Recent publications by members of the Vertebrate Palaeobiology research group, University of Oxford.


  1. Benevento G. L., Benson R.B.J., Friedman M. 2019. Patterns of mammalian jaw ecomorphological disparity during the Mesozoic/Cenozoic transition. Proceedings of the Royal Society B: Biological Sciences, 286: 20190347.
  2. Chapelle K. M., Benson R.B.J., Stiegler J., Otero A., Zhao Q., Choiniere J. In press. A quantitative method for inferring locomotory shifts in amniotes during ontogeny, its application to dinosaurs, and its bearing on the evolution of posture. Palaeontology.
  3. Chen A., White N.D., Benson R.B.J., Braun M.J., Field D.J. 2019. Total-evidence framework reveals complex morphological evolution in nightbirds (Strisores). Diversity, 11(9): 143.
  4. Choo B., Lu J., Giles S., Trinajstic K., Long J. A. 2019. A new actinopterygian from the late Devonian Gogo formation, Western Australia. Papers in Palaeontology, 5(2): 343-363.
  5. Close R. A., Benson R. B. J., Alroy J., Behrensmeyer A., Benito J., Carrano M., Cleary T. J., Butler R. J. 2019. Diversity dynamics of Phanerozoic terrestrial tetrapods at the local-community scale. Nature Ecology and Evolution, 3: 590–597.
  6. Brocklehurst N. 2019. Morphological evolution in therocephalians breaks the hypercarnivore bracket. Proceedings of the Royal Society B: Biological Sciences, 286: 20190590.
  7. Davesne D.,  Meunier F.J., Schmitt A.D., Friedman M., Otero O., Benson R.B.J. 2019. The phylogenetic origin and evolution of acellular bone in teleost fishes: insights into osteocyte function in bone metabolism. Biological Reviews, 94(4): 1338-1363.
  8. Dobson C., Giles S., Johanson Z., Liston J., Friedman M. In press. Cranial osteology of the Middle Jurassic (Callovian) Martillichthys renwickae (Neopterygii, Pachycormiformes) with comments on the evolution and ecology of edentulous pachycormiforms. Papers in Palaeontology, DOI: 10.1002/spp2.1276
  9. Evers S. W., Benson R. B. J. 2019. A new phylogenetic hypothesis of turtles with implications for the timing and number of evolutionary transitions to marine lifestyles in the group. Palaeontology, 62(1): 93-134.
  10. Evers S. W., Barrett P. M., Benson R. B. J. 2019. Anatomy of Rhinochelys pulchriceps (Protostegidae) and marine adaptation during the early evolution of chelonioids. PeerJ, 7: e6811.
  11. Evers S. W., Neenan J. M., Ferreira G. S., Werneburg I., Barrett P. M., Benson R. B. J. In press. Neurovascular anatomy of the protostegid turtle Rhinochelys pulchriceps and comparisons of the membranous and endosseous labyrinth shape in an extant turtle. Zoological Journal of the Linnean Society, DOI: 10.1093/zoolinnean/zlz063
  12. Ford D. P., Benson R. B. J. 2019. A redescription of Orovenator mayorum (Sauropsida: Diapsida) using high-resolution μCT, and its consequences for early amniote phylogeny. Papers in Palaeontology, 5(2): 197-239.
  13. Foth C., Evers S. W., Joyce W. G., Volpato V. S., Benson R. B. J. In press. Comparative analysis of the shape and size of the middle ear cavity of turtles reveals no correlation with habitat ecology. Journal of Anatomy, DOI: 10.1111/joa.13071
  14. Friedman M., Feilich K. L., Beckett H. T., Alfaro M. E., Faircloth B. C., Černý D., Miya M., Near T. J., Harrington R. C. 2019. A phylogenomic framework for pelagiarian fishes (Acanthomorpha: Percomorpha) highlights mosaic radiation in the open oceanProceedings of the Royal Society B: Biological Sciences, 286: 20191502.
  15. Friedman M., Pierce S. E., Coates M., Giles S. 2019. Feeding structures in the ray-finned fish Eurynotus crenatus (Actinopterygii: Eurynotiformes): implications for trophic diversification among Carboniferous actinopterygians. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 109(1-2):  33-47.
  16. Godoy P.L., Benson R.B.J., Bronzati M., Butler R.J. 2019. The multi-peak adaptive landscape of crocodylomorph body size evolution. BMC Evolutionary Biology, 19: 167.
  17. Grunert H. R., Brocklehurst N., Fröbisch J. 2019. Diversity and disparity of Therocephalia: macroevolutionary patterns through two mass extinctions. Scientific Reports, 9: 5063.
  18. Hedrick B. P., Antalek-Schrag P., Conith A. J., Natanson L. J., Brennan P. L. R. 2019. Variability and asymmetry in the shape of the spiny dogfish vagina revealed by 2D and 3D geometric morphometrics. Journal of Zoology, 308(1): 16-27.
  19. Hedrick B. P., Goldsmith E., Rivera-Sylva H., Fiorillo A.R., Tumarkin-Deratzian A.R., Dodson P. In press. Filling in gaps in the ceratopsid histologic database: histology of two basal centrosaurines and an assessment of the utility of rib histology in the Ceratopsidae. The Anatomical Record, DOI: 0.1002/ar.24099
  20. Hedrick B. P., Schachner E. R., Rivera G., Dodson P., Pierce S. E. 2019. The effects of skeletal asymmetry on interpreting biologic variation and taphonomy in the fossil record. Paleobiology, 45(1): 154-166.
  21. Legendre L.J., Davesne D. In press. The evolution of mechanisms involved in vertebrate endothermy. Philosophical Transactions of the Royal Society B. DOI: 10.1098/rstb.2019.0136
  22. Lukic-Walther M., Brocklehurst N., Kammerer C. F., Fröbisch J. 2019.  Diversity patterns of nonmammalian cynodonts (Synapsida, Therapsida) and the impact of taxonomic practice and research history on diversity estimates. Palaeobiology, 45(1): 56-69.
  23. MacDougall M. J., Brocklehurst N., Fröbisch J. 2019. Species richness and disparity of parareptiles across the end-Permian mass extinction. Proceedings of the Royal Society B: Biological Sciences, 286: 20182572.
  24. Neenan J. M., Chapelle K. M., Fernandez V., Choiniere J. 2019. Ontogeny of the Massospondylus labyrinth: implications for locomotory shifts in a basal sauropodomorph dinosaur. Palaeontology, 62(2): 255-265.
  25. Panciroli E., Benson R. B. J., Luo Z.-X. In press. The mandible and dentition of Borealestes serendipitus (Docodonta) from the Middle Jurassic of Skye, Scotland. Journal of Vertebrate Paleontology, DOI: 10.1080/02724634.2019.1621884


  1. Albino A. M., Rothschild B., Carrillo-Briceño J. D., Neenan J. M. 2018. Spondyloarthropathy in vertebrae of the aquatic Cretaceous snake Lunaophis aquaticus, and its first recognition in modern snakes. The Science of Nature, 105: 51.
  2. Beckett H. T., Giles S., Friedman M. 2018. Comparative anatomy of the gill skeleton of fossil Aulopiformes (Teleostei: Eurypterygii). Journal of Systematic Palaeontology, 16 (14): 1221-1245.
  3. Beckett H. T., Giles S., Johanson Z., Friedman M. 2018. Morphology and phylogenetic relationships of fossil snake mackerels and cutlassfishes (Trichiuroidea) from the Eocene (Ypresian) London Clay Formation. Papers in Palaeontology, 4 (4): 577-603.
  4. Benson R. B. J. 2018. Dinosaur macroevolution and macroecology. Annual Review of Ecology, Evolution, and Systematics, 49: 379-408.
  5. Benson R. B. J., Hunt G., Carrano M. T., Campione N. 2018. Cope’s rule and the adaptive landscape of dinosaur body size evolution. Palaeontology, 61 (1): 13-48.
  6. Bronzati M., Benson R. B. J., Rauhut O. W. M. 2018. Rapid transformation in the braincase of sauropod dinosaurs: integrated evolution of the braincase and neck in early sauropods? Palaeontology, 61 (2): 289-302.
  7. Cleary T. J., Benson R. B. J., Evans S. E., Barrett P. M. 2018. Lepidosaurian diversity in the Mesozoic–Paleogene: the potential roles of sampling biases and environmental drivers. Royal Society Open Science, 5: 171830.
  8. Clement A. M., King B., Giles S., Choo B., Ahlberg P. E., Young G. C., Long J. A. 2018. Neurocranial anatomy of an enigmatic Early Devonian fish sheds light on early osteichthyan evolution. eLife, 7: e34349.
  9. Close R. A., Evers S. W., Alroy J., Butler R. J. 2018. How should we estimate diversity in the fossil record? Testing richness estimators using sampling‐standardised discovery curves. Methods in Ecology and Evolution, 9 (6): 1386-1400.
  10. Davesne D., Gueriau P., Dutheil D. B., Bertrand L. 2018. Exceptional preservation of a Cretaceous intestine provides a glimpse of the early ecological diversity of spiny-rayed fishes (Acanthomorpha, Teleostei). Scientific Reports, 8: 8509.
  11. Davesne D., Meunier F. J., Friedman M., Benson R. B. J., Otero O. 2018. Histology of the endothermic opah (Lampris sp.) suggests a new structure-function relationship in teleost fish bone. Biology Letters, 14: 20180270.
  12. Day M. O., Benson R. B. J., Kammerer C. F., Rubidge B. S. 2018. Evolutionary rates of mid-Permian tetrapods from South Africa and the role of temporal resolution in turnover reconstruction. Paleobiology, 44(3): 447-467.
  13. Fischer V., Benson R. B. J., Druckenmiller P. S., Ketchum H. F., Bardet N. 2018. The evolutionary history of polycotylid plesiosaurians. Royal Society Open Science, 5: 172177.
  14. Giles S., Rogers M., Friedman M. 2018. Bony labyrinth morphology in early neopterygian fishes (Actinopterygii: Neopterygii). Journal of Morphology, 279: 426–440.
  15. Latimer A., Giles S. 2018. A giant dapediid from the Late Triassic of Switzerland and insights into neopterygian phylogeny. Royal Society Open Science, 5: 180497.
  16. McPhee B., Benson R. B. J., Botha-Brink J., Bordy E. M., Choiniere J. N. 2018. A giant dinosaur from the earliest Jurassic of South Africa and the transition to quadrupedality in early sauropodomorphs. Current Biology, 28 (19): 3143-3151.
  17. Panciroli E., Benson R. B. J., Butler R. J. 2018. New partial dentaries of amphitheriid mammal Palaeoxonodon ooliticus from Scotland, and posterior dentary morphology in early cladotheriansActa Palaeontologica Polonica, 63 (2): 197-206.
  18. Schwarzhans W., Beckett H. T., Schein J. D., Friedman M. 2018. Computed tomography scanning as a tool for linking the skeletal and otolith-based fossil records of teleost fishes. Palaeontology, 61: 511-541.
  19. Xu X., Choiniere J., Tan Q., Benson R. B. J., Clark J., Sullivan C., Zhao Q., Han F., Ma Q., He Y., Wang S., Xing H., Tan L. 2018. Two Early Cretaceous fossils document transitional stages in alvarezsaurian dinosaur evolution. Current Biology, 28(7): 2853-2860.
  20. Zverkov N. G., Fischer V., Madzia D., Benson R. B. J. 2018. Increased pliosaurid dental disparity across the Jurassic-Cretaceous transition. Palaeontology, 61 (6): 825-846.


  1. Benson R. B. J., Starmer-Jones E., Close R. A., Walsh S. A. 2017. Comparative analysis of vestibular ecomorphology in birds. Journal of Anatomy, 231: 990–1018.
  2. Bjarnason A., Soligo C., Elton S. 2017. Phylogeny, phylogenetic inference, and cranial evolution in pitheciids and Aotus. American Journal of Primatology, 79: e22621.
  3. Close R. A., Benson R. B. J., Upchurch P., Butler R. J. 2017. Controlling for the species-area effect supports constrained long-term Mesozoic terrestrial vertebrate diversification. Nature Communications, 8: 15381.
  4. Crofts S. B., Neenan J. M., Scheyer T. M., & Summers A. P. 2017. Tooth occlusal morphology in the durophagous marine reptiles, Placodontia (Reptilia: Sauropterygia). Paleobiology, 43 (1): 114-128.
  5. Davesne D. 2017. A fossil unicorn crestfish (Teleostei, Lampridiformes, Lophotidae) from the Eocene of Iran. PeerJ, 5: e3381.
  6. Davesne D., Carnevale G., Friedman M. 2017. Bajaichthys elegans from the Eocene of Bolca and the overlooked anatomical diversity of Zeiformes (Teleostei, Acanthomorpha). Palaeontology, 60 (2): 255-268.
  7. Davies T. G., Rahman I. A., Lautenschlager S., Cunningham J. A., Asher R. J., Barrett P. M., Bates K. T., Bengtson S., Benson R. B. J., Boyer D. M., Braga J., Bright J. A., Claessens L. P. A. M., Cox P. G., Dong X-P., Evans A. R., Falkingham P. L., Friedman M., Garwood R. J., Goswami A., Hutchinson J. R., Jeffery N. S., Johanson Z., Lebrun R., Martínez-Pérez C., Marugán-Lobón J., O’Higgins P. M., Metscher B., Orliac M., Rowe T. B., Rücklin M., Sánchez-Villagra M. R., Shubin N. H., Smith S. Y., Starck J. M., Stringer C., Summers A. P., Sutton M. D., Walsh S. A., Weisbecker V., Witmer L. M., Wroe S., Yin Z., Rayfield E. J., Donoghue P. C. J. 2017. Open data and digital morphology. Proceedings of the Royal Society B: Biological Sciences, 284: 20170194.
  8. Fischer V., Benson R. B. J., Zverkov N. G., Soul L. C., Arkhangelsky M. S., Lambert O., Stenshin I. M., Uspensky G. N., Druckenmiller P. S. 2017. Plasticity and convergence in the evolution of short-necked plesiosaurs. Current Biology, 27: 1667–1676
  9. Giles S., Xu G.-H., Near T. J., Friedman M. 2017. Early members of ‘living fossil’ lineage imply later origin of modern ray-finned fishes. Nature, 549: 265–268.
  10. Lu J., Giles S., Friedman M., Zhu M. 2017. A new stem-sarcopterygian illuminates patterns of character evolution in early bony fishes. Nature Communications, 8: 1932.
  11. Martín-Serra A., Figueirido B., Palmqvist P. 2017. Non-decoupled morphological evolution of the fore- and hindlimb of sabretooth predators. Journal of Anatomy, 231: 532–542.
  12. Mennecart B., DeMiguel D., Bibi F., Rössner G. E., Métais G., Neenan J. M., Wang S., Schulz G., Müller B., Costeur L. 2017. Bony labyrinth morphology clarifies the origin and evolution of deer. Scientific Reports, 7: 13176.
  13. Neenan J. M., Reich T., Evers S. W., Druckenmiller P. S., Voeten D. F. A. E., Choiniere J. N., Barrett P. M., Pierce S. E., Benson R. B. J. 2017. Evolution of the sauropterygian labyrinth with increasingly pelagic lifestylesCurrent Biology, 27: 3852–3858.
  14. Panciroli E., Benson R. B. J., Walsh S. 2017. The dentary of Wareolestes rex (Megazostrodontidae): a new specimen from Scotland and implications for morganucodontan tooth replacement. Papers in Palaeontology, 3: 373–386.
  15. Pierce S. E., Williams M., Benson R. B. J. 2017. Virtual reconstruction of the endocranial anatomy of the early Jurassic marine crocodylomorph Pelagosaurus typus (Thalattosuchia)PeerJ, 5: e3225.
  16. Sallan L., Giles S., Sansom R. S., Clarke J. T., Johanson Z., Sansom I. J., Janvier P. 2017. The ‘Tully Monster’ is not a vertebrate: characters, convergence and taphonomy in Palaeozoic problematic animals. Palaeontology, 60 (2): 149-157.
  17. Scheyer T. M., Neenan J. M., Bodogan T., Furrer H., Obrist C., Plamondon M. 2017. A new, exceptionally preserved juvenile specimen of Eusaurosphargis dalsassoi (Diapsida) and implications for Mesozoic marine diapsid phylogeny. Scientific Reports, 7: 4406.
  18. Serratos D. J., Druckenmiller P., Benson R. B. J. 2017. A new elasmosaurid (Sauropterygia, Plesiosauria) from the Bearpaw Shale (Late Cretaceous, Maastrichtian) of Montana demonstrates multiple evolutionary reductions of neck length within Elasmosauridae. Journal of Vertebrate Paleontology, e1278608.
  19. Soul L. C., Benson R. B. J. 2017. Developmental mechanisms of macroevolutionary change in the tetrapod axis: a case study of Sauropterygia. Evolution, 71: 1164–1177.
  20. Soul L. C., Friedman M. 2017. Bias in phylogenetic measurements of extinction and a case study of end‐Permian tetrapods. Palaeontology, 60 (2): 169-185.


  1. Albino A., Carrillo-Briceño J. D., Neenan J. M. 2016. An enigmatic aquatic snake from the Cenomanian of Northern South America. PeerJ, 4: e2027.
  2. Beckett H. T., Friedman M. 2016. The one that got away from Smith Woodward: cranial anatomy of Micrornatus (Acanthomorpha: Scombridae) revealed using computed microtomography. Geological Society, London, Special Publications, 430: 337–353.
  3. Benson R. B., Butler R. J., Alroy J., Mannion P. D., Carrano M. T., & Lloyd G. T. 2016. Near-stasis in the long-term diversification of Mesozoic tetrapods. PLoS Biology, 14 (1): e1002359.
  4. Clarke J. T., Lloyd G. T., Friedman M. 2016. Little evidence for enhanced phenotypic evolution in early teleosts relative to their living fossil sister group. Proceedings of the National Academy of Sciences, 113 (41): 11531–11536.
  5. Close R. A., Davis B. M., Walsh S., Wolniewicz A. S., Friedman M., Benson R. B. J. 2016. A lower jaw of Palaeoxonodon from the Middle Jurassic of the Isle of Skye, Scotland, sheds new light on the diversity of British stem therians. Palaeontology, 59 (1): 155-169.
  6. Close R. A., Johanson Z., Tyler J. C., Harrington R. C., Friedman M. 2016. Mosaicism in a new Eocene pufferfish highlights rapid morphological innovation near the origin of crown tetraodontiforms. Palaeontology, 59: 499–514.
  7. Davesne D., Gallut C., Barriel V., Janvier P., Lecointre G., Otero O. 2016. The phylogenetic intrarelationships of spiny-rayed fishes (Acanthomorpha, Teleostei, Actinopterygii): fossil taxa increase the congruence of morphology with molecular data. Frontiers in Ecology and Evolution, 4: 129.
  8. Delbarre D. J., Davesne D., Friedman M. 2016. Anatomy and relationships of †Aipichthys pretiosus and †‘Aipichthysnuchalis (Acanthomorpha: Lampridomorpha), early Late Cretaceous relatives of oarfishes and their allies. Journal of Systematic Palaeontology, 14 (7): 545-567.
  9. Friedman M., Beckett HT., Close RA., Johanson Z. 2016. The English Chalk and London Clay: two remarkable British bony fish Lagerstätten. Geological Society, London, Special Publications, 430: 165–200.
  10. Friedman M., Giles S. 2016. Actinopterygians: The Ray-Finned Fishes—An Explosion of Diversity. In Evolution of the Vertebrate Ear (pp. 17-49). Springer International Publishing.
  11. Fischer V., Bardet N., Benson R. B. J., Arkhangelsky M. S., Friedman M. 2016. Extinction of fish-shaped marine reptiles associated with reduced evolutionary rates and global environmental volatility. Nature Communications, 7: 10825.
  12. Harrington R. C., Faircloth B. C., Eytan R. I., Smith W. L., Near T. J., Alfaro M. E., Friedman M. 2016. Phylogenomic analysis of carangimorph fishes reveals flatfish asymmetry arose in a blink of the evolutionary eye. BMC Evolutionary Biology, 16: 224.
  13. Lloyd G. T., Bapst D. W., Friedman M., Davis K. E. 2016. Probabilistic divergence time estimation without branch lengths: dating the origins of dinosaurs, avian flight and crown birds. Biology Letters, 12 (11): 20160609.
  14. Lu J., Giles S., Friedman M., den Blaauwen J. L., Zhu M. 2016. The oldest actinopterygian highlights the cryptic early history of the hyperdiverse ray-finned fishes. Current Biology, 26: 1602–1608.
  15. Sobral G., Reisz R., Neenan J. M., Müller J., Scheyer T. M. 2016. Basal Reptilians, Marine Diapsids, and Turtles: The Flowering of Reptile Diversity. In Evolution of the Vertebrate Ear (pp. 207-243). Springer International Publishing.