count":88,"publisher":"Oxford University Press (OUP)","issue":"2","license":[{"start":{"date-parts":[[2019,5,23]],"date-time":"2019-05-23T00:00:00Z","timestamp":1558569600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"}],"funder":[{"DOI":"10.13039\/100010663","name":"European Research Council","doi-asserted-by":"publisher","award":["STG-2014-637171"],"award-info":[{"award-number":["STG-2014-637171"]}],"id":[{"id":"10.13039\/100010663","id-type":"DOI","asserted-by":"publisher"}]},{"name":"SYNTHESYS","award":["FR-TAF-5635"],"award-info":[{"award-number":["FR-TAF-5635"]}]},{"name":"Macaulay family endowment to the AMNH","award":["NSF DEB-1457181"],"award-info":[{"award-number":["NSF DEB-1457181"]}]},{"DOI":"10.13039\/100000001","name":"NSF","doi-asserted-by":"publisher","award":["IOS-1050154"],"award-info":[{"award-number":["IOS-1050154"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"NSF","doi-asserted-by":"publisher","award":["IOS-1456503"],"award-info":[{"award-number":["IOS-1456503"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"NSF","doi-asserted-by":"publisher","award":["EAR-1349825"],"award-info":[{"award-number":["EAR-1349825"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"NSF","doi-asserted-by":"publisher","award":["EAR-1525915"],"award-info":[{"award-number":["EAR-1525915"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2019,8,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:p>Complex structures, like the vertebrate skull, are composed of numerous elements or traits that must develop and evolve in a coordinated manner to achieve multiple functions. The strength of association among phenotypic traits (i.e., integration), and their organization into highly-correlated, semi-independent subunits termed modules, is a result of the pleiotropic and genetic correlations that generate traits. As such, patterns of integration and modularity are thought to be key factors constraining or facilitating the evolution of phenotypic disparity by influencing the patterns of variation upon which selection can act. It is often hypothesized that selection can reshape patterns of integration, parceling single structures into multiple modules or merging ancestrally semi-independent traits into a strongly correlated unit. However, evolutionary shifts in patterns of trait integration are seldom assessed in a unified quantitative framework. Here, we quantify patterns of evolutionary integration among regions of the archosaur skull to investigate whether patterns of cranial integration are conserved or variable across this diverse group. Using high-dimensional geometric morphometric data from 3D surface scans and computed tomography scans of modern birds (n\u2009=\u2009352), fossil non-avian dinosaurs (n\u2009=\u200927), and modern and fossil mesoeucrocodylians (n\u2009=\u200938), we demonstrate that some aspects of cranial integration are conserved across these taxonomic groups, despite their major differences in cranial form, function, and development. All three groups are highly modular and consistently exhibit high integration within the occipital region. However, there are also substantial divergences in correlation patterns. Birds uniquely exhibit high correlation between the pterygoid and quadrate, components of the cranial kinesis apparatus, whereas the non-avian dinosaur quadrate is more closely associated with the jugal and quadratojugal. Mesoeucrocodylians exhibit a slightly more integrated facial skeleton overall than the other grades. Overall, patterns of trait integration are shown to be stable among archosaurs, which is surprising given the cranial diversity exhibited by the clade. At the same time, evolutionary innovations such as cranial kinesis that reorganize the structure and function of complex traits can result in modifications of trait correlations and modularity.<\/jats:p>","DOI":"10.1093\/icb\/icz052","type":"journal-article","created":{"date-parts":[[2019,5,13]],"date-time":"2019-05-13T19:12:28Z","timestamp":1557774748000},"page":"371-382","source":"Crossref","is-referenced-by-count":72,"title":["Evolutionary Integration and Modularity in the Archosaur Cranium"],"prefix":"10.1093","volume":"59","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9201-9213","authenticated-orcid":false,"given":"Ryan N","family":"Felice","sequence":"first","affiliation":[{"name":"Centre for Integrative Anatomy, Department of Cell and Developmental Biology, University College London, London, WC1E 6BT, UK"},{"name":"Life Sciences Department, Vertebrates Division, Natural History Museum, London, SW7 5BD, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Akinobu","family":"Watanabe","sequence":"first","affiliation":[{"name":"Life Sciences Department, Vertebrates Division, Natural History Museum, London, SW7 5BD, UK"},{"name":"Department of Anatomy, New York Institute of Technology College of Osteopathic Medicine, Old Westbury, NY 11568, USA"},{"name":"Division of Paleontology, American Museum of Natural History, New York, NY 10024, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrew R","family":"Cuff","sequence":"first","affiliation":[{"name":"Structure and Motion Laboratory, Department of Comparative Biomedical Sciences, Royal Veterinary College, Hawkshead Lane, North Mymms, Hertfordshire, AL9 7TA, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eve","family":"Noirault","sequence":"first","affiliation":[{"name":"Life Sciences Department, Vertebrates Division, Natural History Museum, London, SW7 5BD, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Diego","family":"Pol","sequence":"first","affiliation":[{"name":"CONICET. Museo Paleontol\u00f3gico Egidio Feruglio, Av. Fontana 140, Trelew, Chubut, U9100GYO, Argentina"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lawrence M","family":"Witmer","sequence":"first","affiliation":[{"name":"Department of Biomedical Sciences, Ohio University Heritage College of Osteopathic Medicine, Athens, OH, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mark A","family":"Norell","sequence":"first","affiliation":[{"name":"Division of Paleontology, American Museum of Natural History, New York, NY 10024, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Patrick M","family":"O'Connor","sequence":"first","affiliation":[{"name":"Department of Biomedical Sciences, Ohio University Heritage College of Osteopathic Medicine, Athens, OH, USA"},{"name":"Ohio Center for Ecology and Evolutionary Studies, Ohio University, Athens, OH, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anjali","family":"Goswami","sequence":"first","affiliation":[{"name":"Life Sciences Department, Vertebrates Division, Natural History Museum, London, SW7 5BD, UK"},{"name":"Department of Genetics, Evolution, and Environment, University College London, London, WC1E 6BT, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2019,5,23]]},"reference":[{"key":"2020052220095517300_icz052-B1","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1111\/2041-210X.12511","article-title":"Evaluating modularity in morphometric data: challenges with the RV coefficient and a new test measure","volume":"7","author":"Adams","year":"2016","journal-title":"Methods Ecol Evol"},{"key":"2020052220095517300_icz052-B2","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1111\/2041-210X.12035","article-title":"geomorph: an r package for the collection and analysis of geometric morphometric shape data","volume":"4","author":"Adams","year":"2013","journal-title":"Methods Ecol Evol"},{"key":"2020052220095517300_icz052-B3","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1111\/ede.12190","article-title":"Competing influences on morphological modularity in biomechanical systems: a case study in mantis shrimp: morphological covariation","volume":"18","author":"Anderson","year":"2016","journal-title":"Evol Dev"},{"key":"2020052220095517300_icz052-B4","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1111\/j.2041-210X.2012.00223.x","article-title":"paleotree\u202f: an R package for paleontological and phylogenetic analyses of evolution","volume":"3","author":"Bapst","year":"2012","journal-title":"Methods Ecol Evol"},{"key":"2020052220095517300_icz052-B5","doi-asserted-by":"crossref","DOI":"10.1186\/s12862-018-1342-7","article-title":"Morphological evolution and modularity of the caecilian skull","volume":"19","author":"Bardua","year":"2019","journal-title":"BMC Evol Biol"},{"key":"2020052220095517300_icz052-B6","doi-asserted-by":"crossref","first-page":"501","DOI":"10.1038\/nature21700","article-title":"A new hypothesis of dinosaur relationships and early dinosaur evolution","volume":"543","author":"Baron","year":"2017","journal-title":"Nature"},{"key":"2020052220095517300_icz052-B7","first-page":"45","volume-title":"Handbook of avian anatomy: nomina anatomica avium.","author":"Baumel","year":"1993"},{"key":"2020052220095517300_icz052-B8","doi-asserted-by":"crossref","first-page":"20131780.","DOI":"10.1098\/rspb.2013.1780","article-title":"Rates of dinosaur limb evolution provide evidence for exceptional radiation in Mesozoic birds","volume":"280","author":"Benson","year":"2013","journal-title":"Proc R Soc Lond B Biol Sci"},{"key":"2020052220095517300_icz052-B9","first-page":"295","volume-title":"The phylogeny and classification of the tetrapods, Volume 1: amphibians, reptiles, birds. 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