(For links to papers, see my Google Scholar page)

In press



Navalón G, Marugán‐Lobón J, Bright JA, Cooney CR & Rayfield EJ. 2020. The consequences of craniofacial integration for the adaptive radiations of Darwin’s finches and Hawaiian honeycreepers. Nature Ecology & Evolution 4 (2), 270-278.


Morales-García NM, Burgess, TD, Hill, JJ, Gill, PG & Rayfield EJ. 2019. The use of extruded finite-element models as a novel alternative to tomography-based models: a case study using early mammal jaws. Journal of the Royal Society Interface 16 (161), 20190674

Schaeffer J, Benton MJ, Rayfield EJ, Stubbs TL. 2019. Morphological disparity in theropod jaws: comparing discrete characters and geometric morphometrics. Palaeontology 

Bright JA, Marugán-Lobón J, Rayfield EJ, Cobb SN. 2019. The multifactorial nature of beak and skull shape evolution in parrots and cockatoos (Psittaciformes). BMC Evolutionary Biology 19(1): 104.

Brocklehurst R, Porro LB, Herrel A, Adriaens D, Rayfield EJ. 2019. A digital dissection of two teleost fishes: comparative functional anatomy of the cranial musculoskeletal system in pike (Esox lucius) and eel (Anguilla anguilla). Journal of Anatomy 10.1111/joa.13007.

Ballell A, Moon BC, Porro LB, Benton MJ, Rayfield EJ. 2019. Convergence and functional evolution of longirostry in crocodylomorphs. Palaeontology 10.1111/pala.12432

Adams NF, Rayfield EJ, Cox PG, Cobb SN, Corfe IJ. 2019. Functional tests of the competitive exclusion hypothesis for multituberculate extinction. Royal Society Open Science 6 (3), 181536.

Navalón G, Bright JA, Marugán‐Lobón J, Rayfield EJ. 2019. The evolutionary relationship among beak shape, mechanical advantage, and feeding ecology in modern birds. Evolution 73 (3), 422-435.

Rayfield EJ. 2019. What Does Musculoskeletal Mechanics Tell Us About Evolution of Form and Function in Vertebrates? In: Bels V., Whishaw I. (eds) Feeding in Vertebrates. Fascinating Life Sciences. Springer, Cham.

Hill JJ, Puttick MN, Stubbs TL, Rayfield EJ, Donoghue PCJ. 2019. Evolution of jaw disparity in fishes. Palaeontology 61 (6), 847-854.

Rodrigues PG, Martinelli AG, Schultz CL, Corfe IJ, Gill PG, Soares MB. & Rayfield EJ. 2018. Digital cranial endocast of Riograndia guaibensis (Late Triassic, Brazil) sheds light on the evolution of the brain in non-mammalian cynodonts. Historical Biology 31 (9), 1195-1212


Lautenschlager S, Gill P, Luo Z-X, Fagan MJ & Rayfield, EJ. 2018. The role of miniaturisation in the evolution of the mammalian jaw and middle ear. Nature 561: 533-537.

Melbourne LA, Denny MW, Harniman RL, Rayfield EJ & Schmidt DN. 2018. The importance of wave exposure on the structural integrity of rhodoliths. Journal of Experimental Marine Biology and Ecology 503: 109-111.

Roddy KA, Skinner REH, Brunt LH, Kague E, Cross S, Rayfield EJ & Hammond CL. 2017. A zebrafish model of developmental joint dysplasia: Manipulating the larval mechanical environment to drive the malformation and recovery of joint shape. bioRxiv doi:

Lautenschlager S, Gill PG, Luo Z-X, Fagan MF, Rayfield EJ.I 2017. Morphological evolution of the mammalian jaw adductor complex. 92(4): 1910-1940 Biological Reviews DOI: 10.1111/brv.12314

Button DJ, Barrett PM & Rayfield EJ. 2017. Craniodental functional evolution in sauropodomorph dinosaurs. Paleobiology 1-28.

Caromel AGM, Schmidt DN & Rayfield EJ. 2017. Ontogenetic constraints on foraminiferal test construction. Evolution and Development 19(3): 157-168.

Taylor A, Lautenschlager S, Zhao Q & Rayfield EJ 2017 Biomechanical evaluation of different musculoskeletal arrangements in Psittacosaurus and implications for cranial function. Anatomical Record 300(1): 49-61.

Madan MA, Rayfield EJ & Bright JA. 2017. Scaling and functional morphology in strigiform hind limbs. Scientific Reports 7: 44920.

Davies TG, Rahman IA, Lautenschlager S, Cunningham JA, Asher RJ, Barrett PM, Bates KT, Bengtson S, Benson RBJ, Boyer DM, Braga J, Bright JA, Claessens LPAM, Cox PG, Dong X-P, Evans AR, Falkingham PL, Friedman M, Garwood RJ, Goswami A, Hutchinson JR, Jeffery NS, Johanson Z, Lebrun R, Martínez-Pérez C, Marugán-Lobón J, O’Higgins PM, Metscher B, Orliac M, Rowe TB, Rücklin M, Sánchez-Villagra MR, Shubin NH, Smith SY, Starck JM, Stringer C, Summers AP, Sutton MD, Walsh SA, Weisbecker V, Witmer LM, Wroe S, Yin Z, Rayfield EJ, Donoghue PCJ. 2017. Open data and digital morphology. Proceedings of the Royal Society of London, B 284(1852): 20170194.


MacLaren JA, Anderson PSL, Barrett PM & Rayfield EJ. 2016. Herbivorous dinosaur jaw disparity and its relationship to extrinsic evolutionary drivers. Paleobiology 43(1): 15-33.

Button DJ, Barrett PM & Rayfield EJ. 2016. Comparative cranial myology and biomechanics of Plateosaurus and Camarasaurus and evolution of the sauropod feeding apparatus. Palaeontology 59(6): 887-913.

Vinther J, Nicholls R, Lautenschlager S, Pittman M, Kaye TG, Rayfield EJ, Mayr G & Cuthill IC. 2016. 3D camouflage in an ornithischian dinosaur. Current Biology 26(18): 2456-2462.

Brunt LH, Skinner REH, Roddy KA, Araujo NM, Rayfield EJ & CL Hammond. 2016. Differential effects of altered patterns of movement and strain on joint cell behaviour and skeletal morphogenesis. Osteoarthritis and Cartilage 24(11): 1940-1950.

Bright JA, Marugán-Lobón J, Cobb SN & Rayfield EJ. 2016. The shapes of bird beaks are highly controlled by nondietary factors. Proceedings of the National Academy of Sciences 113(19): 5352-5357.

Martinez-Perez C, Rayfield EJ, Botella H, Donoghue PCJ. 2016 Translating taxonomy into the evolution of conodont feeding ecology. Geology 44: 247-250.

Polly D, Stayton T, Dumont E, Pierce SE, Rayfield EJ, Angielczyk K 2016 Combining geometric morphometrics and finite element analysis with evolutionary modeling: towards a synthesis. Journal of Vertebrate Paleontology e1111225.

Brunt LH, Roddy KA, Rayfield EJ & CL Hammond. 2016. Building finite element models to investigate zebrafish jaw biomechanics. Journal of Visualised Experiments. 118.


Caromel, AGM, Schmidt DN, Fletcher I & Rayfield EJ. 2015 Morphological change during the ontogeny of the planktic foraminifera. Journal of Micropalaeontology 35: 2-19.

Apostolaki NE, Rayfield EJ & Barrett PM. 2015 Osteological and soft-tissue evidence for pneumatization in the cervical column of the ostrich (Struthio camelus) and observations on the vertebral columns of non-volant, semi-volant and semi-aquatic birds. PLoS ONE 10(12) e0143834.

Murdock DJE, Rayfield EJ & Donoghue PCJ. 2015 Functional adaptation underpinned the evolutionary assembly of the earliest vertebrate skeleton (vol 16, pg354, 2014). Evolution and Development 17: 172-172.

Cuff AR, Bright JA & Rayfield EJ. 2015. Validation experiments on finite element models of an ostrich (Struthio camelus) cranium. PeerJ 3, e1294.

Brunt LH, Norton JL, Bright JA, Rayfield EJ & Hammond CL. 2015. Finite Element modelling predicts changes in joint shape and cell behaviour due to loss of muscle strain in jaw development. Journal of Biomechanics 48(12): 3112-3122.

Porro LB, Rayfield EJ & Clack JA. 2015 Computed tomography, anatomical description and three-dimensional reconstruction of the lower jaw of Eusthenopteron foordi Whiteaves, 1881 from the Upper Devonian of Canada. Palaeontology 58(6): 1031-1047.

Melbourne LA, Griffin J, Schmidt DN & Rayfield EJ 2015 Potential and limitations of finite element modelling in assessing structural integrity of coralline algae under future global change. Biogeosciences 12(19): 5871-5883.

Cuff AR & Rayfield EJ 2015 Retrodeformation and muscular reconstruction of ornithomimosaurian dinosaur crania PeerJ e1093

 Porro LB, Rayfield EJ & Clack JA 2015 Descriptive anatomy and three-dimensional reconstruction of the skull of the early tetrapod Acanthostega gunnari Jarvik, 1952. PloS One 10 (3), e0118882


Button DJ, Rayfield EJ & Barrett PM 2014 Cranial biomechanics underpins high sauropod diversity in resource-poor environments. Proceedings of the Royal Society of London, B. 281: 20142114.

Murdock DJE, Rayfield EJ & Donoghue PCJ 2014 Functional adaptation underpinned the evolutionary assembly of the earliest vertebrate skeleton. Evolution and Development 16: 354-361. DOI: 10.1111/ede.12096

Neenan JM, Ruta M, Clack JA & Rayfield EJ 2014 Feeding biomechanics in Acanthostega and across the fish-tetrapod transition. Proceedings of the Royal Society B 281 1781

Lautenschlager S, Witmer LM, Altangerel P, Zanno LE & Rayfield EJ 2014 Cranial anatomy of Erlikosaurus andrewsi (Dinosauria, Therizinosauria): new insights based on digital reconstruction. Journal of Vertebrate Paleontology 34: 1263-1291. [Cover image]

Gill PG, Purnell MA, Crumpton N, Robson Brown K, Gostling NJ, Stampanoni M & Rayfield EJ. 2014 Dietary specializations and diversity in feeding ecology of the earliest stem mammals. Nature 512: 303-305. [Cover image]

Cunningham JA, Rahman IA, Lautenschlager S, Rayfield EJ & Donoghue PCJ 2014 A virtual world of paleontology. Trends in Ecology and Evolution 29(6): 347-357. [Cover image]

Foffa D, Cuff AR, Sassoon J, Rayfield EJ, Mavrogordato MN & Benton MJ 2014 Functional anatomy and feeding biomechanics of a giant Upper Jurassic pliosaur (Reptilia: Sauropterygia) from Weymouth Bay, Dorset, UK. Journal of Anatomy 225: 209-219.

Anderson PSL, Renaud, S. & Rayfield EJ 2014 Adaptive plasticity in the mouse mandible. BMC Evolutionary Biology 14(9): 85

Lautenschlager S, Bright JA & Rayfield EJ 2014 Digital dissection – using contrast enhanced CT scanning to elucidate hard and soft tissue anatomy in the common buzzard Buteo Buteo Journal of Anatomy 224: 412-431.

Caromel AGM, Schmidt DS, Phillips JC. & Rayfield EJ 2014 Hydrodynamic constraints on the evolution and ecology of planktic foraminifera. Marine Micropaleontology 106: 69-78.

Martinez-Perez C, Rayfield EJ, Purnell MA & Donoghue PCJ 2014 Finite element, occlusal, microwear and microstructural analyses indicate that conodont microstructure is adapted to dental function. Palaeontology 57: 1059-1066.


Lautenschlager S, Altangerel P, Witmer LM. & Rayfield EJ 2013 Edentulism, beaks and biomechanical innovations in the early evolution of theropod dinosaurs. Proceedings of the National Academy of Sciences 110: 20657-20662.

Stubbs TL, Pierce SE, Rayfield EJ & Anderson PSL 2013 Morphological and biomechanical disparity of crocodile-line archosaurs following the end-Triassic extinction. Proceedings of the Royal Society B 280(1770) 20131940.

Purnell MA, Crumpton N, Gill PG, Jones G & Rayfield EJ 2013 Within-guild dietary discrimination from 3-D textural analysis of tooth microwear in insectivorous mammals. Journal of Zoology 291: 249-257.

Cobley MJ, Rayfield EJ & Barrett PM. 2013 Inter-vertebral flexibility of the ostrich neck: implications for estimating sauropod neck flexibility. PLOS ONE 8(8) e72187.

Schmidt DN, Rayfield EJ, Cocking A & Marone F. 2013 Linking evolution and development: synchrotron radiation X-ray tomographic microscopy of planktic foraminifers. Palaeontology 56(4) 741-749.

Cuff AR & Rayfield EJ 2013 Feeding mechanics in spinosaurid theropods and extant crocodilians. PLOS ONE 8(5) e65295.


Jones D, Evans AR, Rayfield EJ, Sui KKW & Donoghue PCJ. 2012 Testing microstructural adaptation in the earliest dental tools. Biology Letters 8(6) 952-955.

Lautenschlager S, Rayfield EJ, Altangerel P, Zanno LE & Witmer LM. 2012. The endocranial anatomy of Therizinosauria and its implications for sensory and cognitive function. PLOS ONE 7(12) e52289

Cox PG, Rayfield EJ, Fagan MJ, Herrel A, Pataky TC & Jeffrey N. 2012 Functional evolution of the feeding system in rodents. PLoS ONE 7(4): e36299.

Birn-Jeffery AV, Miller CE, Naish D, Rayfield EJ, Hone DWE. 2012 Pedal claw curvature in birds, lizards and Mesozoic dinosaurs – complicated categories and compensating for mass-specific and phylogenetic control. PLOS ONE 7(12): e50555.

Anderson PSL & Rayfield EJ. 2012 Virtual experiments, physical validation: dental morphology at the intersection of experiment and theory. Journal of the Royal Society Interface 9(73): 1846-1855.

Young MT, Rayfield EJ, Holliday CM, Witmer LM, Button DJ, Upchurch P & Barrett PM. 2012. Cranial biomechanics of Diplodocus (Dinosauria, Sauropoda): testing hypotheses of feeding behaviour in an extinct megaherbivore. Naturwissenschaften 99: 637-643 Awarded the Arnold Berliner Award for best research article published in Naturwissenschaften in 2012.

Jones D, Evans AR, Sui KKW, Rayfield EJ & Donoghue PCJ. 2012 The sharpest tools in the box? Quantitative analysis of conodont element functional morphology. Proceedings of the Royal Society, B 279: 2849-54 DOI: 10.1098/rspb.2012.0147 [21]

Close RA & Rayfield EJ. 2012 Functional morphometric analysis of the furcula in Mesozoic birds. PLoS ONE 7(5): e36664.


Bright JA & Rayfield EJ. 2011. Sensitivity and ex vivo validation of finite element models of the domestic pig cranium. Journal of Anatomy 219: 456-471.

Anderson PSL, Bright JA, Gill PG, Palmer C & Rayfield EJ. 2012. Models in palaeontological functional analysis. Biology Letters 8: 119-122.

Anderson PSL, Friedman M, Brazeau MD & Rayfield EJ. 2011. Initial radiation of jaws demonstrated stability despite faunal and environmental change. Nature 476: 206-209.

Cox PG, Fagan MJ, Rayfield EJ & Jeffery N. 2011. Finite element modelling of squirrel, guinea pig and rat skulls: using geometric morphometrics to assess sensitivity Journal of Anatomy 219: 696-709.

Rayfield EJ. 2011. Structural performance of tetanuran theropod skulls, with emphasis on the Megalosauridae, Spinosauridae and Carcharodontosauridae. Special Papers in Palaeontology 86: 241-253.

Bright JA & Rayfield EJ. 2011. The response of cranial biomechanical finite element models to variations in mesh density. Anatomical Record 294(4): 610-620.

Rayfield EJ. 2011. Strain in the ostrich mandible during simulated pecking and validation of specimen-specific finite element models. Journal of Anatomy 218: 47-58.

Anderson PSL, Gill PG & Rayfield EJ. 2011. Modeling the effects of cingula structure on strain patterns and potential fracture in tooth enamel. Journal of Morphology 272: 50-65.


Fletcher TM, Janis CM & Rayfield EJ. 2010. Finite element analysis of ungulate jaws: can mode of digestive physiology be determined? Palaeontologica Electronica 13(3) 21A, 15p.

Jasinoski SC, Rayfield EJ & Chinsamy A. 2010. Mechanics of the scarf premaxilla-nasal suture in the snout of Lystrosaurus. Journal of Vertebrate Paleontology. 30(4): 1283-1288.

Jasinoski SC, Rayfield EJ & Chinsamy A. 2010. Functional implications of dicynodont cranial suture morphology. Journal of Morphology. 271: 705-728.


Pierce SE, Angielczyk KD & Rayfield EJ. 2009. Shape and mechanics in thalattosuchian (Crocodylomorpha) skulls: implications for feeding behavior and niche partitioning. Journal of Anatomy. 215: 555-576.

Pierce SE, Angielczyk KD & Rayfield EJ. 2009. Morphospace occupation in thalattosuchian crocodylomorphs: skull shape variation, species delineation, and temporal patterns. Palaeontology 52(5): 1057-1097.

Jasinoski SC, Rayfield EJ & Chinsamy A. 2009. Comparative feeding biomechanics of Lystrosaurus and the generalized dicynodont Oudenodon. The Anatomical Record 292: 862-874.

Rayfield EJ, Barrett PM, Milner AR. 2009. Utility and validity of Triassic ‘Land Vertebrate Faunachrons’. Journal of Vertebrate Paleontology 29(1): 80-87.

Head JJ, Barrett, PM & Rayfield EJ. 2009. Neurocranial osteology and systematic relationships of Varanus (Megalania) prisca Owen, 1859 (Squamata: Varanidae). Zoological Journal of the Linnean Society 155: 445-457.


Rayfield EJ & Milner AC 2008. Establishing a framework for archosaur cranial mechanics. Paleobiology 34: 494-515.

Pierce SE, Angielczyk KD & Rayfield EJ 2008. Patterns of morphospace occupation and mechanical performance in extant crocodilian skulls: a combined geometric morphometric and finite element modelling approach. Journal of Morphology 269: 840-864.

Moreno K, Wroe S, Clausen P, McHenry C, D’Amore DC, Rayfield EJ & Cunningham E. 2008. Cranial performance in the Komodo dragon (Varanus komodoensis) as revealed by high-resolution 3-D finite element analysis. Journal of Anatomy 212: 736-746.


Rayfield EJ, Milner AC, Bui Xuan V, Young PG. 2007. Functional morphology of spinosaur ‘crocodile-mimic’ dinosaurs. Journal of Vertebrate Paleontology 27(4): 892-901.

Rayfield EJ. 2007. Finite element analysis and understanding the biomechanics and evolution of living and fossil organisms. Annual Review of Earth and Planetary Sciences 35: 541-576.


Barrett PM & Rayfield EJ. 2006. Ecological and evolutionary implications of dinosaur feeding behaviour. Trends in Ecology and Evolution 21: 217-24.

Evans SE, Barrett PM, Hilton J, Butler R, Jones MEH, Liang MM, Parish J, Rayfield EJ, Sigogneau-Russell D and Underwood C. 2006. The Middle Jurassic vertebrate assemblage of Skye, Scotland. In Barrett PM, Evans SE, eds. Ninth international symposium on Mesozoic terrestrial ecosystems and biota. London, UK: Natural History Museum. 36–39.


Rayfield EJ. 2005. Aspects of comparative cranial mechanics in the theropod dinosaurs Coelophysis, Allosaurus and Tyrannosaurus. Zoological Journal of the Linnean Society 144, 309-316.

Rayfield EJ. 2005. Using Finite Element Analysis to investigate suture morphology: a case study using large, carnivorous dinosaurs. The Anatomical Record 283A 349-365.

Rayfield EJ, Barrett PM, McDonnell RA & Wills KJ. 2005. A geographical information system (GIS) study of Triassic vertebrate biochronology. Geological Magazine 142, 327-354.


Rayfield EJ. 2004. Cranial mechanics and feeding in Tyrannosaurus rex. Proceedings of the Royal Society of London, B. 271: 1451-1459.


Rayfield EJ, Norman DB, Upchurch P. 2002. Prey attack by a large theropod dinosaur. Nature  416: 388.

Rayfield EJ. 2002 Cranial form and function in a large theropod dinosaur: a study using Finite Element Analysis. PhD Thesis, University of Cambridge.


Rayfield EJ, Norman DB, Horner CC, Horner JR, Smith PM, Thomason JJ & Upchurch P. 2001. Cranial design and function in a large theropod dinosaur. Nature 409, 1033-1037.



Popular authored books and book contributions 

Rayfield EJ. 2015. Finite Element Analysis (Paleontology). In McGraw-Hill Yearbook of Science and Technology pp108-110 McGraw-Hill, New York

Rayfield EJ. 2007. ‘Allosaurus feeding habits’. p107. in The Random House Look-It-Up Book of Dinosaurs. Random House Books For Young Readers, Random House, Jersey City, NJ.

Rayfield EJ. 2004. Finite Element Analysis (Paleontology). In McGraw-Hill Yearbook of Science and Technology pp108-110 McGraw-Hill, New York

Rayfield EJ. 2003. Dinosaurs Connections and Axis Publishing Ltd, London.

Barrett PM. 2001. National Geographic Dinosaurs. Firecrest Books Ltd, Amersham, UK and National Geographic Society, Washington DC, USA. 192 pp. Contributing writer.