{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,24]],"date-time":"2026-10-24T15:11:16Z","timestamp":1792854676275,"version":"4.5.0"},"reference-count":53,"publisher":"Public Library of Science (PLoS)","issue":"8","license":[{"start":{"date-parts":[[2011,8,10]],"date-time":"2011-08-10T00:00:00Z","timestamp":1312934400000},"content-version":"unspecified","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"DOI":"10.1371\/journal.pone.0023393","type":"journal-article","created":{"date-parts":[[2011,8,10]],"date-time":"2011-08-10T21:05:17Z","timestamp":1313010317000},"page":"e23393","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":52,"title":["Variation, Variability, and the Origin of the Avian Endocranium: Insights from the Anatomy of Alioramus altai (Theropoda: Tyrannosauroidea)"],"prefix":"10.1371","volume":"6","author":[{"given":"Gabe S.","family":"Bever","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stephen L.","family":"Brusatte","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amy M.","family":"Balanoff","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mark A.","family":"Norell","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2011,8,10]]},"reference":[{"key":"ref1","doi-asserted-by":"crossref","first-page":"967","DOI":"10.2307\/2410639","article-title":"Complex adaptations and the evolution of evolvability.","volume":"50","author":"GP Wagner","year":"1996","journal-title":"Evolution"},{"key":"ref2","first-page":"1","article-title":"Characters, units and natural kinds: an introduction.","author":"GP Wagner","year":"2001"},{"key":"ref3","doi-asserted-by":"crossref","first-page":"8420","DOI":"10.1073\/pnas.95.15.8420","article-title":"Evolvability.","volume":"95","author":"M Kirscher","year":"1998","journal-title":"Proceedings of the National Academy of Sciences, USA"},{"key":"ref4","first-page":"384","article-title":"Scaling, allometry, and skull design.","author":"SB Emerson","year":"1993"},{"key":"ref5","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1206\/330.1","article-title":"Postnatal ontogeny of the skull in the extant North American turtle <italic>Sternotherus odoratus<\/italic> (Cryptodira: Kinosternidae).","volume":"330","author":"GS Bever","year":"2009","journal-title":"Bulletin of the American Museum of Natural History"},{"key":"ref6","doi-asserted-by":"crossref","first-page":"1481","DOI":"10.1126\/science.1193304","article-title":"Tyrannosaur paleobiology: new research on ancient exemplar organisms.","volume":"329","author":"SL Brusatte","year":"2010","journal-title":"Science"},{"key":"ref7","doi-asserted-by":"crossref","first-page":"1100","DOI":"10.1017\/S0022336000026706","article-title":"The phylogenetic position of the Tyrannosauridae: implications for theropod systematics.","volume":"68","author":"TR Holtz Jr","year":"1994","journal-title":"Journal of Paleontology"},{"key":"ref8","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1017\/S1477201907002143","article-title":"A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda).","volume":"5","author":"P Senter","year":"2007","journal-title":"Journal of Systematic Palaeontology"},{"key":"ref9","doi-asserted-by":"crossref","first-page":"1378","DOI":"10.1126\/science.1144066","article-title":"A basal dromaeosaurid and size evolution preceeding avian flight.","volume":"317","author":"AH Turner","year":"2007","journal-title":"Science"},{"key":"ref10","first-page":"3505","article-title":"A new North American therizinosaurid and the role of herbivory in \u2018predatory\u2019 dinosaur evolution.","volume":"276","author":"LE Zanno","year":"2009","journal-title":"Proceedings of the National Academy of Sciences, Series B"},{"key":"ref11","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1126\/science.1182143","article-title":"A basal alvarezsauroid theropod from the Early Late Jurassic of Xinjiang, China.","volume":"327","author":"JN Choiniere","year":"2010","journal-title":"Science"},{"key":"ref12","doi-asserted-by":"crossref","first-page":"15357","DOI":"10.1073\/pnas.1006970107","article-title":"An aberrant island-dwelling theropod dinosaur from the Late Cretaceous of Romania.","volume":"107","author":"Z Csiki","year":"2010","journal-title":"Proceedings of the National Academy of Sciences, USA"},{"key":"ref13","first-page":"7","article-title":"Feathered dinosaurs, flying dinosaurs, crown dinosaurs, and the name \u201cAves\u201d.","author":"J Gauthier","year":"2001"},{"key":"ref14","first-page":"1","article-title":"Crania of <italic>Tyrannosaurus<\/italic> and <italic>Allosaurus<\/italic>.","volume":"1","author":"HF Osborn","year":"1912","journal-title":"Memoirs of the American Museum of Natural History"},{"key":"ref15","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1086\/282627","article-title":"Brain evolution and dinosaur brains.","volume":"103","author":"HT Jerison","year":"1969","journal-title":"American Naturalist"},{"key":"ref16","doi-asserted-by":"crossref","first-page":"1","DOI":"10.2307\/3889334","article-title":"Osteology of <italic>Tyrannosaurus rex<\/italic>: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull.","volume":"7","author":"CA Brochu","year":"2003","journal-title":"Memoir of the Society of Vertebrate Paleontology"},{"key":"ref17","doi-asserted-by":"crossref","first-page":"615","DOI":"10.1671\/0272-4634(2000)020[0615:FEANTD]2.0.CO;2","article-title":"Forebrain enlargement among nonavian theropod dinosaurs.","volume":"20","author":"HCE Larsson","year":"2000","journal-title":"Journal of Vertebrate Paleontology"},{"key":"ref18","first-page":"111","article-title":"Tyrannosauroidea.","author":"TR Holtz Jr","year":"2004"},{"key":"ref19","doi-asserted-by":"crossref","first-page":"479","DOI":"10.1111\/j.1096-3642.2004.00130.x","article-title":"Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America.","volume":"142","author":"TD Carr","year":"2004","journal-title":"Zoological Journal of the Linnean Society"},{"key":"ref20","doi-asserted-by":"crossref","first-page":"772","DOI":"10.1038\/nature02699","article-title":"Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs.","volume":"430","author":"GM Erickson","year":"2004","journal-title":"Nature"},{"key":"ref21","doi-asserted-by":"crossref","first-page":"1266","DOI":"10.1002\/ar.20983","article-title":"New insights into the brain, braincase, and ear region of tyrannosaurs (Dinosauria, Theropoda), with implications for sensory organization and behavior.","volume":"292","author":"LM Witmer","year":"2009","journal-title":"The Anatomical Record"},{"key":"ref22","first-page":"61","article-title":"The Cleveland tyrannosaur skull (<italic>Nanotyrannus<\/italic> or <italic>Tyrannosaurus<\/italic>): New findings based on CT scanning, with special reference to the braincase.","volume":"57","author":"LM Witmer","year":"2010","journal-title":"Kirtlandia"},{"key":"ref23","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1080\/02724634.1999.10011161","article-title":"Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria).","volume":"19","author":"TD Carr","year":"1999","journal-title":"Journal of Vertebrate Paleontology"},{"key":"ref24","doi-asserted-by":"crossref","first-page":"17261","DOI":"10.1073\/pnas.0906911106","article-title":"A long-snouted multihorned tyrannosaurid from the Late Cretaceous of Mongolia.","volume":"106","author":"SL Brusatte","year":"2009","journal-title":"Proceedings of the National Academy of Sciences, USA"},{"key":"ref25","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1671\/0272-4634(2008)28[123:HOTECO]2.0.CO;2","article-title":"Homology of the \u201cethmoid complex\u201d of tyrannosaurids and its implications for the reconstruction of the olfactory apparatus of non-avian theropods.","volume":"28","author":"F Ali","year":"2008","journal-title":"Journal of Vertebrate Paleontology"},{"key":"ref26","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1671\/0272-4634(2000)020[0001:ADREFT]2.0.CO;2","article-title":"A digitally-rendered endocast for <italic>Tyrannosaurus rex<\/italic>.","volume":"20","author":"CA Brochu","year":"2000","journal-title":"Journal of Vertebrate Paleontology"},{"key":"ref27","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1134\/S0031030107030070","article-title":"A new study of the brain of the predatory dinosaur <italic>Tarbosaurus bataar<\/italic> (Theropoda, Tyrannosauridae).","volume":"41","author":"SV Saveliev","year":"2007","journal-title":"Paleontological Journal"},{"key":"ref28","doi-asserted-by":"crossref","first-page":"667","DOI":"10.1098\/rspb.2008.1075","article-title":"Olfactory acuity in theropods: palaeobiological and evolutionary implications.","volume":"276","author":"DK Zelenitsky","year":"2009","journal-title":"Proceedings of the Royal Society B"},{"key":"ref29","first-page":"225","article-title":"Craniofacial anatomy of <italic>Majungasaurus crenatissimus<\/italic> (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. In: Sampson SD, Krause DW, eds. <italic>Majungasaurus crenatissimus<\/italic> (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar.","volume":"8","author":"SD Sampson","year":"2007","journal-title":"Society of Vertebrate Paleontology Memoir"},{"key":"ref30","first-page":"137","article-title":"The cranial morphology of <italic>Tyrannosaurus rex<\/italic>.","volume":"217","author":"RE Molnar","year":"1991","journal-title":"Paleontographica Abteilung A"},{"key":"ref31","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1080\/02724634.2011.557116","article-title":"Cranial osteology of a juvenile specimen of <italic>Tarbosaurus bataar<\/italic> (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia.","volume":"31","author":"T Tsuihiji","year":"2011","journal-title":"Journal of Vertebrate Paleontology"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"680","DOI":"10.1038\/nature02855","article-title":"Basal tyrannosauroids from China and evidence for protofeathers in tyrannosauroids.","volume":"431","author":"X Xu","year":"2004","journal-title":"Nature"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1038\/nature04511","article-title":"A basal tyrannosauroid dinosaur from the Late Jurassic of China.","volume":"439","author":"X Xu","year":"2006","journal-title":"Nature"},{"key":"ref34","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1206\/650.1","article-title":"The perinate skull of <italic>Byronosaurus<\/italic> (Troodontidae) with observations on the cranial ontogeny of paravian theropods.","volume":"3657","author":"GS Bever","year":"2009","journal-title":"American Museum Novitates"},{"key":"ref35","article-title":"Anatomy, evolution, and functional significance of cephalic vasculature in Archosauria.","author":"JC Sedlmayr","year":"2002"},{"key":"ref36","first-page":"81","article-title":"Braincase anatomy.","author":"PJ Currie","year":"1997","journal-title":"Encyclopedia of Dinosaurs"},{"key":"ref37","first-page":"275","article-title":"The anatomy of the palatoquadrate in the Lower Triassic <italic>Proterosuchus fergusi<\/italic> (Reptilia, Archosauromorpha) and its morphological transformation within the archosauriform clade.","volume":"89","author":"J Klembara","year":"2008","journal-title":"Acta Zoologica"},{"key":"ref38","first-page":"585","article-title":"Organa sensuum [Organa sensoria].","volume":"No. 23","author":"HE Evans","year":"1993"},{"key":"ref39","first-page":"161","article-title":"Giant theropod dinosaurs from Asia and North America: skulls of <italic>Tarbosaurus bataar<\/italic> and <italic>Tyrannosaurus rex<\/italic> compared.","volume":"48","author":"JH Hurum","year":"2003","journal-title":"Acta Palaeontologica Polonica"},{"key":"ref40","first-page":"93","article-title":"A new late Cretaceous carnosaur from Nogon-Tsav Mongolia.","volume":"3","author":"SM Kurzanov","year":"1976","journal-title":"Sovmestna\u00e2 Sovetsko-Mongolska\u00e2 Paleontologiceska\u00e2 Ekspeditci\u00e2, Trudy"},{"key":"ref41","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1159\/000151473","article-title":"Brains, lifestyles, and cognition: are there general trends?","volume":"72","author":"L Lefebvre","year":"2008","journal-title":"Brain, Behavior, and Evolution"},{"key":"ref42","article-title":"The Triple Helix: Gene, Organism and Environment.","author":"R Lewontin","year":"2000"},{"key":"ref43","first-page":"67","article-title":"The relationship between homoplasy and the confidence in a phylogenetic tree.","author":"MJ Sanderson","year":"1996"},{"key":"ref44","first-page":"91","article-title":"Homoplasy increases phylogenetic structure.","volume":"15","author":"V K\u00e4llersj\u00f6","year":"1999","journal-title":"Cladistics"},{"key":"ref45","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1111\/j.1525-142X.2011.00478.x","article-title":"Finding the frameshift: digit loss, developmental variability, and the origin of the avian hand.","volume":"13","author":"GS Bever","year":"2011","journal-title":"Evolution & Development"},{"key":"ref46","doi-asserted-by":"crossref","first-page":"1042","DOI":"10.1002\/dvdy.22595","article-title":"Identity of the avian wing digits: problems resolved and unsolved.","volume":"240","author":"RL Young","year":"2011","journal-title":"Developmental Dynamics"},{"key":"ref47","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1111\/j.1525-142X.2009.00371.x","article-title":"Development of beak polymorphism in the African seedcracker, <italic>Pyrenestes ostrinus<\/italic>.","volume":"11","author":"C Clabaut","year":"2009","journal-title":"Evolution & Development"},{"key":"ref48","doi-asserted-by":"crossref","first-page":"1485","DOI":"10.1126\/science.1161833","article-title":"Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs.","volume":"321","author":"SL Brusatte","year":"2008","journal-title":"Science"},{"key":"ref49","doi-asserted-by":"crossref","first-page":"431","DOI":"10.1146\/annurev.es.20.110189.002243","article-title":"The importance of fossils in phylogeny reconstruction.","volume":"20","author":"MJ Donoghue","year":"1989","journal-title":"Annual Review of Ecology and Systematics"},{"key":"ref50","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1002\/jez.b.21078","article-title":"Developmental constraints vs. variational properties: how pattern formation can help to understand evolution and development.","volume":"306","author":"I Salazar-Ciudad","year":"2006","journal-title":"Journal of Experimental Zoology B"},{"key":"ref51","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1002\/jmor.10677","article-title":"The postnatal skull of the extant North American turtle <italic>Pseudemys texana<\/italic> (Cryptodira: Emydidae), with comments on the study of discrete intraspecific variation.","volume":"270","author":"GS Bever","year":"2009","journal-title":"Journal of Morphology"},{"key":"ref52","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1002\/ar.20865","article-title":"New insight on the anatomy and architecture of the avian neurocranium.","volume":"292","author":"J Margug\u00e1n-Lob\u00f3n","year":"2009","journal-title":"The Anatomical Record"},{"key":"ref53","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/jez.b.41","article-title":"What is the promise of developmental evolution? III. The crucible of developmental evolution.","volume":"300B","author":"GP Wagner","year":"2003","journal-title":"Journal of Experimental Zoology"}],"container-title":["PLoS ONE"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.pone.0023393","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2018,10,19]],"date-time":"2018-10-19T23:16:06Z","timestamp":1539990966000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0023393"}},"subtitle":[],"editor":[{"given":"Daphne","family":"Soares","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2011,8,10]]},"references-count":53,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2011,8,10]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0023393","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2011,8,10]]}}}