Here is a list of my publications. It’s here for completeness’ sake, but you may find it more useful to view on Google Scholar.

Alternatively, Fulltext downloads are available from:
ResearchGate or, or you can just email me.


Falkingham, P. L., Marty, D., and Richter, A. Dinosaur Tracks: The next steps. Indiana University Press, Bloomington.  Link

Journal Articles:


Flannery-Sutherland J. T., Kogan I., Trubin Y. S., Falkingham P. L., Winkler A., De Sousa D. D., Krylov K. D., Pokhaznikova A. A., Derbisheva M., Kapitany T. and Dudashvili A. 2023. Dinosaur trackways from the Upper Cretaceous Nichkesai Formation near Mayluu Suu City, Southern Tien Shan Mountains, north-western Kyrgyzstan. Royal Society Open Science.10230311230311 [Link]

Hatala, K.G., Gatesy, S.M. and Falkingham, P.L., 2023. Arched footprints preserve the motions of fossil hominin feet. Nature Ecology & Evolution, pp.1-10. [Link]


Grant, B., Charles, J., Geraghty, B., Gardiner, J., D’Août, K., Falkingham, P.L. and Bates, K.T., 2022. Why does the metabolic cost of walking increase on compliant substrates? Journal of the Royal Society Interface, 19(196), p.20220483. [Link]

Lallensack, J.N., Romilio, A. and Falkingham, P.L., (2022). A machine learning approach for the discrimination of theropod and ornithischian dinosaur tracks. Journal of the Royal Society Interface19(196), p.20220588 [Link]

Lallensack, J.N., Owais, A. Falkingham, P.L., Breithaupt, B.H. and Sander, M.P. (2022) ‘How to verify fossil tracks: the first record of dinosaurs from Palestine’, Historical Biology, pp. 1–11. doi:10.1080/08912963.2022.2069020.

Turner, M.L., Falkingham, P.L. and Gatesy, S.M. (2022) What is stance phase on deformable substrates? Integrative and Comparative Biology. [Link]

Lallensack, J.N. and Falkingham, P.L., 2022. A new method to calculate limb phase from trackways reveals gaits of sauropod dinosaurs. Current Biology, 32(7), pp.1635-1640. [Link]

Falkingham, P.L., Maidment, S.C., Lallensack, J.N., Martin, J.E., Suan, G., Cherns, L., Howells, C. and Barrett, P.M., (2022). Late Triassic dinosaur tracks from Penarth, south Wales. Geological Magazine, pp.1-12. [Link]

Lallensack, J.N., Farlow, J.O. and Falkingham, P.L., 2022. A new solution to an old riddle: elongate dinosaur tracks explained as deep penetration of the foot, not plantigrade locomotion. Palaeontology65(1), p.e12584. [Link]

Peri, E., Falkingham, P.L., Collareta, A. and Bianucci, G., 2022. Biting in the Miocene seas: Estimation of the bite force of the macroraptorial sperm whale Zygophyseter varolai using finite element analysis. Historical Biology, 34(10), pp.1916-1927[Link]


Cieri, R.L., Turner, M.L., Carney, R.M., Falkingham, P.L., Kirk, A.M., Wang, T., Jensen, B., Novotny, J., Tveite, J., Gatesy, S.M., Laidlaw, D.H., Kaplan, H., Moorman, A.F.M., Howell, M., Engel, B., Cruz, C., Smith, A., Gerichs, W., Lian, Y., Schultz, J.T. and Farmer, C.G. (2021), Virtual and Augmented Reality: new tools for visualizing, analyzing, and communicating complex morphology. Journal of Morphology. 282(12):1785-1800 [Link]

Hatala K. G., Gatesy S. M. and Falkingham, P. L. (2021) Integration of biplanar X-ray, three-dimensional animation and particle simulation reveals details of human ‘track ontogeny.’ Interface Focus. 11(5) p.202000075 [Link]

Mármol-Guijarro, A., Nudds, R., Folkow, L., Sellers, W., Falkingham, P. L., and Codd, J., (2021) The Influence of snow properties on speed and gait choice in the Svalbard rock ptarmigan (Lagopus muta hyperborea), Integrative Organismal Biology, 3(1), p.obab021.[Link]

Farlow, J.O., Falkingham, P.L. and Therrien, F. (2021) Pedal Proportions of Small and Large Hadrosaurs and Other Potentially Bipedal Ornithischian Dinosaurs. Cretaceous Research 127, p.104945 [Link]

Marek, R.D., Falkingham, P.L., Benson, R.B.J., Gardiner, J.D., Maddox, T.W., and Bates, K.T., (2021) Evolutionary versatility of the avian neck. Proceedings of the Royal Society B: Biological Sciences 288, 202003150. [Link]

Tanaka, I., Markó, A., Hyodo, M. Strickson, C.E., and Falkingham, P.L. (2021). A re-analysis of Chibanian Pleistocene tracks from Vértesszólós, Hungary, employing photogrammetry and 3D analysis. Annales Societatis Geologorum Poloniae 91. DOI: 10.14241/asgp.2021.02 [Link]

Falkingham, P.L. and Rae, R., (2021). 3D morphology of nematode encapsulation in snail shells, revealed by micro-CT imaging. Scientific Reports. 11(1): 2523 [Link]


Farlow, J.O, Richards, R.L., Johnson, B.D. and Falkingham, P.L. (2020) ‘A Set-up for Rapid Screen-Washing Unconsolidated Sediments for Salvage-Collecting Large Vertebrate Fossils (Pipe Creek Sinkhole, Late Neogene, Grant County, Indiana, USA)’. Proceedings of the Indiana Academy of Science 129(2): 88–96. [Link]

Milàn, J., Falkingham, P.L., Mueller-Töwe, I.J., (2020). Small ornithopod dinosaur tracks and crocodilian remains from the Middle Jurassic Bagå Formation, Born-holm, Denmark: Important additions to the rare Middle Jurassic vertebrate faunas of Northern Europe. Bulletin of the Geological Society of Denmark, 68: 245–253. Link.

Gatesy, S. M., and Falkingham, P. L., (2020). Hitchcock’s Leptodactyli, penetrative tracks, and dinosaur footprint diversity. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2020.1781142. Link.

Larkin, N. R., Duffin, C. J., Dey, S., Stukins, S., and Falkingham, P. L. (2020). The first tetrapod track recorded from the Rhaetian in the British Isles. Proceedings of the Geologists’ Association. DOI: 10.1016/j.pgeola.2020.07.012. Link.

Falkingham, P. L., Turner, M. L. and Gatesy, S. M. (2020). Constructing and testing hypotheses of dinosaur foot motions from fossil tracks, using digitization and simulation., Palaeontology, pp. 1–16. DOI: 10.1111/pala.12502. Link.

Falkingham, P. L. and Gatesy, S. M. (2020). Discussion: Defining the morphological quality of fossil footprints. Problems and principles of preservation in tetrapod ichnology with examples from the Palaeozoic to the present by Lorenzo Marchetti et al., Earth-Science Reviews, p. 103320. DOI: 10.1016/j.earscirev.2020.103320. Link.

Turner, M. L., Falkingham, P. L., and Gatesy, S. M. (2020). It’s in the loop: shared sub-surface foot kinematics in birds and other dinosaurs shed light on a new dimension of fossil track diversity. Biology Letters 16(7). DOI: 10.1098/rsbl.2020.0309. Link.

Otero, A. Pérez Moreno, A., Falkingham, P. L., Cassini, G., Ruella, A., Militello, M., and Toledo, N. (2020). Three-dimensional image surface acquisition in vertebrate paleontology: a review of principal techniques. Publicación Electrónica de la Asociación Paleontológica Argentina 20 (1): 1–14 DOI: 10.5710/PEAPA.04.04.2020.310 Link

Strickson, E. C., Hutchinson, J. R., Wilkinson, D. M. and Falkingham, P. L. (2020). Can skeletal surface area predict in vivo foot surface area? Journal of Anatomy. 236:72-84 DOI: 10.1111/joa.13090. Link

Farlow, J. O., Bakker, R. T., Dattilo, B. F. , Deschner, E. , Falkingham, P. L. Harter, C., Solis, R., Temple, D. and Ward, W. (2020). Thunder lizard handstands: Manus-only sauropod trackways from the Glen Rose Formation (Lower Cretaceous, Kendall County, Texas). Ichnos 27:2, 167-199. Link


Falkingham, P.L. and Gatesy, S. M. (2019). Track formation mechanisms elucidated by computer simulation and bi-planar X-ray. 3rd International Conference of Continental Ichnology, abstract Volume & field trip guide p24-29. Link

Troelsen, P. V., Wilkinson, D. M., Seddighi, M., Allanson, D. R. and Falkingham, P.L. (2019). Functional morphology and hydrodynamics of plesiosaur necks: Does size matter? Journal of Vertebrate Paleontology e1594850 Link

Crandell, K. E., Howe, R. O., Falkingham, P. L. (2019). Repeated evolution of drag reduction at the air–water interface in diving kingfishers. Journal of The Royal Society Interface. 16(154):20190125 DOI: 10.1098/rsif.2019.0125 Link

Novotny, J., Tveite, J., Turner, M. L., Gatesy, S., Drury, F. Falkingham, P. L., Laidlaw, D H. (2019). Developing Virtual Reality Visualizations for Unsteady Flow Analysis of Dinosaur Track Formation using Scientific Sketching. IEEE Transactions on Visualization and Computer Graphics. 25(5):2145 – 2154. DOI: 10.1109/TVCG.2019.2898796 Link


Bates, K. T. and Falkingham, P. L. (2018). The importance of muscle architecture in biomechanical reconstructions of extinct animals: a case study using Tyrannosaurus rex. Journal of Anatomy. 233: 625-635 DOI: 10.1111/joa.12874 Link

Falkingham, P. L., K. T. Bates, M. Avanzini, M. Bennett, E. M. Bordy, B. H. Breithaupt, D. Castanera, P. Citton, I. Díaz-Martínez, J. O. Farlow, A. R. Fiorillo, S. M. Gatesy, P. Getty, K. G. Hatala, J. J. Hornung, J. A. Hyatt, H. Klein, J. N. Lallensack, A. J. Martin, D. Marty, N. A. Matthews, C. A. Meyer, J. Milàn, N. J. Minter, N. L. Razzolini, A. Romilio, S. W. Salisbury, L. Sciscio, I. Tanaka, A. L. A. Wiseman, L. D. Xing, and M. Belvedere. (2018). A standard protocol for documenting modern and fossil ichnological data. Palaeontology 61:469-480. Link.

Farlow J.O., Robinson, N.J., Kumagai, C.J., Paladino, F.V., Falkingham, P.L., Elsey, R.M. and Martin, A.J. (2018). Trackways of the American Crocodile (Crocodylus acutus) in Northwestern Costa Rica: Implications for Crocodylian Ichnology. Ichnos. 25(1):30-65 Link.


Gatesy S.M. and Falkingham, P.L. (2017). Neither Bones nor Beet: Track Morphological Variation and ‘Preservation Quality’. Journal of Vertebrate Paleontology: 37(3): e1314298. Link.

Lomax D. R., Falkingham P. L., Schweigert G., Jiménez A. P. (2017). An 8.5 m long ammonite drag mark from the Upper Jurassic Solnhofen Lithographic Limestones, Germany. PLoS ONE 12(5): e0175426. Link.

Davies, T. G., I. A. Rahman, S. Lautenschlager, J. A. Cunningham, R. J. Asher, P. M. Barrett, K. T. Bates, S. Bengtson, R. B. J. Benson, D. M. Boyer, J. Braga, J. A. Bright, L. P. A. M. Claessens, P. G. Cox, X.-P. Dong, A. R. Evans, P. L. Falkingham, M. Friedman, R. J. Garwood, A. Goswami, J. R. Hutchinson, N. S. Jeffery, Z. Johanson, R. Lebrun, C. Martínez-Pérez, J. Marugán-Lobón, P. M. O’Higgins, B. Metscher, M. Orliac, T. B. Rowe, M. Rücklin, M. R. Sánchez-Villagra, N. H. Shubin, S. Y. Smith, J. M. Starck, C. Stringer, A. P. Summers, M. D. Sutton, S. A. Walsh, V. Weisbecker, L. M. Witmer, S. Wroe, Z. Yin, E. J. Rayfield and P. C. J. Donoghue. (2017) Open data and digital morphology. Proceedings of the Royal Society B: Biological Sciences 284(1852): 20170194. Link


Xing, L., Lockley, M. G., Klein, H., Falkingham, P. L., Kim, J. Y. U. L., McCrea, R. T., Zhang, J., Iv, W. S. P., Wang, T. A. O., and Wang, Z. (2016). “First early Jurassic small ornithischian tracks from Yunnan Province , Southwestern China.” PALAIOS 31(11): 516-524. Link.

Falkingham, P. L. and Horner, A. M. (2016). Trackways Produced by Lungfish During Terrestrial Locomotion. Nature Scientific Reports 6:33734. Link

Klein, H. Wizevich, M.C., Thuring, B., Marty, D., Thuring, S., Falkingham P. L., and Meyer, C. A. (2016). Triassic chirotheriid footprints from the Swiss Alps: ichnotaxonomy and depositional environment (Cantons Wallis & Glarus). Swiss Journal of Palaeontology 135(2): 295-314 Link

Bates, K. T. Mannion, P. D., Falkingham, P. L., Brusatte, S. L., Hutchinson, J. R., Otero, A., Sellers, W. I., Sullivan, C., Stevens, K., Allen, V. (2016). Temporal and phylogenetic evolution of the sauropod dinosaur body plan. Royal Society Open Science 3:150636 Link

Xing, L., Li, D., Falkingham, P. L., Lockley, M. G., Benton, M. J., Klein, H., Zhang, J., Ran, H., Persons, S, Dai, H. (2016). Digit-only sauropod pes trackways from China – evidence of swimming or a preservational phenomenon? Nature Scientific Reports 6: 21138. Link

Basu, C., Falkingham, P. L., and Hutchinson, J. R. (2016). The extinct, giant giraffid Sivatherium giganteum: skeletal reconstruction and body mass estimation. Biology letters 12: 20150940. Link


Rahman, I.A. Zamora, S., Falkingham, P.L., and Phillips, J.C. (2015). Cambrian cinctan echinoderms shed light on feeding in the ancestral deuterostome. Proceedings of the Royal Society B 282:20151964 Link

Bates, K.T., Falkingham, P.L., Macauley, S., Brassey, C., and Maidment, S.C.R. (2015). Downsizing a giant: Re-evaluating Dreadnoughtus body mass. Biology Letters (11):20150215


Falkingham, P.L., and Gatesy, S.M. (2014). The birth of a dinosaur footprint: Subsurface 3D motion reconstruction and discrete element simulation reveal track ontogeny. PNAS 111(51): 18279-18284. Link

Falkingham, P.L. Hage, J., Bäker, M. (2014). Mitigating the Goldilocks effect: the effects of different substrate models on track formation potential. Royal Society Open Science 1(3):140225 Link

Bennett, M.R., Morse, S.A., Falkingham, P.L. (2014). Tracks made by swimming Hippopotami: An example from Koobi Fora (Turkana Basin, Kenya). Palaeogeography, Palaeoclimatology, Palaeoecology 409: 9-23. Link.

Falkingham, P.L., Bates, K.T., Farlow, J.O. (2014). Historical Photogrammetry: Bird’s Paluxy River Dinosaur Chase Sequence Digitally Reconstructed as It Was prior to Excavation 70 Years Ago. PLoS ONE 9: e93247. Link.

Falkingham, P. L. (2014). Interpreting ecology and behaviour from the vertebrate fossil track record. Journal of Zoology: 222-228. Link.

Razzolini, N.L., Vila, B., Castanera, D., Falkingham, P. L., Barco JL, et al. (2014). Intra-Trackway Morphological Variations Due to Substrate Consistency: The El Frontal Dinosaur Tracksite (Lower Cretaceous, Spain). PLoS ONE 9: e93708. Link.


Maidment, S.R.C., Bates, K.T, Falkingham, P.L., van Buren, C., Arbour, V., and Barrett, P.M. (2013). Locomotion in Ornithischian Dinosaurs: An Assessment Using Three-Dimensional Computational Modelling. Biological Reviews. 89(3): 588-617. Link.

Falkingham, P L. (2013). Low cost 3D scanning using off-the-shelf video gaming peripherals. Journal of Paleontological Techniques 11: 1-9. Link.

Bates, K. T., Savage, R., Pataky, T. C., Morse, S. A., Webster, E., and Falkingham, P. L., et al. (2013). Does footprint depth correlate with foot motion and pressure? Journal of the Royal Society Interface 10(83): 20130009. Link.

Bennett, M.R., Morse, S.A., Falkingham, P.L., Bates, K.T., and Crompton, R. H. (2013). Preserving the impossible: conservation of soft-sediment hominin footprint sites and strategies for three-dimensional digital data capture. PLoS ONE 8:e60755. Link.

White, M. A., Falkingham, P. L., Cook, A. G., Hocknull, S. A., and Elliot, D. A. (2013). Morphological comparisons of metacarpal I for Australovenator wintonensis and Rapator ornitholestoides and implications for their relationship. Alcheringa. 37(4): 435-441. Link.

Castanera D, Vila B, Razzolini N. L., Falkingham P.L., Canudo J. I., et al. (2013). Manus Track Preservation Bias as a Key Factor for Assessing Trackmaker Identity and Quadrupedalism in Basal Ornithopods. PLoS ONE 8: e54177. Link.


Falkingham P. L., Bates K. T., Mannion P. D. (2012). Temporal and palaeoenvironmental distribution of manus- and pes-dominated sauropod trackways. Journal of the Geological Society 169: 365-370. Link.

Farlow J. O., O’Brien M., Kuban G. J., Dattilo B. F., Bates K. T., Falkingham, P.L. et al. (2012). Dinosaur Tracksites of the Paluxy River Valley ( Glen Rose Formation , Lower Cretaceous ), Dinosaur Valley State Park , Somervell County , Texas. Proceedings of the V International Symposium about Dinosaur Palaeontology and their Environment. pp. 41-69. Link

Sellers W. I., Hepworth-Bell J., Falkingham P. L., Bates K. T., Brassey C. A., et al. (2012). Minimum convex hull mass estimations of complete mounted skeletons. Journal of the Royal Society: Biology Letters, 8(5):842-845. Link.

Bates K. T., and Falkingham P. L. (2012). Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics. Journal of the Royal Society: Biology Letters, 8(4):660-664. Link.

Falkingham, P. L. (2012). Acquisition of high resolution 3D models using free, open source, photogrammetry. Palaeontologia Electronica. 15. 1T:15. Link.

Bates, K.T., Benson, R.B.J., and Falkingham, P. L. (2012). The evolution of body size, stance and gait in the Allosauroidea (Dinosauria: Theropoda). Palaeobiology 38: 486-507.


Falkingham, P. L., Bates, K. T., Margetts, L., and Manning, P. L. (2011). The Goldilocks effect: Preservational bias in dinosaur track assemblages. Journal of the Royal Society: Interface. 8:1142-1154. DOI: 10.1098/rsif.2010.0634. Link.

Falkingham, P. L. ,Bates, K. T., Margetts, L., Manning, P. L. (2011). Simulating sauropod manus only trackway formation using finite element analysis. Journal of the Royal Society: Biology Letters, 7(1): 142-145. DOI: 10.1098/rsbl.2010.040 Link


Bates, K. T., Falkingham, P. L., Rarity, F., Hodgetts, D., Purslow, A. and Manning, P. L. (2010). Application of high-resolution laser scanning and photogrammetric techniques to data acquisition, analysis and interpretation in palaeontology. International Archives of the Photogrammetry, Remote Sensing, and Spatial Information Sciences, XXXVIII, Part 5: 68-73

Falkingham, P. L., Milan, J., and Manning, P. L. (2010). A re-description of a crocodile trace previously interpreted as a dinosaur tail drag mark from Wyoming, U.S.A. New Mexico Museum of Natural History Bulletin 51: 171-174

Falkingham, P. L., Margetts, L., and Manning, P. L. (2010). Fossil Vertebrate Tracks as Palaeopenetrometers: confounding effects of foot morphology. PALAIOS, 25(6): 356-360

Falkingham, P. L., Thompson, K., Agenbroad, L. D. and Manning, P. L. (2010). Bird tracks at the Mammoth Site of Hot Springs, S.D. Ichnos 17(1): 34-39


Bates, K. T., Breithaupt, B. H., Falkingham, P. L., Matthews, N., Hodgetts, D., Rarity, F., and Manning, P. L. (2009). Integrated Lidar & Photogrammetric documentation of the Red Gulch Dinosaur Tracksite (Wyoming, U.S.A.). In:Foss, S.E., Cavin, J.L., Brown, T., Kirkland, J.I., and Santucci, V.L., eds., Proceedings of the Eighth Conference on Fossil Resources: Utah, p. 101-103.

Bates, K. T., Falkingham, P. L., Breithaupt, B. H., Hodgetts, D., Sellers, W. I., Manning, P. L., (2009). How Big was Big Al? Quantifying the effect of soft tissue and osteological unknowns on mass predictions of Allosaurus (Dinosauria, Theropoda) Palaeontologia Electronica 12: 33.

Manning, P. L., Margetts, L., Johnson, M. R., Withers, P., Codd, J., Sellers, W. I., Falkingham, P. L., Mummery, P. M., and Barrett, P. M. (2009). Biomechanics of Dromaeosaurid Dinosaur Claws: Application of X-Ray Microtomography, Nano-Indentation and Finite Element Analysis. The Anatomical Record, 292: 1397-1405.

Falkingham, P. L., Margetts, L., Smith, I. M., and Manning, P. L. (2009). A Reinterpretation of Palmate and Semi-Palmate (Webbed) Fossil Tracks; Insights from Finite Element Modelling. Palaeogeography, Palaeoclimatology, Palaeoecology 271(1-2): 69-76.


Manning, P.L., Ott, C. and Falkingham, P. L. (2008). A Probable Tyrannosaurid track from the Hell Creek Formation (Upper Cretaceous), Montana, United States. PALAIOS 23: 64-647.

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