Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

Cretaceous ornithurine supports a neognathous crown bird ancestor


The bony palate diagnoses the two deepest clades of extant birds: Neognathae and Palaeognathae1,2,3,4,5. Neognaths exhibit unfused palate bones and generally kinetic skulls, whereas palaeognaths possess comparatively rigid skulls with the pterygoid and palatine fused into a single element, a condition long considered ancestral for crown birds (Neornithes)3,5,6,7,8. However, fossil evidence of palatal remains from taxa close to the origin of Neornithes is scarce, hindering strong inferences regarding the ancestral condition of the neornithine palate. Here we report a new taxon of toothed Late Cretaceous ornithurine bearing a pterygoid that is remarkably similar to those of the extant neognath clade Galloanserae (waterfowl + landfowl). Janavis finalidens, gen. et sp. nov., is generally similar to the well-known Mesozoic ornithurine Ichthyornis in its overall morphology, although Janavis is much larger and exhibits a substantially greater degree of postcranial pneumaticity. We recovered Janavis as the first-known well-represented member of Ichthyornithes other than Ichthyornis, clearly substantiating the persistence of the clade into the latest Cretaceous9. Janavis confirms the presence of an anatomically neognathous palate in at least some Mesozoic non-crown ornithurines10,11,12, suggesting that pterygoids similar to those of extant Galloanserae may be plesiomorphic for crown birds. Our results, combined with recent evidence on the ichthyornithine palatine12, overturn longstanding assumptions about the ancestral crown bird palate, and should prompt reevaluation of the purported galloanseran affinities of several bizarre early Cenozoic groups such as the ‘pseudotoothed birds’ (Pelagornithidae)13,14,15.

This is a preview of subscription content, access via your institution

Access options

Rent or buy this article

Get just this article for as long as you need it


Prices may be subject to local taxes which are calculated during checkout

Fig. 1: Skeletal reconstruction, axial pneumaticity and phylogenetic position of J. finalidens.
Fig. 2: Comparative pterygoid and palatal morphology of J. finalidens and representatives of the main neornithine lineages.
Fig. 3: Three-dimensional geometric morphospace of avian pterygoid morphology.

Data availability

Scan data and surface meshes of all preserved elements of Janavis are housed on MorphoSource (; project ID 000444955), along with surface meshes of comparative taxa (; project ID 000444956). See Supplementary Information for a complete list of skeletal models and relevant links. Phylogenetic matrices and morphometric landmark coordinates are provided at Zenodo (; The Life Science Identifier for Janavis is

Code availability

The code underpinning our morphometric analyses is provided at Zenodo (;


  1. Huxley, T. H. On the classification of birds; and on the taxonomic value of the modifications of certain of the cranial bones observed in that class. J. Anat. Physiol. 2, 390 (1868).

    Google Scholar 

  2. Pycraft, W. P. On the morphology and phylogeny of the Palæognathæ (Ratitæand Crypturi) and Neognathæ (Carinatæ). Trans. Zool. Soc. Lond. 15, 149–290 (1900).

    Article  Google Scholar 

  3. Pycraft, W. P. Some points in the morphology of the palate of the Neognathæ. Zool. J. Linn. Soc. 28, 343–357 (1901).

    Article  Google Scholar 

  4. McDowell, S. The bony palate of birds. Part I. The Palaeognathae. Auk 65, 520–549 (1948).

    Article  Google Scholar 

  5. Simonetta, A. M. On the mechanical implications of the avian skull and their bearing on the evolution and classification of birds. Q. Rev. Biol. 35, 206–220 (1960).

    Article  Google Scholar 

  6. Houde, P. W. Paleognathous birds from the early Tertiary of the Northern Hemisphere. Pub. Nuttall Ornithol. Club 22, 1–148 (1988).

    Google Scholar 

  7. Hu, H. et al. Evolution of the vomer and its implications for cranial kinesis in Paraves. Proc. Natl Acad. Sci. USA 116, 19571–19578 (2019).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  8. Witmer, L. M. & Martin, L. D. The primitive features of the avian palate, with special reference to Mesozoic birds. Travaux et Documents des Laboratoires de Géologie de Lyon 99, 21–40 (1987).

    Google Scholar 

  9. Longrich, N. R., Tokaryk, T. & Field, D. J. Mass extinction of birds at the Cretaceous–Paleogene (K–Pg) boundary. Proc. Natl Acad. Sci. USA 108, 15253–15257 (2011).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  10. Zusi, R. L. & Livezey, B. C. Variation in the os palatinum and its structural relation to the palatum osseum of birds (Aves). Ann. Carnegie Mus. 75, 137–180 (2006).

    Article  Google Scholar 

  11. Bell, A. & Chiappe, L. M. Anatomy of Parahesperornis: evolutionary mosaicism in the Cretaceous Hesperornithiformes (Aves). Life 10, 62 (2020).

    Article  PubMed  PubMed Central  Google Scholar 

  12. Torres, C. R., Norell, M. A. & Clarke, J. A. Bird neurocranial and body mass evolution across the end-Cretaceous mass extinction: the avian brain shape left other dinosaurs behind. Sci. Adv. 7, eabg7099 (2021).

    Article  PubMed  PubMed Central  Google Scholar 

  13. Bourdon, E. Osteological evidence for sister group relationship between pseudo-toothed birds (Aves: Odontopterygiformes) and waterfowls (Anseriformes). Naturwissenschaften 92, 586–591 (2005).

    Article  CAS  PubMed  Google Scholar 

  14. Mayr, G. Cenozoic mystery birds—on the phylogenetic affinities of bony‐toothed birds (Pelagornithidae). Zool. Scr. 40, 448–467 (2011).

    Article  Google Scholar 

  15. Mayr, G., De Pietri, V. L., Love, L., Mannering, A. & Scofield, R. P. 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. Pap. Palaeontol. 7, 217–233 (2019).

    Article  Google Scholar 

  16. Elżanowski, A. On the role of basipterygoid processes in some birds. Verh. Anat. Ges. 71, 1303–1307 (1977).

    Google Scholar 

  17. Mayr, G. Paleogene Fossil Birds 2nd edn (Springer, 2022).

  18. Gingerich, P. D. Skull of Hesperornis and early evolution of birds. Nature 243, 70–73 (1973).

    Article  Google Scholar 

  19. Dyke, G. J. et al. Europe’s last Mesozoic bird. Naturwissenschaften 89, 408–411 (2002).

    Article  CAS  PubMed  Google Scholar 

  20. Vellekoop, J. et al. A new age model and chemostratigraphic framework for the Maastrichtian type area (southeastern Netherlands, northeastern Belgium). Newsl. Stratigr. 2022, 0703 (2022).

    Google Scholar 

  21. Clarke, J. A. Morphology, phylogenetic taxonomy, and systematics of Ichthyornis and Apatornis (Avialae: Ornithurae). Bull. Am. Mus. Nat. Hist. 2004, 1–179 (2004).

    Article  Google Scholar 

  22. Benito, J. et al. 40 New specimens of Ichthyornis provide unprecedented insight into the postcranial morphology of crownward stem group birds. PeerJ 10, e13919 (2022).

    Google Scholar 

  23. Marsh, O. C. Odontornithes: a Monograph on the Extinct Toothed Birds of North America: with Thirty-four Plates and Forty Woodcuts. Memoirs of the Peabody Museum of Natural History 1 (US Government Printing Office, 1880).

  24. Field, D. J. et al. Complete Ichthyornis skull illuminates mosaic assembly of the avian head. Nature 557, 96–100 (2018).

    Article  CAS  PubMed  Google Scholar 

  25. Witmer, L. M. The craniofacial air sac system of Mesozoic birds (Aves). Zool. J. Linn. Soc. 100, 327–378 (1990).

    Article  Google Scholar 

  26. Tahara, R. & Larsson, H. C. Head pneumatic sinuses in Japanese quail and zebra finch. Zool. J. Linn. Soc. 186, 742–792 (2019).

    Google Scholar 

  27. Field, D. J., Benito, J., Chen, A., Jagt, J. W. M. & Ksepka, D. T. Late Cretaceous neornithine from Europe illuminates the origins of crown birds. Nature 579, 397–401 (2020).

    Article  CAS  PubMed  Google Scholar 

  28. Berv, J. S. & Field, D. J. Genomic signature of an avian Lilliput effect across the K–Pg extinction. Syst. Biol. 67, 1–13 (2018).

    Article  PubMed  Google Scholar 

  29. Field, D. J. et al. in Pennaraptoran Theropod Dinosaurs: Past Progress and New Frontiers Vol. 440 (eds Pittman, M. & Xu, X.) 160–181 (Bulletin of the American Museum of Natural History, 2020).

  30. Field, D. J. et al. Early evolution of modern birds structured by global forest collapse at the end-Cretaceous mass extinction. Curr. Biol. 28, 1825–1831 (2018).

    Article  CAS  PubMed  Google Scholar 

  31. Prum, R. O. et al. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526, 569–573 (2015).

    Article  CAS  PubMed  Google Scholar 

  32. Feduccia, A. Explosive evolution in Tertiary birds and mammals. Science 267, 637–638 (1995).

    Article  CAS  PubMed  Google Scholar 

  33. Bock, W. J. Kinetics of the avian skull. J. Morphol. 114, 1–42 (1964).

    Article  Google Scholar 

  34. Bühler, P., Martin, L. D. & Witmer, L. M. Cranial kinesis in the Late Cretaceous birds Hesperornis and Parahesperornis. Auk 105, 111–122 (1988).

    Article  Google Scholar 

  35. Bout, R. G. & Zweers, G. A. The role of cranial kinesis in birds. Comp. Biochem. Physiol. A Mol. Integr. Physiol. 131, 197–205 (2001).

    Article  CAS  PubMed  Google Scholar 

  36. Gussekloo, S. W. & Bout, R. G. Cranial kinesis in palaeognathous birds. J. Exp. Biol. 208, 3409–3419 (2005).

    Article  PubMed  Google Scholar 

  37. Lucas, F. A. Notes on the osteology and relationship of the fossil birds of the genera Hesperornis Hargeria Baptornis and Diatryma. Proc. US Natl Mus. 26, 545–556 (1903).

    Article  Google Scholar 

  38. Elzanowski, A. New observations of the skull of Hesperornis with reconstructions of the bony palate and otic region. Postilla 207, 1–20 (1991).

    Google Scholar 

  39. Elżanowski, A. Skulls of Gobipteryx (Aves) from the upper Cretaceous of Mongolia. Acta Palaeontol. Pol. 37, 153–165 (1977).

    Google Scholar 

  40. Elżanowski, A. & Wellnhofer, P. Cranial morphology of Archaeopteryx: evidence from the seventh skeleton. J. Vertebr. Paleontol. 16, 81–94 (1996).

    Article  Google Scholar 

  41. Chiappe, L. M., Norell, M. & Clark, J. A new skull of Gobipteryx minuta (Aves: Enantiornithes) from the Cretaceous of the Gobi Desert. Am. Mus. Novit. 2001, 1–15 (2001).

    Article  Google Scholar 

  42. Zhang, Z., Chiappe, L. M., Han, G. & Chinsamy, A. A large bird from the Early Cretaceous of China: new information on the skull of enantiornithines. J. Vertebr. Paleontol. 33, 1176–1189 (2013).

    Article  CAS  Google Scholar 

  43. Xu, L. et al. A new, remarkably preserved, enantiornithine bird from the Upper Cretaceous Qiupa Formation of Henan (central China) and convergent evolution between enantiornithines and modern birds. Geol. Mag. 158, 2087–2094 (2021).

    Article  Google Scholar 

  44. Wang, M., Stidham, T. A., Li, Z., Xu, X. & Zhou, Z. Cretaceous bird with dinosaur skull sheds light on avian cranial evolution. Nat. Commun. 12, 3890 (2021).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  45. De Beer, S. G. The evolution of ratites. Bull. Brit. Mus. Nat. Hist. Zool. 4, 59–70 (1956).

    Google Scholar 

  46. Maxwell, E. E. Comparative ossification and development of the skull in palaeognathous birds (Aves: Palaeognathae). Zool. J. Linn. Soc. 156, 184–200 (2009).

    Article  Google Scholar 

  47. Worthy, T. H., Degrange, F. J., Handley, W. D. & Lee, M. S. The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres). R. Soc. Open Sci. 4, 170975 (2017).

    Article  PubMed  PubMed Central  Google Scholar 

  48. Dyke, G. J., Schulp, A. S. & Jagt, J. W. Bird remains from the Maastrichtian type area (Late Cretaceous). Neth. J. Geosci. 87, 353–358 (2008).

    Google Scholar 

  49. Dumont, M. et al. Synchrotron imaging of dentition provides insights into the biology of Hesperornis and Ichthyornis, the “last” toothed birds. BMC Evol. Biol. 16, 178 (2016).

    Article  PubMed  PubMed Central  Google Scholar 

  50. Wang, M., Li, Z., Liu, Q. & Zhou, Z. Two new early cretaceous ornithuromorph birds provide insights into the taxonomy and divergence of Yanornithidae (Aves: Ornithothoraces). J. Syst. Paleontol. 18, 1805–1827 (2020).

    Article  Google Scholar 

  51. Bell, A. & Chiappe, L. M. A species-level phylogeny of the Cretaceous Hesperornithiformes (Aves: Ornithuromorpha): implications for body size evolution amongst the earliest diving birds. J. Syst. Paleontol. 14, 239–251 (2016).

    Article  Google Scholar 

  52. Bell, A. & Chiappe, L. M. The Hesperornithiformes: a review of the diversity, distribution, and ecology of the earliest diving birds. Diversity 14, 267 (2022).

    Article  Google Scholar 

  53. Tanaka, T., Kobayashi, Y., Kurihara, K. I., Fiorillo, A. R. & Kano, M. The oldest Asian hesperornithiform from the Upper Cretaceous of Japan, and the phylogenetic reassessment of Hesperornithiformes. J. Syst. Paleontol. 16, 689–709 (2018).

    Article  Google Scholar 

  54. Goloboff, P. A. & Catalano, S. A. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32, 221–238 (2016).

    Article  PubMed  Google Scholar 

  55. Ronquist, F. et al. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol. 61, 539–542 (2012).

    Article  PubMed  PubMed Central  Google Scholar 

  56. Miller, M. A., Pfeiffer, W. & Schwartz, T. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In 2010 Gateway Computing Environments Workshop (GCE)pp. 1–8 (IEEE, 2010).

  57. Lewis, P. O. A likelihood approach to estimating phylogeny from discrete morphological character data. Syst. Biol. 50, 913–925 (2001).

    Article  CAS  PubMed  Google Scholar 

  58. Wang, M., O’Connor, J. K., Pan, Y. & Zhou, Z. A bizarre Early Cretaceous enantiornithine bird with unique crural feathers and an ornithuromorph plough-shaped pygostyle. Nat. Commun. 8, 14141 (2017).

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  59. Atterholt, J., Hutchison, J. H. & O’Connor, J. K. The most complete enantiornithine from North America and a phylogenetic analysis of the Avisauridae. PeerJ 6, e5910 (2018).

    Article  PubMed  PubMed Central  Google Scholar 

  60. O’Connor, J. K., Chiappe, L. M. & Bell, A. in Living Dinosaurs: the Evolutionary History of Modern Birds (eds Dyke, G. J. & Kaiser, G.) 39–114 (Wiley-Blackwell, 2011).

  61. O’Connor, P. M. et al. Late Cretaceous bird from Madagascar reveals unique development of beaks. Nature 588, 272–276 (2020).

    Article  PubMed  Google Scholar 

  62. Zelenkov, N. V., Averianov, A. O. & Popov, E. V. An Ichthyornis-like bird from the earliest Late Cretaceous (Cenomanian) of European Russia. Cretaceous Res. 75, 94–100 (2017).

    Article  Google Scholar 

  63. Mohsen, A., Hirayama, R., AbdelGawad, M., Sileem, A. & Aly, M. in Journal of Vertebrate Paleontology, Program and Abstracts. Society of Vertebrate Paleontology 80 (2020).

  64. Olson, S. L. et al. The anseriform relationships of Anatalavis Olson and Parris (Anseranatidae), with a new species from the Lower Eocene London Clay. Smithson. Contrib. Paleobiol. 89, 231–243 (1999).

    Google Scholar 

  65. Harrison, J. O. & Walker, C. A. A review of the bony-toothed birds (Odontopterygiformes): with descriptions of some new species. Tertiary Res. Special Pap. 2, 1–62 (1976).

    Google Scholar 

  66. Bourdon, E., Amaghzaz, M. & Bouya, B. Pseudotoothed birds (Aves, Odontopterygiformes) from the early Tertiary of Morocco. Am. Mus. Novit. 2010, 1–71 (2010).

    Article  Google Scholar 

  67. Adams, D. C., Collyer, M., Kaliontzopoulou, A. & Sherratt, E. Geomorph: software for geometric morphometric analyses. R package version 4.0. 2021. (2021).

  68. R Core Team. R: A Language and Environment for Statistical Computing. (R Foundation for Statistical Computing, 2020).

  69. Bjarnason, A. & Benson, R. A 3D geometric morphometric dataset quantifying skeletal variation in birds. MorphoMuseuM 7, e125 (2021).

    Article  Google Scholar 

  70. Zelditch, M. L., Swiderski, D. L. & Sheets, H. D. Geometric Morphometrics for Biologists: a Primer (Academic Press, 2004).

  71. Foth, C., Hedrick, B. P. & Ezcurra, M. D. Cranial ontogenetic variation in early saurischians and the role of heterochrony in the diversification of predatory dinosaurs. PeerJ 4, e1589 (2016).

    Article  PubMed  PubMed Central  Google Scholar 

  72. Field, D. J., Lynner, C., Brown, C. & Darroch, S. A. Skeletal correlates for body mass estimation in modern and fossil flying birds. PLoS ONE 8, e82000 (2013).

    Article  PubMed  PubMed Central  Google Scholar 

  73. Dunning Jr, J. B. CRC Handbook of Avian Body Masses (CRC Press, 2007).

Download references


We thank R. Dortangs for collecting the specimen, K. Smithson for scanning support, M. Lowe for collections assistance, B. Creisler for etymological information, S. Finney for preparation assistance, G. Navalón for assistance with morphometric methods, A. Chen and L. Steell for proofreading and discussion, and R. Olivé for permission to use his artwork. J.B. acknowledges support from the Hesse Award from the American Ornithological Society and grants from the Jurassic Foundation and Paleontological Society. This work was funded by UKRI grant MR/S032177/1 to D.J.F. For the purpose of open access, the authors have applied a Creative Commons Attribution (CC-BY) license to any Author Accepted Manuscript version arising.

Author information

Authors and Affiliations



D.J.F. and J.B. conceived and designed the project. D.J.F. supervised the project. J.W.M.J. facilitated sample access and provided provenance data. J.B., K.E.W. and P.-C.K. segmented CT scans. J.B. and D.J.F. identified the pterygoid. J.B. and P.-C.K. performed analyses. The manuscript was written by J.B. and D.J.F., with input from all of the authors.

Corresponding authors

Correspondence to Juan Benito or Daniel J. Field.

Ethics declarations

Competing interests

The authors declare no competing interests.

Peer review

Peer review information

Nature thanks Luis Chiappe and Christopher Torres for their contribution to the peer review of this work.

Additional information

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Extended data figures and tables

Extended Data Fig. 1 Best-preserved cervical and thoracic vertebrae of Janavis finalidens.

(a) 6th or 7th cervical vertebra; (b) 7th or 8th cervical vertebra; (c) 1st thoracic vertebra (12th presacral); (d) 2nd thoracic vertebra (13th presacral); (e) 3rd thoracic vertebra (14th presacral); (f) 4th thoracic vertebra (15th presacral); (g) indeterminate mid-thoracic vertebra; (h) fragmentary indeterminate mid-thoracic vertebra. Scale bar equals 10 mm.

Extended Data Fig. 2 Additional non-vertebral skeletal elements of Janavis finalidens and comparative elements of Ichthyornis.

All elements belong to Janavis unless otherwise specified. (a) Isolated tooth; (b) left pterygoid; (c) left scapula; (d) detailed view of omal end of left scapula; (e) thoracic ribs; (f) ribcage (previously identified as lower jaws and portion of the zygoma; Dyke et al.19), with the five ribs coloured orange, green, yellow, and red, and the two thoracic vertebrae in blue and purple; (g) Cross sections of omal end of Janavis and Ichthyornis scapulae, showing the well preserved bone surface around the acromion region of Janavis; left, Janavis scapula with cross-sectional planes 1-4 indicated; top, cross-sections through Janavis scapula (including surrounding rock matrix); bottom, cross-sections through Ichthyornis scapula at equivalent positions (FHSM VP-18702, mirrored); (h) reconstructed left humerus; (i) distorted left humerus as preserved; (j) mirrored right humerus of Ichthyornis specimen KUVP 119673; (k) right manual phalanx II:1; (l) right manual phalanx II:1 of Ichthyornis specimen ALMNH PV93.2.133; (m) proximal portion of right femur; (n) fragmentary pedal phalanx. Ichthyornis specimen figures (I and K) modified from Benito et al.22. All scale bars equal 10 mm.

Extended Data Fig. 3 CT-cross sections and additional views illustrating pneumaticity of the thoracic vertebrae of Janavis finalidens and the apneumatic condition of Ichthyornis.

(a) 15th vertebra of Janavis; (b) 15th vertebra of Ichthyornis; (c) indeterminate mid-thoracic vertebra of Janavis; (d) indeterminate mid-thoracic vertebra of Ichthyornis. Position of cross-sections 1-6 shown on the left for each specimen. (e) Additional views of Janavis vertebrae illustrating the position and extent of pneumatic foramina; left to right: indeterminate mid-thoracic vertebra in caudolateral view, indeterminate mid-thoracic vertebra from (c) in dorsolateral (left) and caudolateral (right) views, and 15th vertebra from (b) in caudolateral view. Small red arrows in (a) and (c) indicate pneumatic foramina situated within the pleurocoels, and large red arrow in (a) corresponds to the large ventral pneumatic opening. Scale bars equal 10 mm.

Extended Data Fig. 4 Detailed pterygoid morphology of Janavis finalidens and selected neornithines.

All are left pterygoids except those of Anatalavis, Anas, Megapodius and Lophophorus, which are mirrored right pterygoids. BPC, contact with basipterygoid process; PC, contact with palatine (or hemipterygoid); PF, pneumatic foramen; QAS, quadrate articular surface; RP, rostral process. All scale bars equal 2.5 mm.

Extended Data Fig. 5 Detailed pterygoid morphology of selected neornithines.

Illustrated pterygoids from right side, except that of Pelecanus, which is a mirrored left pterygoid. BPC, contact with basipterygoid process; PC, contact with palatine (or hemipterygoid); PF, pneumatic foramen; QAS, quadrate articular surface. All scale bars equal 2.5 mm.

Extended Data Fig. 6 Phylogenetic position of Janavis finalidens.

(a) topology recovered from Bayesian analysis of a modified version of the morphological matrix from Wang et al.50 and Benito et al.22; (b) topology recovered from Bayesian analysis of a modified version of the morphological matrix from Torres et al.12 and Benito et al.22. Node values indicate Bayesian posterior probabilities. Genus names in white indicate taxa illustrated on the right side of the figure, with corresponding illustrations indicated by superscript numbers (illustrations not to scale). Branches in white indicate the position of Janavis, the focal taxon of this study. Illustrations are courtesy of R. Olivé, used with permission.

Extended Data Fig. 7 Landmarking scheme for ornithurine pterygoids.

Taxa shown exemplify the range of variation of the principal morphological structures within our sample. (a) pseudo-landmark curve delimiting the quadrate contact of the pterygoid in caudal view; (b) pseudo-landmark curve delimiting the hemipterygoid-palatine contact of the pterygoid in rostral view; (c) pseudo-landmark curve characterizing the shape of the “dorsal crest” of the pterygoid in dorsal view; (d) pseudo-landmark curve characterizing the shape of the “medial crest” of the pterygoid, which includes the rostral process in those taxa which possess one, in medial view; (e) pseudo-landmark curve characterizing the shape of the “ventral crest” of the pterygoid in ventral view; (f) pseudo-landmark curve delimiting the shape of the basipterygoid process contact of the pterygoid.

Extended Data Fig. 8 Bivariate morphospaces of three-dimensional avian pterygoid geometry.

Plots illustrate morphospace distribution on the three first principal component axes; proportion of geometric variance explained by each principal component indicated on axis labels. (a) PC1 vs PC2; (b) PC1 vs PC3; and (c) PC2 vs PC3. Colours indicate the phylogenetic affinities of the included taxa: Palaeognathae (yellow), Galliformes (burgundy), Anseriformes (pink), Neoaves (blue); fossil taxa indicated in grey and numbered (1 – Janavis, 2 – Dasornis, 3 – Anatalavis).

Extended Data Fig. 9 Geometric similarity among avialan pterygoids summarized using total Procrustes distance.

(a) heatmap of total Procrustes distance. All taxa included in our morphometric analyses are illustrated. Strong geometric similarity is indicated by blue portions of the heatmap (low Procrustes distances), weak geometric similarity is indicated by red portions of the heatmap (high Procrustes distances), and intermediate degrees of geometric similarity are indicated by shades of yellow. Heatmap illustrates strong geometric similarity between Janavis and Anseriformes, Galliformes, and Dasornis, weak geometric similarity with Palaeognathae, and limited similarity with Neoaves. Cladogram beneath the heatmap illustrates the phylogenetic relationships of all taxa included in the analyses (Dasornis is illustrated as an early stem anseriform, though this is only one of several potentially viable phylogenetic placements for Pelagornithidae; see main text). (b) summarized Procrustes distance values from Janavis to major neornithine clades, showing the minimum, maximum and mean distances between Janavis and representatives of Galliformes, Anseriformes, Neoaves and Palaeognathae. Distances are lowest between Janavis and members of Galliformes and Anseriformes; cell background colours correspond to the heatmap colour scale from (a). (c) Box plots illustrating the Procrustes distance distribution between Janavis and major neornithine clades. PD = Procrustes Distance. n = 32 independent taxa compared with Janavis (Dasornis excluded due to its uncertain phylogenetic position); Galliformes: mean = 0.29, median = 0.29, minimum = 0.24, 1st quartile (25th percentile) = 0.26, 3rd quartile (75th percentile) = 0.32, maximum = 0.34; Anseriformes: mean = 0.37, median = 0.36, minimum = 0.32, 1st quartile (25th percentile) = 0.33, 3rd quartile (75th percentile) = 0.41, maximum = 0.44; Neoaves: mean = 0.44, median = 0.44, minimum = 0.35, 1st quartile (25th percentile) = 0.38, 3rd quartile (75th percentile) = 0.49, maximum = 0.55; Palaeognathae: mean = 0.78, median = 0.79, minimum = 0.70, 1st quartile (25th percentile) = 0.72, 3rd quartile (75th percentile) = 0.83, maximum = 0.84.

Extended Data Fig. 10 Reconstructed ichthyornithine palatal morphology and comparative neornithine taxa.

(a) Skulls in palatal view, from left to right: composite Ichthyornithes skull reconstruction (see main text), Lophophorus impejanus, and Struthio camelus. Ichthyornithes palatal reconstruction produced using Ichthyornis dispar skull model from Field et al.24; the quadrate is a composite of I. dispar specimens BHI 6421 and AMNH FARB 32773 (yellow), palatine and hemipterygoid of I. dispar AMNH FARB 32773 (green, from Torres et al.11), and pterygoid from Janavis finalidens at 50% scale (blue). (b) Isolated views of composite Ichthyornithes palate morphology from A; (c) focused views of the quadrate-pterygoid contact (top) and the pterygoid-hemipterygoid/palatine contact (bottom) in anatomical connection and separated, in approximately ventral view, showing the close match between the contacts of the Ichthyornis quadrate and palatine-hemipterygoid and the Janavis pterygoid at 50% scale. Janavis pterygoid in (a) illustrated at 100%; Ichthyornis skull components in (a) illustrated at 200%. Scale bars equal 10 mm.

Supplementary information

Supplementary Information

Merged PDF including table of contents, additional background/provenance data, further detail on phylogenetic analyses, further detail on geometric morphometric analyses, morphological variation in the pterygoid and basisphenoid, additional morphological description and discussion, institutional abbreviations, scan parameters, specimens included in the morphometric analyses, other comparative specimens, morphometric character descriptions, synapomorphies diagnosing key clades, morphological character descriptions, MorphoSource links, supplementary references.

Reporting Summary

Rights and permissions

Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.

Reprints and Permissions

About this article

Check for updates. Verify currency and authenticity via CrossMark

Cite this article

Benito, J., Kuo, PC., Widrig, K.E. et al. Cretaceous ornithurine supports a neognathous crown bird ancestor. Nature 612, 100–105 (2022).

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI:

This article is cited by


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.


Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing