{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T07:28:39Z","timestamp":1790062119187,"version":"4.0.1"},"reference-count":135,"publisher":"Public Library of Science (PLoS)","issue":"7","license":[{"start":{"date-parts":[[2024,7,30]],"date-time":"2024-07-30T00:00:00Z","timestamp":1722297600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["DGE-16-4486"],"award-info":[{"award-number":["DGE-16-4486"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100016451","name":"Department of Ornithology, American Museum of Natural History","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100016451","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100013118","name":"Jackson School of Geosciences,University of Texas at Austin","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100013118","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100013118","name":"Jackson School of Geosciences,University of Texas at Austin","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100013118","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"abstract":"<jats:p>Despite making up one of the most ecologically diverse groups of living birds, comprising soaring, diving and giant flightless taxa, the evolutionary relationships and ecological evolution of Anseriformes (waterfowl) remain unresolved. Although Anseriformes have a comparatively rich, global Cretaceous and Paleogene fossil record, morphological datasets for this group that include extinct taxa report conflicting relationships for all known extinct taxa. Correct placement of extinct taxa is necessary to understand whether ancestral anseriform feeding ecology was more terrestrial or one of a set of diverse aquatic ecologies and to better understand avian evolution around the K-T boundary. Here, we present a new morphological dataset for Anseriformes that includes more extant and extinct taxa than any previous anseriform-focused dataset and describe a new anseriform species from the early Eocene Green River Formation of North America. The new taxon has a mediolaterally narrow bill which is rarely found in previously described anseriform fossils. The matrix created to assess the placement of this taxon comprises 41 taxa and 719 discrete morphological characters describing skeletal morphology, musculature, syringeal morphology, ecology, and behavior. We additionally combine the morphological dataset with published sequences using Bayesian methods and perform ancestral state reconstruction for select morphological, ecological and behavioral characters. We recover the new Eocene taxon as the sister taxon to (Anseranatidae+Anatidae) across all analyses, and find that the new taxon represents a novel ecology within known Anseriformes and the Green River taxa. Results provide insight into avian evolution during and following the K-Pg mass extinction and indicate that Anseriformes were likely ancestrally aquatic herbivores with rhamphothecal lamellae..<\/jats:p>","DOI":"10.1371\/journal.pone.0278737","type":"journal-article","created":{"date-parts":[[2024,7,30]],"date-time":"2024-07-30T13:24:01Z","timestamp":1722345841000},"page":"e0278737","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":10,"title":["A new Paleogene fossil and a new dataset for waterfowl (Aves: Anseriformes) clarify phylogeny, ecological evolution, and avian evolution at the K-Pg Boundary"],"prefix":"10.1371","volume":"19","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9476-5757","authenticated-orcid":true,"given":"Grace","family":"Musser","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julia A.","family":"Clarke","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2024,7,30]]},"reference":[{"key":"pone.0278737.ref001","first-page":"63","article-title":"On the geologic age of the vertebrate fauna of the Eocene of New Mexico","volume":"28","author":"P. Cope","year":"1876","journal-title":"Proc Acad Nat Sci Phila"},{"key":"pone.0278737.ref002","unstructured":"H\u00e9bert M. Note sur le f\u00e9mur du Gastornis parisiensis. Mallet-Bachelier; 1855."},{"key":"pone.0278737.ref003","first-page":"616","article-title":"Nouveaux documents sur le gisement du Gastornis parisiensis et considerations generals sur les vestiges laisses par des oiseaux dans les terrains des divers ages","volume":"40","author":"C. Pr\u00e9vost","year":"1855","journal-title":"C R Acad Sci"},{"key":"pone.0278737.ref004","unstructured":"Ml\u00edkovsk\u00fd J. 2002. Cenozoic Birds of the World, Part 1: Europe. Praha: Ninox Press; 2002."},{"key":"pone.0278737.ref005","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-030-87645-6","volume-title":"Paleogene Fossil Birds","author":"G. Mayr","year":"2022","edition":"2"},{"issue":"2","key":"pone.0278737.ref006","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1017\/S0094837300010435","article-title":"Biomechanics of the jaw apparatus of the gigantic Eocene bird Diatryma: implications for diet and mode of life","volume":"17","author":"LM Witmer","year":"1991","journal-title":"Paleobiology"},{"key":"pone.0278737.ref007","unstructured":"Andors AV. Reappraisal of the Eocene groundbird Diatryma (Aves: Anserimorphae). In: Campbell Jr KE, editor. Papers in Avian Paleontology Honoring Pierce Brodkorb. Los Angeles (CA): Science Series of the Natural History Museum of Los Angeles County No. 3; 1992. p. 109\u2013125."},{"issue":"101","key":"pone.0278737.ref008","first-page":"313","article-title":"Isotopic and anatomical evidence of an herbivorous diet in the early Tertiary giant bird Gastornis. Implications for the structure of Paleocene terrestrial ecosystems","volume":"2\u201314","author":"D Angst","journal-title":"Naturwissenschaften"},{"key":"pone.0278737.ref009","doi-asserted-by":"crossref","first-page":"1313","DOI":"10.1080\/02724634.2010.501465","article-title":"Osteology of a new giant bony-toothed bird from the Miocene of Chile, with a revision of the taxonomy of Neogene Pelagornithidae","volume":"30","author":"G Mayr","year":"2010","journal-title":"J Vertebr Paleontol"},{"issue":"29","key":"pone.0278737.ref010","doi-asserted-by":"crossref","first-page":"10624","DOI":"10.1073\/pnas.1320297111","article-title":"Flight performance of the largest volant bird","volume":"111","author":"D. Ksepka","year":"2014","journal-title":"PNAS"},{"key":"pone.0278737.ref011","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1038\/s41586-020-2096-0","article-title":"Late Cretaceous neornithine from Europe illuminates the origin of crown birds","volume":"579","author":"DJ Field","year":"2020","journal-title":"Nature"},{"key":"pone.0278737.ref012","doi-asserted-by":"crossref","first-page":"391","DOI":"10.5962\/p.231090","article-title":"Fossil birds from the Green River Deposits of Eastern Utah","volume":"16","author":"A. Wetmore","year":"1926","journal-title":"Ann Carnegie Mus"},{"key":"pone.0278737.ref013","first-page":"361","article-title":"A phylogenetic analysis of basal Anseriformes, the fossil Presbyornis, and the interordinal relationships of waterfowl","volume":"121","author":"BC Livezey","year":"1997","journal-title":"Zool J Linn Soc"},{"key":"pone.0278737.ref014","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1111\/j.1096-3642.1997.tb01286.x","article-title":"Systematic relationships of the palaeogene family Presbyornithidae (Aves: Anseriformes)","volume":"121","author":"Per. GP Ericson","year":"1997","journal-title":"Zool J Linn Soc"},{"key":"pone.0278737.ref015","volume-title":"The Waterfowl of the World","author":"J Delacour","year":"1945"},{"key":"pone.0278737.ref016","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1093\/auk\/103.4.737","article-title":"A phylogenetic analysis of recent anseriform genera using morphological characters","volume":"103","author":"BC Livezey","year":"1986","journal-title":"Auk"},{"key":"pone.0278737.ref017","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1007\/s11692-015-9345-4","article-title":"The craniolingual morphology of waterfowl (Aves, Anseriformes) and its relationship with feeding mode through contrast-enhanced x-ray computed tomography and 2D morphometrics","volume":"43","author":"Z Li","year":"2015","journal-title":"Evol Biol"},{"issue":"21","key":"pone.0278737.ref018","doi-asserted-by":"crossref","first-page":"5016","DOI":"10.1002\/ece3.1787","article-title":"Exceptional avian herbivores: multiple transitions toward herbivory in the bird order Anseriformes and its correlation with body mass","volume":"5","author":"AM Olsen","year":"2015","journal-title":"Ecol Evol"},{"key":"pone.0278737.ref019","doi-asserted-by":"crossref","first-page":"1985","DOI":"10.1111\/1365-2435.12890","article-title":"Feeding ecology is the primary driver of beak shape diversification in waterfowl","volume":"31","author":"AM Olsen","year":"2017","journal-title":"Funct Ecol"},{"key":"pone.0278737.ref020","first-page":"130","article-title":"Evolutionary trends in the behavior and morphology of the Anatidae","volume":"13","author":"PA Johnsgard","year":"1962","journal-title":"Wildfowl Trust Ann Rep"},{"key":"pone.0278737.ref021","volume-title":"Waterfowl: ducks, geese and swans of the world","author":"FS Todd","year":"1979"},{"key":"pone.0278737.ref022","doi-asserted-by":"crossref","first-page":"1","DOI":"10.5479\/si.00810282.323","article-title":"Presbyornis and the origin of the Anseriformes (Aves: Charadrimorphae)","volume":"323","author":"SL Olson","year":"1980","journal-title":"Smithson Contrib to Zool"},{"key":"pone.0278737.ref023","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1038\/nature03150","article-title":"Definitive fossil evidence for the extant avian radiation in the Cretaceous","volume":"433","author":"JA Clarke","year":"2005","journal-title":"Nature"},{"key":"pone.0278737.ref024","doi-asserted-by":"crossref","first-page":"586","DOI":"10.1007\/s00114-005-0047-0","article-title":"Osteological evidence for sister group relationship between pseudo-toothed birds (Aves: Odontopterygiformes) and waterfowls (Anseriformes)","volume":"92","author":"E. Bourdon","year":"2005","journal-title":"Naturwissenschaften"},{"key":"pone.0278737.ref025","doi-asserted-by":"crossref","first-page":"448","DOI":"10.1111\/j.1463-6409.2011.00484.x","article-title":"Cenozoic mystery birds\u2014on the phylogenetic affinities of bony-toothed birds (Pelagornithidae)","volume":"40","author":"G. Mayr","year":"2011","journal-title":"Zool. Scr"},{"key":"pone.0278737.ref026","doi-asserted-by":"crossref","first-page":"e80372","DOI":"10.1371\/journal.pone.0080372","article-title":"Structure and growth pattern of pseudoteeth in Pelagornis mauretanicus (Aves, Odontopterygiformes, Pelagornithidae)","volume":"8","author":"A Louchart","year":"2013","journal-title":"PLoS One"},{"key":"pone.0278737.ref027","doi-asserted-by":"crossref","first-page":"170975","DOI":"10.1098\/rsos.170975","article-title":"The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres)","volume":"4","author":"TH Worthy","year":"2017","journal-title":"R Soc Open Sci"},{"key":"pone.0278737.ref028","first-page":"1","article-title":"A stem anseriform from the early Palaeocene of Antarctica provides new key evidence in the early evolution of waterfowl","author":"CP Tambussi","year":"2019","journal-title":"Zool J Linn"},{"key":"pone.0278737.ref029","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1071\/ZO9960047","article-title":"Molecular relationships within Australasian waterfowl (Anseriformes)","volume":"44","author":"M Sraml","year":"1996","journal-title":"Aust J Zool"},{"key":"pone.0278737.ref030","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1016\/S1055-7903(02)00019-2","article-title":"A molecular phylogeny of Anseriformes based on mitochondrial DNA analysis","volume":"23","author":"C Donne-Gouss\u00e9","year":"2002","journal-title":"Mol Phylogenet and Evol"},{"issue":"9","key":"pone.0278737.ref031","doi-asserted-by":"crossref","first-page":"e0184529","DOI":"10.1371\/journal.pone.0184529","article-title":"Rapid and recent diversification patterns in Anseriformes birds: inferred from molecular phylogeny and diversification analyses","volume":"12","author":"Z Sun","year":"2017","journal-title":"PLoS One"},{"key":"pone.0278737.ref032","doi-asserted-by":"crossref","DOI":"10.7208\/chicago\/9780226922980.001.0001","volume-title":"The Lost World of Fossil Lake: Snapshots from Deep Time","author":"L. Grande","year":"2013"},{"key":"pone.0278737.ref033","doi-asserted-by":"crossref","first-page":"1","DOI":"10.11646\/zootaxa.1046.1.1","article-title":"New specimens of the early Eocene frigatebird Limnofregata (Pelecaniformes: Fregatidae), with the description of a new species","volume":"1046","author":"SL Olson","year":"2005","journal-title":"Zootaxa"},{"key":"pone.0278737.ref034","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1111\/pala.12134","article-title":"A new species of Limnofregata (Pelecaniformes: Fregatidae) from the Early Eocene Wasatch Formation of Wyoming: implications for palaeoecology and palaeobiology","volume":"58","author":"TA Stidham","year":"2015","journal-title":"Palaeontology"},{"key":"pone.0278737.ref035","first-page":"57","article-title":"An early Eocene Oilbird from the Green River Formation of Wyoming (Caprimulgiformes: Steatornithidae)","volume":"99","author":"SL Olson","year":"1987","journal-title":"Documents des Laboratoires de G\u00e9ologie de Lyon"},{"key":"pone.0278737.ref036","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1080\/02724634.2012.722898","article-title":"A new species of Threskiornithidae-like bird (Aves, Ciconiiformes) from the Green River Formation (Eocene) of Wyoming","volume":"33","author":"NA Smith","year":"2013","journal-title":"J Vertebr Paleontol"},{"issue":"7","key":"pone.0278737.ref037","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1017\/S0016756800181701","article-title":"New fossil bird and fish remains from the Middle Eocene of Wyoming","volume":"4","author":"CR Eastman","year":"1900","journal-title":"Geol Mag"},{"key":"pone.0278737.ref038","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1186\/s12862-018-1212-3","article-title":"A North American stem turaco, and the complex biogeographic history of modern birds","volume":"18","author":"DJ Field","year":"2018","journal-title":"BMC Evol Biol"},{"key":"pone.0278737.ref039","doi-asserted-by":"crossref","first-page":"527","DOI":"10.1130\/G30630.1","article-title":"Eocene clocks agree: coeval 40Ar\/39Ar, U-Pb, and astronomical ages from the Green River Formation","volume":"38","author":"ME Smith","year":"2010","journal-title":"Geology"},{"key":"pone.0278737.ref040","doi-asserted-by":"crossref","first-page":"1","DOI":"10.5479\/si.00810266.63.1","article-title":"The Cretaceous birds of New Jersey","volume":"63","author":"SL Olson","year":"1987","journal-title":"Smithsonian Contributions to Paleobiology"},{"key":"pone.0278737.ref041","first-page":"736","article-title":"Note sur un hum\u00e9rus fossile d\u2019oiseau, attribu\u00e9 \u00e0 un tr\u00e8s-grand palmip\u00e8de de la section des Longipennes","volume":"44","author":"E. Lartet","year":"1857","journal-title":"Comptes rendus hebdomadaires des S\u00e9ances de l\u2019Acad\u00e9mie des Sciences (Paris)"},{"key":"pone.0278737.ref042","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1038\/nature19852","article-title":"Fossil evidence of the avian vocal organ from the Mesozoic","volume":"538","author":"JA Clarke","year":"2016","journal-title":"Nature"},{"key":"pone.0278737.ref043","doi-asserted-by":"crossref","first-page":"104818","DOI":"10.1016\/j.cretres.2021.104818","article-title":"New data on the Vegavis iaai holotype from the Maastrichtian of Antarctica","volume":"124","author":"CA Hospitaleche","year":"2021","journal-title":"Cret Res"},{"key":"pone.0278737.ref044","doi-asserted-by":"crossref","first-page":"233","DOI":"10.3390\/d15020233","article-title":"Basal Anseriformes from the Early Paleogene of North America and Europe","volume":"15","author":"P Houde","year":"2023","journal-title":"Diversity"},{"key":"pone.0278737.ref045","doi-asserted-by":"crossref","first-page":"21","DOI":"10.3853\/j.0067-1975.62.2010.1537","article-title":"A new avian species with tubercle-bearing cervical vertebrae from the Middle Eocene of Messel (Germany)","volume":"62","author":"G. Mayr","year":"2010","journal-title":"Records of the Australian Museum"},{"key":"pone.0278737.ref046","doi-asserted-by":"crossref","first-page":"1","DOI":"10.2113\/gsrocky.30.1.33","article-title":"Paleontological and sedimentological variation in early Eocene Fossil Lake","volume":"30","author":"L Grande","year":"1994","journal-title":"Rocky Mountain Geol"},{"issue":"406","key":"pone.0278737.ref047","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1206\/0003-0090-406.1.1","article-title":"The anatomy and taxonomy of the exquisitely preserved Green River Formation (Early Eocene) Lithornithids (Aves) and the relationships of Lithornithidae","volume":"2016","author":"SJ Nesbitt","year":"2016","journal-title":"Bull Am Mus Nat Hist"},{"key":"pone.0278737.ref048","first-page":"1","article-title":"A new wading bird from the Eocene of Patagonia","volume":"1710","author":"H. Howard","year":"1955","journal-title":"Am Mus Novit"},{"key":"pone.0278737.ref049","unstructured":"Alvarenga, HMF. A fossil Screamer (Anseriformes: Anhimidae) from the middle Tertiary of southeastern Brazil. In: Olson SL, editor. Avian Paleontology at the Close of the 20th Century: Proceedings of the 5th International Meeting of the Society of Avian Paleontology and Evolution, Washington, D.C., 4\u20137 June 1996. Washington, D.C.: Smithsonian Contributions to Paleobiology 89; 1999. Pp. 223\u2013230."},{"issue":"11","key":"pone.0278737.ref050","first-page":"307","article-title":"The skeleton of Diatryma, a gigantic bird from the lower Eocene of Wyoming","volume":"37","author":"WD Matthew","year":"1917","journal-title":"Bull Am Mus Nat Hist"},{"issue":"1","key":"pone.0278737.ref051","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1002\/spp2.1284","article-title":"Oldest, smallest and phylogenetically most basal Pelagornithid, from the early Paleocene of New Zealand, sheds light on the evolutionary history of the largest flying birds","volume":"7","author":"G Mayr","year":"2021","journal-title":"Pap Palaeontol"},{"key":"pone.0278737.ref052","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1002\/jmor.10799","article-title":"Morphology of the quadrate in the Eocene anseriform Presbyornis and extant galloanserine birds","volume":"271","author":"A Elzanowski","year":"2010","journal-title":"Journal of Morphology"},{"issue":"1725","key":"pone.0278737.ref053","article-title":"Evolution of olfaction in non-avian theropod dinosaurs and birds","volume":"278","author":"DK Zelenitsky","year":"2011","journal-title":"Proc R Soc Lond B Biol Sci"},{"key":"pone.0278737.ref054","doi-asserted-by":"crossref","unstructured":"Olson, SL. The anseriform relationships of Anatalavis Olson and Parris (Anseranatidae), with a species from the lower Eocene London Clay. In: Olson SL, editor. Avian Paleontology at the Close of the 20th Century: Proceedings of the 5th International Meeting of the Society of Avian Paleontology and Evolution, Washington, D.C., 4\u20137 June 1996. Washington, D.C.: Smithsonian Contributions to Paleobiology 89. 1999. pp. 231\u2013243.","DOI":"10.5479\/si.00810266.89.1"},{"issue":"2","key":"pone.0278737.ref055","first-page":"211","article-title":"The early Eocene bird Gallinuloides wyomingensis\u2014a stem group representative of Galliformes","volume":"49","author":"G Mayr","year":"2004","journal-title":"Acta Palaeontologica Polonica"},{"key":"pone.0278737.ref056","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1206\/0003-0090(2004)286<0001:MPTASO>2.0.CO;2","article-title":"Morphology, phylogenetic taxonomy, and systematics of Ichthyornis and Apatornis (Avialae: Ornithurae)","volume":"286","author":"JA Clarke","year":"2004","journal-title":"Bull Am Mus Nat Hist"},{"key":"pone.0278737.ref057","volume-title":"Paleognathous Birds from the Early Tertiary of the Northern Hemisphere","author":"PW Houde","year":"1988"},{"issue":"2","key":"pone.0278737.ref058","doi-asserted-by":"crossref","DOI":"10.1098\/rsos.150635","article-title":"The unexpected survival of an ancient lineage of anseriform birds into the Neogene of Australia: the youngest record of Presbyornithidae","volume":"3","author":"VL De Pietri","year":"2016","journal-title":"R Soc Open Sci"},{"key":"pone.0278737.ref059","first-page":"43","volume-title":"Paleornithological Research 2013: Proceedings of the 8th International Meeting of the Society of Avian Paleontology and Evolution","author":"WE Boles","year":"2013"},{"key":"pone.0278737.ref060","doi-asserted-by":"crossref","unstructured":"Mayr G, Carrio V, Kitchener AC. On the \u201cscreamer-like\u201d birds from the British London Clay: An archaic anseriform-galliform mosaic and a non-galloanserine \u201cbarb-necked\u201d species of Perplexicervix. Paleontologica Electronica. 2023; 26.2.a33.","DOI":"10.26879\/1301"},{"key":"pone.0278737.ref061","first-page":"7","volume-title":"New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrum","author":"J Gauthier","year":"2001"},{"key":"pone.0278737.ref062","first-page":"510","article-title":"Einige Mittheilungen \u00fcber Thiere Mexicos","volume":"5","author":"JG Wagler","year":"1831","journal-title":"Isis von Oken"},{"key":"pone.0278737.ref063","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1111\/pala.12270","article-title":"The plumage and colouration of an Enantiornithine bird from the Early Cretaceous of China","volume":"60","author":"JA Peteya","year":"2017","journal-title":"Palaeontology"},{"key":"pone.0278737.ref064","first-page":"1","article-title":"The extent of the preserved feathers on the four-winged dinosaur Microraptor gui under ultraviolet light","volume":"5","author":"DWE Hone","year":"2010","journal-title":"PLoS One"},{"key":"pone.0278737.ref065","doi-asserted-by":"crossref","first-page":"1705","DOI":"10.1139\/e01-050","article-title":"Anatomy of Sinosauropteryx prima from Liaoning, northeastern China","volume":"38","author":"PJ Currie","year":"2001","journal-title":"Can. J. Earth Sci"},{"key":"pone.0278737.ref066","doi-asserted-by":"crossref","first-page":"640","DOI":"10.1038\/nature08322","article-title":"A pre-Archaeopteryx troodontid from China with long feathers on the metatarsus","volume":"461","author":"D Hu","year":"2009","journal-title":"Nature"},{"key":"pone.0278737.ref067","unstructured":"O\u2019Leary MA, Kaufman SG. MorphoBank 3.0: Web Application for Morphological Phylogenetics and Taxonomy [Internet]. 2012 [cited Mar 12 2019]. morphobank.org."},{"key":"pone.0278737.ref068","unstructured":"Shoshoni Dictionary [Internet]. Shoshoni Language Project. 2018 [cited Aug 6 2020]. https:\/\/shoshoniproject.utah.edu\/language-materials\/shoshoni-dictionary\/ dictionary.php."},{"key":"pone.0278737.ref069","first-page":"50","volume-title":"Sedimentology and Geochemistry of Modern and Ancient Saline Lakes","author":"P. Buchheim","year":"1994"},{"key":"pone.0278737.ref070","first-page":"1","article-title":"New birds from the lower Eocene Green River Formation, North America","volume":"62","author":"I. Weidig","year":"2010","journal-title":"Proceedings of the VII International Meeting of the Society of Avian Paleontology and Evolution. Rec. Aust. Mus"},{"key":"pone.0278737.ref071","article-title":"Podargiform Affinities of the Enigmatic Fluvioviridavis platyrhamphus and the Early Diversification of Strisores (\u201cCaprimulgiformes\u201d + Apodiformes)","author":"SJ Nesbitt","year":"2011","journal-title":"PLoS One"},{"key":"pone.0278737.ref072","first-page":"1","article-title":"A Lower Eocene Frigatebird from the Green River Formation of Wyoming (Pelecaniformes: Fregatidae)","volume":"35","author":"SL Olson","year":"1977","journal-title":"Smithsonian Contributions to Paleontology"},{"key":"pone.0278737.ref073","first-page":"171","volume-title":"Contributions in Science: Papers in Avian Paleontotogy Honoring Pierce Brodkorb.","author":"A. Hesse","year":"1992"},{"key":"pone.0278737.ref074","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1080\/02724630903412414","article-title":"Primobucco mcgrewi (Aves: Coracii) from the Eocene Green River Formation: new anatomical data from the earliest constrained record of stem rollers","volume":"30","author":"DT Ksepka","year":"2010","journal-title":"J Vertebr Paleontol"},{"key":"pone.0278737.ref075","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1666\/10-108.1","article-title":"Stem Parrots (Aves, Halcyornithidae) from the Green River Formation and a Combined Phylogeny of Pan-Psittaciformes","volume":"85","author":"DT Ksepka","year":"2011","journal-title":"J Paleontol"},{"key":"pone.0278737.ref076","doi-asserted-by":"crossref","first-page":"101","DOI":"10.5479\/si.00810266.27.101","volume-title":"Collected Papers in Avian Paleontology Honoring the 90th Birthday of Alexander Wetmore","author":"A Feduccia","year":"1976"},{"key":"pone.0278737.ref077","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1016\/j.cub.2018.12.040","article-title":"Oldest finch-beaked birds reveal parallel ecological radiations in the earliest evolution of Passerines","volume":"29","author":"DT Ksepka","year":"2019","journal-title":"Current Biology"},{"key":"pone.0278737.ref078","doi-asserted-by":"crossref","first-page":"700","DOI":"10.1080\/02724634.1998.10011099","article-title":"A well-preserved fossil amphiumid (Lissamphibia: Caudata) from the Eocene Green River Formation of Wyoming","volume":"18","author":"O Rieppel","year":"1998","journal-title":"J Vertebr Paleontol"},{"key":"pone.0278737.ref079","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1206\/3927.1","article-title":"A new morphological dataset reveals a novel relationship for the adzebills of New Zealand (Aptornis) and provides a foundation for total evidence neoavian phylogenetics","volume":"2019","author":"GM Musser","year":"2019","journal-title":"Am Mus Novit"},{"key":"pone.0278737.ref080","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1111\/azo.12117","article-title":"Variations in the hypotarsus morphology of birds and their evolutionary significance","volume":"97","author":"G. Mayr","year":"2016","journal-title":"Acta Zool"},{"key":"pone.0278737.ref081","doi-asserted-by":"crossref","first-page":"e82000","DOI":"10.1371\/journal.pone.0082000","article-title":"Skeletal correlates for body mass estimation in modern and fossil flying birds","volume":"8","author":"DJ Field","year":"2013","journal-title":"PLoS One"},{"key":"pone.0278737.ref082","first-page":"45","volume-title":"Handbook of Avian Anatomy: Nomina Anatomica Avium","author":"JJ Baumel","year":"1993"},{"key":"pone.0278737.ref083","first-page":"344","article-title":"Notice of a new and remarkable fossil bird","volume":"4","author":"OC Marsh","year":"1872","journal-title":"American Journal of Science"},{"key":"pone.0278737.ref084","doi-asserted-by":"crossref","first-page":"559929","DOI":"10.3389\/fevo.2020.559929","article-title":"An exceptionally preserved specimen from the Green River Formation elucidates complex phenotypic evolution in Gruiformes and Charadriiformes","volume":"8","author":"G Musser","year":"2020","journal-title":"Front Ecol Evol"},{"key":"pone.0278737.ref085","unstructured":"The Mesquite Project Team. 2021. Documentation for Mesquite: a modular system for evolutionary analysis. http:\/\/www.mesquiteproject.org."},{"key":"pone.0278737.ref086","first-page":"143","volume-title":"New Perspectives on the Origin and Early Evolution of Birds","author":"JL Cracraft","year":"2001"},{"key":"pone.0278737.ref087","doi-asserted-by":"crossref","first-page":"508","DOI":"10.1038\/35107146","article-title":"Fossils and avian evolution","volume":"414","author":"JA Clarke","year":"2001","journal-title":"Nature"},{"key":"pone.0278737.ref088","doi-asserted-by":"crossref","first-page":"527","DOI":"10.1111\/j.1096-0031.2003.tb00387.x","article-title":"The deep divergences of neornithine birds: a phylogenetic analysis of morphological characters","volume":"19","author":"G Mayr","year":"2003","journal-title":"Cladistics"},{"key":"pone.0278737.ref089","first-page":"1","article-title":"Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy (I. Methods and characters)","volume":"37","author":"BC Livezey","year":"2006","journal-title":"Bulletin of Carnegie Museum of Natural History"},{"key":"pone.0278737.ref090","doi-asserted-by":"crossref","first-page":"D67","DOI":"10.1093\/nar\/gkv1276","article-title":"Genbank","volume":"44","author":"K Clark","year":"2016","journal-title":"Nucleic Acids Res"},{"issue":"5","key":"pone.0278737.ref091","doi-asserted-by":"crossref","DOI":"10.1093\/sysbio\/syx041","article-title":"Why do phylogenomic data sets yield conflicting trees? Data type influences the avian tree of life more than taxon sampling","volume":"66","author":"S Reddy","year":"2017","journal-title":"Syst. Biol"},{"issue":"14","key":"pone.0278737.ref092","doi-asserted-by":"crossref","first-page":"3059","DOI":"10.1093\/nar\/gkf436","article-title":"MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform","volume":"30","author":"K Katoh","year":"2002","journal-title":"Nucleic Acids Res"},{"key":"pone.0278737.ref093","doi-asserted-by":"crossref","first-page":"1253","DOI":"10.1093\/molbev\/msn083","article-title":"jModelTest: Phylogenetic Model Averaging","volume":"25","author":"D. Posada","year":"2008","journal-title":"Mol Biol Evol"},{"key":"pone.0278737.ref094","doi-asserted-by":"crossref","first-page":"1411","DOI":"10.1093\/gbe\/evw086","article-title":"Capturing the Phylogeny of Holometabola with Mitochondrial Genome Data and Bayesian Site-Heterogeneous Mixture Models","volume":"8","author":"F Song","year":"2016","journal-title":"Genome Biology and Evolution"},{"key":"pone.0278737.ref095","volume-title":"Version 4.0a164 (X86)","author":"DL Swofford","year":"2002"},{"key":"pone.0278737.ref096","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1007\/BF02101990","article-title":"A new method for calculating evolutionary substitution rates","volume":"20","author":"C Lanave","year":"1984","journal-title":"J Mol Evol"},{"key":"pone.0278737.ref097","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1007\/BF00160154","article-title":"Maximum likelihood phylogenetic estimation from DNA sequences with variable rates over sites: approximate methods","volume":"39","author":"Z. Yang","year":"1994","journal-title":"J. Mol. Evol"},{"key":"pone.0278737.ref098","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1080\/106351501753462876","article-title":"A likelihood approach to estimating phylogeny from discrete morphological character data","volume":"50","author":"PO Lewis","year":"2001","journal-title":"Syst Biol"},{"key":"pone.0278737.ref099","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1093\/oxfordjournals.molbev.a025811","article-title":"Bayesian phylogenetic inference using DNA sequences: a Markov Chain Monte Carlo Method","volume":"14","author":"Z Yang","year":"1997","journal-title":"Mol Biol Evol"},{"key":"pone.0278737.ref100","doi-asserted-by":"crossref","first-page":"754","DOI":"10.1093\/bioinformatics\/17.8.754","article-title":"MrBayes: bayesian inference of phylogenetic trees","volume":"17","author":"JP Huelsenbeck","year":"2001","journal-title":"Bioinformatics"},{"key":"pone.0278737.ref101","doi-asserted-by":"crossref","first-page":"1572","DOI":"10.1093\/bioinformatics\/btg180","article-title":"MrBayes 3: Bayesian phylogenetic inference under mixed models","volume":"19","author":"F Ronquist","year":"2003","journal-title":"Bioinformatics"},{"key":"pone.0278737.ref102","first-page":"1","volume-title":"Proceedings of the Gateway Computing Environments Workshop (GCE)","author":"MA Miller","year":"2010"},{"key":"pone.0278737.ref103","doi-asserted-by":"crossref","first-page":"79","DOI":"10.3161\/068.041.0113","article-title":"Two additional synapomorphies of grebes Podicipedidae and flamingos Phoenicopteridae","volume":"41","author":"A. Manegold","year":"2006","journal-title":"Acta Ornithol"},{"key":"pone.0278737.ref104","doi-asserted-by":"crossref","DOI":"10.1126\/sciadv.abg7099","article-title":"Bird neurocranial and body mass evolution across the end-Cretaceous mass extinction: The avian brain shape left other dinosaurs behind","volume":"7","author":"CR Torres","year":"2021","journal-title":"Science Advances"},{"key":"pone.0278737.ref105","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1038\/s41586-022-05445-y","article-title":"Cretaceous ornithurine supports a neognathous crown bird ancestor","volume":"612","author":"J Benito","year":"2022","journal-title":"Nature"},{"key":"pone.0278737.ref106","first-page":"247","article-title":"Preliminary diagnoses of two extraordinary new genera of birds from Pleistocene deposits in the Hawaiian Islands","volume":"89","author":"SL Olson","year":"1976","journal-title":"Proceedings of the Biological Society of Washington"},{"key":"pone.0278737.ref107","first-page":"1","article-title":"Descriptions of thirty-two new species of birds from the Hawaiian islands","volume":"45","author":"SL Olson","year":"1991","journal-title":"I. Non-passeriformes. Ornithol Monagr"},{"key":"pone.0278737.ref108","doi-asserted-by":"crossref","first-page":"873","DOI":"10.1038\/35091139","article-title":"The beaks of ostrich dinosaurs","volume":"412","author":"MA Norell","year":"2001","journal-title":"Nature"},{"key":"pone.0278737.ref109","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1111\/j.1475-4983.2005.00448.x","article-title":"The diet of ostrich dinosaurs (the Ornithomimosauria)","volume":"48","author":"PM Barrett","year":"2005","journal-title":"Palaeontology"},{"key":"pone.0278737.ref110","doi-asserted-by":"crossref","first-page":"22281","DOI":"10.1038\/s41598-021-01312-4","article-title":"The first edentulous ceratosaur from South America","volume":"11","author":"G Alves de Souza","year":"2021","journal-title":"Nature Scientific Reports"},{"key":"pone.0278737.ref111","doi-asserted-by":"crossref","first-page":"102","DOI":"10.2307\/1443010","article-title":"Occurrence and significance of the os transiliens in gopher tortises","volume":"1","author":"DM Bramble","year":"1974","journal-title":"Copeia"},{"key":"pone.0278737.ref112","volume-title":"Encyclopedia of turtles","author":"PCH Pritchard","year":"1979"},{"key":"pone.0278737.ref113","first-page":"1","article-title":"Evolution of jaw mechanisms in ornithopod dinosaurs","volume":"87","author":"DB Weishampel","year":"1984","journal-title":"Adv Anat Embryol Cel"},{"key":"pone.0278737.ref114","first-page":"293","volume-title":"Encyclopedia of dinosaurs","author":"CA Forster","year":"1997"},{"key":"pone.0278737.ref115","first-page":"269","article-title":"Comparative mechanics of filter feeding","volume":"108","author":"JGM Kooloos","year":"1989","journal-title":"Anas platyrhynchos, Anas clypeata and Aythya fuligula (Aves, Anseriformes). Zoomorphology"},{"key":"pone.0278737.ref116","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1163\/157075603322539453","article-title":"Conflicting demands on the trophic system of Anseriformes and their evolutionary implications","volume":"53","author":"AHJ Van der Leeuw","year":"2003","journal-title":"Anim Biol"},{"key":"pone.0278737.ref117","first-page":"105","volume-title":"Functional anatomy of the syrinx in Form and Function in Birds","author":"AS King","year":"1989"},{"key":"pone.0278737.ref118","first-page":"9","article-title":"The beginning of the modern avian radiation","volume":"99","author":"LD Martin","year":"1987","journal-title":"Documents des Laboratories de Giologieban"},{"key":"pone.0278737.ref119","doi-asserted-by":"crossref","first-page":"14106","DOI":"10.1038\/s41598-020-70751-2","article-title":"Palaeontological evidence reveals convergent evolution of intervertebral joint types in amniotes","volume":"10","author":"T Wintrich","year":"2020","journal-title":"Nature Scientific Reports"},{"issue":"4","key":"pone.0278737.ref120","doi-asserted-by":"crossref","first-page":"525","DOI":"10.1080\/00206816709474481","article-title":"Functional evolution of the fish skeleton","volume":"9","author":"VN Yakovlev","year":"1967","journal-title":"Int Geol Rev"},{"key":"pone.0278737.ref121","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1111\/j.1096-3642.1976.tb00830.x","article-title":"The development, function, and design of amphicoelous vertebrae in teleost fishes","volume":"58","author":"J. Laerm","year":"1976","journal-title":"Zool J Linn Soc"},{"key":"pone.0278737.ref122","doi-asserted-by":"crossref","first-page":"150439","DOI":"10.1098\/rsos.150439","article-title":"Morphological and functional changes in the vertebral column with increasing aquatic adaptation in crocodylomorphs","volume":"2","author":"JL Molnar","year":"2015","journal-title":"R Soc Open Sci"},{"key":"pone.0278737.ref123","volume-title":"Osteology of the Reptiles","author":"AS Romer","year":"1956"},{"key":"pone.0278737.ref124","first-page":"239","article-title":"Redefinici\u00f3n estratigr\u00e1fica del grupo R\u00edo Chico (Pale\u00f3geno inferior), en el norte de la Cuenca del Golfo San Jorge, Chubut","volume":"67","author":"MS Raigemborn","year":"2010","journal-title":"Revista de la Asociaci\u00f3n Geol\u00f3gica Argentina"},{"key":"pone.0278737.ref125","first-page":"429","article-title":"A Giant Presbyornis (Aves, Anseriformes) and Other Birds from the Paleocene Aquia Formation of Maryland and Virginia","volume":"107","author":"SL Olson","year":"1994","journal-title":"Proceedings of the Biological Society of Washington"},{"key":"pone.0278737.ref126","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1206\/0003-0082(2004)447<0001:NARAAR>2.0.CO;2","article-title":"New Avialan Remains and a Review of the Known Avifauna from the Late Cretaceous Nemegt Formation of Mongolia","volume":"2004","author":"JA Clarke","year":"2004","journal-title":"Am Mus Novit"},{"key":"pone.0278737.ref127","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1002\/gj.1177","article-title":"A large collection of Presbyornis (Aves, Anseriformes, Presbyornithidae) from the late Paleocene and early Eocene of Mongolia","volume":"45","author":"EN Kurochkin","year":"2010","journal-title":"Geol J"},{"key":"pone.0278737.ref128","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1206\/3934.1","article-title":"New Fossil Birds from the Earliest Eocene of Mongolia","volume":"2019","author":"SC Hood","year":"2019","journal-title":"Am Mus Novit"},{"key":"pone.0278737.ref129","doi-asserted-by":"crossref","first-page":"675","DOI":"10.1126\/science.246.4930.675","article-title":"Angiosperm Diversification and Paleolatitudinal Gradients in Cretaceous Floristic Diversity","volume":"246","author":"PR Crane","year":"1989","journal-title":"Science"},{"key":"pone.0278737.ref130","first-page":"1","article-title":"Global climate change driven by soot at the K-Pg boundary as the cause of the mass extinction","volume":"6","author":"K Kaiho","year":"2016","journal-title":"Sci Rep-UK"},{"key":"pone.0278737.ref131","doi-asserted-by":"crossref","first-page":"1825","DOI":"10.1016\/j.cub.2018.04.062","article-title":"Early evolution of modern birds structured by global forest collapse at the end-Cretaceous mass extinction","volume":"28","author":"DJ Field","year":"2018","journal-title":"Curr Biol"},{"key":"pone.0278737.ref132","doi-asserted-by":"crossref","first-page":"28867","DOI":"10.1073\/pnas.2005571117","article-title":"The rise of angiosperms pushed conifers to decline during global cooling","volume":"117","author":"FL Condamine","year":"2020","journal-title":"Proc Natl Acad Sci"},{"key":"pone.0278737.ref133","doi-asserted-by":"crossref","first-page":"25327","DOI":"10.1073\/pnas.2004596117","article-title":"Organic matter from the Chicxulub crater exacerbated the K\u2014Pg impact winter","volume":"117","author":"SL Lyons","year":"2020","journal-title":"PNAS"},{"key":"pone.0278737.ref134","doi-asserted-by":"crossref","first-page":"103400","DOI":"10.1016\/j.gloplacha.2020.103400","article-title":"Response of the lacustrine flora in East Asia to global climate changes across the K\/Pg boundary","volume":"197","author":"S Li","year":"2021","journal-title":"Global Planet 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