Publications and CV

Bhart-Anjan S. Bhullar CV: 

Option one hyoid
Griffin et al. (2022) Nature
Bhullar & Manafzadeh et al. (2019) Nature
Field & Hanson et al. (2018) Nature
Hanson et al. (2022) Nature

* indicates Bhullar Lab student or postdoc

71. Griffin CT*, Botelho JF*, Hanson M*, Fabbri M*, Smith-Paredes D*, Carney RM, Norell MA, Egawa S*, Gatesy SM, Rowe TB, Elsey RM, Nesbitt SJ, Bhullar B-AS. 2022. The developing bird pelvis passes through ancestral dinosaurian conditions. Nature 608: 346–352.


70. Smith-Paredes D*, Vergara-Cereghino ME, Lord A, Moses MM, Behringer RR, Bhullar B-AS. 2022. Embryonic muscle splitting patterns reveal homologies of amniote forelimb muscles. Nature Ecology & Evolution 6(5): 604-613.

69. Egawa S*, Griffin CT*, Bishop P, Pintore R, Tsai HP, Botelho JF*, Smith-Paredes D*, Kuratani S, Norell MA, Nesbitt S, Hutchinson JR, Bhullar B-AS. In press. The femoral head of birds evolved via a shift from torsion to growth along the avian stem. Proceedings of the Royal Society B.


68. Fabbri M*, Navalón G, Benson RBJ, Pol D, O’Connor J, Bhullar B-AS, Erickson GM, Norell MA, Orkney A, Lamanna MC, Zouhri S, Becker J, Emke A, Dal Sasso C, Bindellini G, Maganuco S, Auditore M, Ibrahim N. 2022. Subaqueous foraging among carnivorous dinosaurs. Nature 603(7903): 852-857.

67. Jenkins KM*, Meyer DL, Lewis PJ, Choiniere JN, Bhullar B-AS. 2022. Re-description of the early Triassic diapsid Palacrodon from the lower Fremouw formation of Antarctica. Journal of Anatomy Online Early View.

66. Chen A, Field D*, Benito J, Wilson LE, Bhullar B-AS, Burnham D. 2022. In press. 40 new specimens of Ichthyornis provide unprecedented insight into the postcranial morphology of crownward stem group birds. PeerJ.

65. Brownstein C, Meyer D, Fabbri M*, Bhullar B-AS, Gauthier JA. 2022. In press. Three-dimensionally preserved crown lizard skulls from the Jurassic illuminate the prolonged rise of the extant squamate radiation. Nature Communications. 

64. Fabbri M*, Bhullar B-AS. In press. The endocast of Euparkeria sheds light on the ancestral archosaur nervous system. Palaeontology.

63. Smith KT, Bhullar B-AS, Bloch JI. 2022. New diminutive Eocene lizard reveals high K-Pg survivorship and taxonomic diversity of stem xenosaurs in North America. American Museum Novitates 3986: 1-36.

62. Jenkins KM*, Bhullar B-AS. 2022. Tooth Implantation and Attachment in Scoloparia glyphanodon (Parareptilia: Procolophonidae). Bulletin of the Peabody Museum of Natural History 63(1): 27-30.


61. Hanson M*, Hoffman EA, Norell MA, Bhullar B-AS. 2022. Response to Comment on “The early origin of a birdlike inner ear and the evolution of dinosaurian movement and vocalization”. Science 376(6600): eabl8181.


60. Yohe LR*, Fabbri M*, Lee D, Davies KTJ, Yohe TP, Sánchez MKR, Rengifo EM, Hall R, Mutumi G, Hedrick BP, Sadier A, Simmons NB, Sears KE, Dumont E, Rossiter SJ, Bhullar B-AS, Dávalos LM. 2022. Ecological constraints on highly evolvable olfactory receptor genes and morphology in neotropical bats. Evolution (Online Early View).


59. Luo Z-X, Bhullar B-AS, Crompton AW, Neander AI, Rowe TB. 2022. Reexamination of the mandibular and dental morphology of the Early Jurassic mammaliaform Hadrocodium wui. Acta Palaeontologica Polonica 67(1): 95-113.


58. Powers MJ, Fabbri M*, Doschak MR, Bhullar B-AS, Evans DC, Norell MA, Currie PJ. 2022. A new hypothesis of eudromaeosaurian evolution: CT scans assist in testing and constructing morphological characters. Journal of Vertebrate Paleontology 41 (5): e2010087.

57. Napoli JG, Ruebenstal AA*, Bhullar B-AS, Turner AH, Norell MA. 2021. A new dromaeosaurid (Dinosauria: Coelurosauria) from Khulsan, central Mongolia. American Museum Novitates 3982: 1-47.

56. Hanson M*, Hoffman EA, Norell MA, Bhullar B-AS. 2021. The early origin of a birdlike inner ear and the evolution of dinosaurian movement and vocalization. Science 372(6542): 601-609.

55. Fabbri M*, Navalon G, Mongiardino Koch N, Hanson M*, Petermann H, Bhullar B-AS. 2021. A shift in ontogenetic timing produced the unique sauropod skull. Evolution 75(4): 819-831.

54. Smith-Paredes D*, Lord A*, Meyer D*, Bhullar B-AS. 2021. A developmental staging system and musculoskeletal development sequence of the Common Musk Turtle (Sternotherus odoratus). Developmental Dynamics 250(1): 111-127.

53. Smith-Paredes D*, Griffith O., Fabbri M*, Yohe L*, Blackburn DG, Siler CD, Bhullar B-A.S., Wagner GP. 2021. Hidden limbs in the “limbless skink” Brachymeles lukbani: developmental observations. Journal of Anatomy 239(3): 693-703.

52. Navalón G, Nebreda SM, Bright JA, Fabbri M*, Benson RBJ, Bhullar B.-A.S. 2021. Craniofacial development illuminates the evolution of nightbirds (Strisores). Proceedings of the Royal Society B 288(1948): 20210181.

51. Blackburn DG, Barnes MS, Reimers CD, Appiah FA, Lestz LL, Bonneau LJ, Hanson M*, Smith-Paredes D.*, Bhullar B-AS. 2021. How do crocodylian embryos process yolk? Morphological evidence from the American alligator, Alligator mississippiensis. Journal of Morphology 282(7): 953-958.

50. Louchart A, Bhullar B-AS, Riamon S, Field DJ*. 2021. The true identity of putative tooth alveoli in a Cenozoic crown bird, the gastornithid Omorhamphus. Frontiers in Earth Science 9: 333.

49. Bhullar B-AS, Manafzadeh AR, Miyamae JA*, Hoffman EA, Brainerd EL, Musinsky C, Crompton AW. 2020. Reply to: Jaw roll and jaw yaw in early mammals. Nature 582(7812): E9-E12.

48. Yohe LR*, Fabbri M*, Hanson M*, Bhullar B-AS. 2020. Olfactory receptor gene evolution is unusually rapid across Tetrapoda and outpaces chemosensory phenotypic change. Current Zoology 66(5): 505-514.

47. Ksepka DT, Balanoff AM, Smith NA, Bever GS, Bhullar B-AS et al. 2020. Tempo and pattern of avian brain size evolution. Current Biology 30(11): P2026-P2036.E3.

46. Felice RN, Watanabe A., Cuff AR, Hanson M.*, Bhullar B-AS, Rayfield ER, Witmer LM, Norell MA, Goswami A. 2020. Decelerated dinosaur skull evolution with the origin of birds. PLoS Biology 18(18): e3000801.

45. Sookias RB, Dilkes D, Sobral G, Smith RMH, Wolvaardt FP, Arcucci AB, Bhullar B-AS, Werneburg I. 2020. The craniomandibular anatomy of the early archosauriform Euparkeria capensis and the dawn of the archosaur skull. Royal Society Open Science 7(7): 200116.

44. Brown EE, Butler RJ, Ezcurra MD, Bhullar B-AS, Lautenschlager S. 2020. Endocranial anatomy and life habits of the Early Triassic archosauriform Proterosuchus fergusi. Palaeontology 63(2): 255-282.

43. Bhullar B-AS, Manafzadeh AR, Miyamae JA*, Hoffman E, Brainerd EL, Musinsky C, Crompton AW. 2019. Rolling of the jaw is essential for mammalian chewing and tribosphenic molar function. Nature 566: 528-532.

42. Zhou C.-F., Bhullar B-AS (co-first), Neander AI, Martin T, Luo Z-X. 2019. New Jurassic mammaliaform sheds light on early evolution of mammal-like hyoid bones. Science 365(6450): 276-279.

41. Smith-Paredes D*, Bhullar B-AS. 2019. The skull and head muscles of Archosauria. In: Ziermann JM, Diaz RE Jr., Diogo R. Heads, Jaws, and Muscles, Springer: 229-251.

40. Camacho J, Heyde A, Bhullar B-AS, Haelwaters D, Simmons N, Abzhanov A. 2019. Peramorphosis, an evolutionary developmental mechanism in neotropical bat skull diversity. Developmental Dynamics 248(11): 1129-1143.

39. Field DJ*, Hanson M*, Burnham D, Wilson LE, Super K, Ehret D, Ebersole JA, Bhullar B-AS. 2018. Complete Ichthyornis skull illuminates mosaic assembly of the avian head. Nature 557: 96-100.

38. Pritchard AC*, Gauthier JA, Hanson M*, Bever GS, Bhullar B-AS. 2018. A tiny Triassic saurian from Connecticut and the early evolution of the diapsid feeding apparatus. Nature Communications 9(1): 1213.

37. Smith KT, Bhullar B-AS, Köhler G, Habersetzer J. 2018. The only known jawed vertebrate with four eyes and the bauplan of the pineal complex. Current Biology 28(7): 1101-1107.

36. Wagner GP, Griffith OW, Bergmann PJ, Bello-Hellegouarch, G., Kohlsdorf T, Bhullar A, Siler CD. 2018. Are there general laws for digit evolution in squamates? The loss and re-evolution of digits in a clade of fossorial lizards (Brachymeles, Scincinae). Journal of Morphology 279(8): 1104-1119.

35. Crompton AW, Musinsky C, Rougier GW, Bhullar B-AS, Miyamae J*. 2018. Origin of the lateral wall of the mammalian skull: fossils, monotremes and therians revisited. Journal of Mammalian Evolution 25: 301-313.

34. Rodrigues HG, Cornette R, Clavel J, Cassini G, Bhullar B-AS, Monescillo MF, Moreno K, Herrel A, Billet G. 2018. Differential influences of allometry, phylogeny and environment on the rostral shape diversity of extinct South American notoungulates. Royal Society Open Science 5(1): 171816.

33. Fabbri M*, Mongiardino Koch N, Pritchard AC*, Hanson M*, Hoffman E, Bever GS, Balanoff AM, Morris ZS*, Field DJ*, Camacho J, Rowe TB, Norell MA, Smith RM, Abzhanov A, Bhullar B-AS. 2017. The skull roof tracks the brain during the evolution and development of reptiles including birds. Nature Ecology & Evolution 1(10): 1543-1550.

32. Bhullar B-AS. 2017. Catastrophe triggers diversification. Nature 542: 304-305.

31. Schultz JA, Bhullar B-AS, Luo Z-X. 2017. Re-examination of the Jurassic mammaliaform Docodon victor by computed tomography and occlusal functional analysis. Journal of Mammalian Evolution 26: 9-38.

30. Clark EG, Bhullar B-AS, Darroch SAF, Briggs DEG. 2017. Water vascular system architecture in an Ordovician ophiuroid. Biology Letters. 13(12): 635.

29. Crompton AW, Owerkowicz T, Bhullar B-AS, Musinsky C. 2017. Structure of the nasal region of non-mammalian cynodonts and mammaliaforms: Speculations on the evolution of mammalian endothermy. Journal of Vertebrate Paleontology 37: e1269116.

28. Bhullar B-AS, Hanson M*, Fabbri M*, Pritchard A*, Bever GS, Hoffman E. 2016. How to make a bird skull: Major transitions in the evolution of the avian cranium, paedomorphosis, and the beak as a surrogate hand. Integrative and Comparative Biology 56(3): 389-403.

27. Sefton EM, Bhullar B-AS, Mohaddes Z, Hanken J. 2016. Evolution of the head-trunk interface in tetrapod vertebrates. eLife 5: e09972.

26. Chavan AR, Bhullar B-AS, Wagner GP. 2016. What was the ancestral function of decidual stromal cells? A model for the evolution of eutherian pregnancy. Placenta 40: 40-51.

25. Sobral G, Sookias RB, Bhullar B-AS, Smith R, Butler RJ, Müller J. 2016. New information on the braincase and inner ear of Euparkeria capensis Broom: implications for diapsid and archosaur evolution. Royal Society Open Science 3(7): 160072.

24. Bever GS, Lyson TR, Field DJ*, Bhullar B-AS. 2016. The amniote temporal roof and the diapsid origin of the turtle skull. Zoology 119(6): 471-473.

23. Dumont M, Tafforeau P, Bertin T, Bhullar BA, Field D*, Schulp A, Strilisky B, Thivichon-Prince B, Viriot L, Louchart A. 2016. Synchrotron imaging of dentition provides insights into the biology of Hesperornis and Ichthyornis, the “last” toothed birds. BMC Evolutionary Biology 16(1): 1-178.

22. Bhullar B-AS, Morris ZS*, Sefton EM, Tok A, Tokita M, Namkoong B, Camacho J, Burnham DA, Abzhanov A. 2015. A molecular mechanism for the origin of a key evolutionary innovation, the bird beak and palate, revealed by an integrative approach to major transitions in vertebrate history. Evolution 69(7): 1665-1677.  

21. Bever GS, Lyson T, Field DJ*, Bhullar B-AS. 2015. Evolutionary origin of the turtle skull. Nature 525: 239-242.

20. Molnar J, Pierce S, Bhullar B-AS, Turner A, Hutchinson J. 2015. Morphological and functional changes in the vertebral column with increasing aquatic adaptation in crocodylomorphs. Royal Society Open Science 2(11).

19. Scanferla CA, Bhullar B-AS. 2014. Postnatal development of the skull of Dinilysia patagonica (Squamata-stem Serpentes). The Anatomical Record 297(3): 560-573.

18. Kirk EC, Daghighi P, Macrini TE, Bhullar B-AS, Rowe TB. 2014. Cranial anatomy of the Duchesnean primate Rooneyia viejaensis: New insights from high-resolution computed tomography. Journal of Human Evolution 74: 82-95.

17. Roček Z, Wuttke M, Gardner JD, Bhullar B-AS. 2014. The Euro-American genus Eopelobates, and a re-definition of the family Pelobatidae (Amphibia, Anura). Palaeobiodiversity and Palaeoenvironments 94: 529-567.

16. Weeks O, Bhullar B-AS, Abzhanov A. 2013. Molecular characterization of dental development in a toothed archosaur, the American alligator Alligator mississippiensis. Evolution and Development 15(6): 393-405.

15. Lyson TR, Bhullar B-AS, Bever GS, Joyce WG, de Queiroz K, Abzhanov A, Gauthier JA. 2013. Homology of the enigmatic nuchal reveals the novel reorganization of the shoulder girdle in the evolution of the turtle shell. Evolution and Development 15(5): 317-325.

14. Bhullar B-AS, Marugán-Lobón J, Racimo F, Bever GS, Norell MA, Rowe TB, Abzhanov A. 2012. Birds have paedomorphic dinosaur skulls. Nature 487: 223-226.

13. Longrich NR, Bhullar B-AS, Gauthier JA. 2012. A transitional snake from the Late Cretaceous of North America. Nature 488: 205-208.

12. Longrich NR, Bhullar B-AS, Gauthier JA. 2012. Mass extinction of lizards and snakes at the Cretaceous-Paleogene boundary. Proceedings of the National Academy of Sciences 109(52): 21396-21401.

11. Bhullar B-AS. 2012. A phylogenetic approach to ontogeny and heterochrony in the fossil record: Cranial evolution and development in anguimorphan lizards (Reptilia: Squamata). Journal of Experimental Zoology, Part B: Molecular and Developmental Evolution 318b: 521-530.

10. Kennedy AM, Bhullar B-AS, Lewis PJ. In press. Preliminary analysis of the fossil herpetofauna from the Koanaka Hills site, Ngamiland, Northwestern Botswana. Palaeontologia Africana.

9. Ortiz D, Lewis PJ, Kennedy AM, Bhullar B-AS, Hancox J. In press. Preliminary analysis of Dipnoi (lungfish) fossils from Driefontein, South Africa. Palaeontologia Africana.

8. Bhullar B-AS. 2011. The power and utility of morphological characters in systematics: A fully resolved phylogeny of Xenosaurus and its fossil relatives (Squamata: Anguimorpha). Bulletin of the Museum of Comparative Zoology 160(3): 65-181.

7. Bhullar B-AS. 2010. Cranial osteology of Exostinus serratus, fossil sister taxon to the enigmatic clade Xenosaurus. Zoological Journal of the Linnean Society 159: 921-953.

6. Lyson, TR, Bever BS, Bhullar B-AS, Joyce WG, Gauthier JA. 2010. Transitional fossils and the origin of turtles. Biology Letters 6: 830-833.

5. Bhullar B-AS, Bever GS. 2009. An archosaur-like laterosphenoid in early turtles (Reptilia: Pantestudines). Breviora of the Museum of Comparative Zoology 518: 1-11.

4. Bhullar B-AS. 2009. A reevaluation of the unusual abdominal musculature of squamate reptiles (Reptilia: Squamata). The Anatomical Record 292(8): 1154-1161.

3. Bhullar B-AS, Bell CJ. 2008. Osteoderms of the California legless lizard Anniella (Squamata: Anguidae) and their relevance for considerations of miniaturization. Copeia 2008(4): 785-793.

2. Bhullar B-AS, Smith KT. 2008. Helodermatid lizard from the Miocene of Florida, the evolution of the dentary in Helodermatidae, and comments on dentary morphology in Varanoidea. Journal of Herpetology 42(2): 286-302.

1. Smith KT, Bhullar B-AS, Holroyd PA. 2008. Earliest record of the Varanus stem-clade (Squamata: Varanidae) from the Early Oligocene of Egypt. Journal of Vertebrate Paleontology 28(3): 909-913.