My background is in biology, paleontology, and art. I was an undergraduate at Colgate University (B.A., 1983) before a year-long Watson Fellowship studying dinosaurs in European museums. At Harvard (Ph.D. 1989) I did X-ray and muscle activity studies of alligators and birds to explore the evolution of hind limb function in bipedal, meat-eating dinosaurs (theropods). I received postdoctoral training in nerve-muscle development at Emory University, bird flight at the University of Montana, and avian embryology at Harvard before joining the faculty at Wake Forest University. Since 1995 I've been a professor at Brown, where I teach human anatomy to medical students. My research uses 3-D animation tools to reconstruct dinosaur foot movements based on fossil tracks, to measure skeletal motion in walking and flying birds, and to find new ways to study locomotor evolution. I strive to be a bridge between biologists exclusively studying living animals and paleontologists working solely with fossils.
|Hatala KG, Perry DA, Gatesy SM A biplanar X-ray approach for studying the 3D dynamics of human track formation.. Journal of Human Evolution/Journal of Human Evolution. 2018; 121 : 104-118.|
|Falkingham, Peter L., Bates, Karl T., Avanzini, Marco, Bennett, Matthew, Bordy, Emese M., Breithaupt, Brent H., Castanera, Diego, Citton, Paolo, Díaz-Martínez, Ignacio, Farlow, Jim O., Fiorillo, Anthony R., Gatesy, Stephen M., Getty, Patrick, Hatala, Kevin G., Hornung, Jahn J., Hyatt, James A., Klein, Hendrik, Lallensack, Jens N., Martin, Anthony J., Marty, Daniel, Matthews, Neffra A., Meyer, Christian A., Milàn, Jesper, Minter, Nicholas J., Razzolini, Novella L., Romilio, Anthony, Salisbury, Steven W., Sciscio, Lara, Tanaka, Ikuko, Wiseman, Ashleigh L. A., Xing, L. D., Belvedere, Matteo A standard protocol for documenting modern and fossil ichnological data. Palaeontology/Palaeontology. 2018; 61 (4) : 469-480.|
|FARLOW, JAMES O., ROBINSON, NATHAN J., TURNER, MORGAN L., BLACK, JENELL, GATESY, STEPHEN M. FOOTFALL PATTERN OF A BOTTOM-WALKING CROCODILE (CROCODYLUS ACUTUS). PALAIOS/PALAIOS. 2018; 33 (9) : 406-413.|
|Kambic RE, Roberts TJ, Gatesy SM 3-D range of motion envelopes reveal interacting degrees of freedom in avian hind limb joints. Journal of Anatomy/J. Anat.. 2017; 231 (6) : 906-920.|
|Gearing effects of the patella (knee extensor muscle sesamoid) of the helmeted guineafowl during terrestrial locomotion. 2017;|
|Gatesy, Stephen M., Falkingham, Peter L. Neither bones nor feet: track morphological variation and ‘preservation quality’. Journal of Vertebrate Paleontology/Journal of Vertebrate Paleontology. 2017; : e1314298.|
|Panagiotopoulou O, Rankin JW, Gatesy SM, Hutchinson JR A preliminary case study of the effect of shoe-wearing on the biomechanics of a horse's foot.. PeerJ. 2016; 4 : e2164.|
Gatesy, Stephen M. and Richard G. Ellis
Beyond Surfaces: A Particle-Based Perspective on Track Formation.
2016; : 82-91.
|Knörlein BJ, Baier DB, Gatesy SM, Laurence-Chasen JD, Brainerd EL Validation of XMALab software for marker-based XROMM.. The Journal of Experimental Biology. 2016; 219 (Pt 23) : jeb.145383.|
|Kambic RE, Roberts TJ, Gatesy SM Guineafowl with a twist: asymmetric limb control in steady bipedal locomotion. The Journal of Experimental Biology. 2015; 218 (Pt 23) : 3836-3844.|
|Luo, Zhe-Xi, Gatesy, Stephen M., Jenkins, Farish A., Amaral, William W., Shubin, Neil H. Mandibular and dental characteristics of Late Triassic mammaliaform Haramiyavia and their ramifications for basal mammal evolution . Proceedings of the National Academy of Sciences. 2015; : 201519387.|
|Kambic, R. E., Roberts, T. J., Gatesy, S. M. Long-axis rotation: a missing degree of freedom in avian bipedal locomotion. The Journal of Experimental Biology. 2014; 217 (15) : 2770-2782.|
|Falkingham, Peter L., Gatesy, Stephen M. The birth of a dinosaur footprint: Subsurface 3D motion reconstruction and discrete element simulation reveal track ontogeny. Proceedings of the National Academy of Sciences. 2014; : 201416252.|
|Ellis, R.G. and Gatesy, S.M. A biplanar X-ray method for three-dimensional analysis of track formation.. Palaeontologia Electronica. 2013; 16 (1) : 16.|
|Pittman, Michael, Gatesy, Stephen M., Upchurch, Paul, Goswami, Anjali, Hutchinson, John R. Shake a Tail Feather: The Evolution of the Theropod Tail into a Stiff Aerodynamic Surface. PLoS ONE. 2013; 8 (5) : e63115.|
|Baier, David B., Gatesy, Stephen M. Three-dimensional skeletal kinematics of the shoulder girdle and forelimb in walking Alligator . J. Anat.. 2013; : n/a-n/a.|
|Baier, David B., Gatesy, Stephen M., Dial, Kenneth P. Three-Dimensional, High-Resolution Skeletal Kinematics of the Avian Wing and Shoulder during Ascending Flapping Flight and Uphill Flap-Running. PLoS ONE. 2013; 8 (5) : e63982.|
|Gatesy SM, Pollard NS Apples, oranges, and angles: Comparative kinematic analysis of disparate limbs.. Journal of Theoretical Biology. 2011; 282 (1) : 7-13.|
|Gatesy SM, Baier DB, Jenkins FA, Dial KP Scientific rotoscoping: a morphology-based method of 3-D motion analysis and visualization.. Journal of experimental zoology. Part A, Ecological genetics and physiology. 2010; 313 (5) : 244-61.|
|Brainerd, Elizabeth L., Baier, David B., Gatesy, Stephen M., Hedrick, Tyson L., Metzger, Keith A., Gilbert, Susannah L., Crisco, Joseph J. X-ray reconstruction of moving morphology (XROMM): precision, accuracy and applications in comparative biomechanics research.. Journal of experimental zoology. Part A, Ecological genetics and physiology. 2010; 9999A (5) : 262-79.|
|Gatesy, Stephen M., Bäker, Martin, Hutchinson, John R. Constraint-based exclusion of limb poses for reconstructing theropod dinosaur locomotion. Journal of Vertebrate Paleontology. 2009; 29 (2) : 535-544.|
|Keefe, D. F., O'Brien, T. M., Baier, D. B., Gatesy, S. M., Brainerd, E. L., Laidlaw, D. H. Exploratory Visualization of Animal Kinematics Using Instantaneous Helical Axes. Computer Graphics Forum. 2008; 27 (3) : 863-870.|
|Jenkins, Farish A., Shubin, Neil H., Gatesy, Stephen M., Warren, Anne Gerrothorax pulcherrimus from the Upper Triassic Fleming Fjord Formation of East Greenland and a reassessment of head lifting in temnospondyl feeding . Journal of Vertebrate Paleontology. 2008; 28 (4) : 935-950.|
|Baier DB, Gatesy SM, Jenkins FA A critical ligamentous mechanism in the evolution of avian flight.. J. Geophys. Res.. 2007; 445 (7125) : 307-10.|
|Brainerd, E., Gatesy, S., Baier, D., Hedrick, T. A method for accurate 3-D reconstruction of skeletal morphology and movement. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology. 2007; 146 (4) : S119.|
Gatesy, S.M. and Middleton, K.M.
Skeletal adaptations for flight.
2007; : 269-283.
|Hutchinson, John R., Gatesy, Stephen M. Dinosaur Locomotion: Beyond the bones. J. Geophys. Res.. 2006; 440 (7082) : 292-294.|
|Gatesy, S.M., Shubin, N.H. and Jenkins, F.A., Jr. Anaglyph stereo imaging of dinosaur track morphology and microtopography.. Palaeontologia Electronica. 2005; 8 (1) : 10.|
|Gatesy, S.M. and Baier, D.B. The origin of the avian flight stroke: a kinematic and kinetic perspective.. Paleobiology. 2005; 31 (3) : 382-399.|
|Gatesy, Stephen Direct and Indirect Track Features: What Sediment Did a Dinosaur Touch?. Ichnos. 2003; 10 (2-4) : 91-98.|
Locomotor evolution on the line to modern birds..
2002; : 432-447.
|Bimber, O., Gatesy, S.M., Witmer, L.M., Raskar, R., Miguel Encarnacao, L. Merging fossil specimens with computer-generated information. Computer. 2002; 35 (9) : 25-30.|
Jenkins, F.A. Jr., Shubin, N.H., Gatesy, S.M. and Padian, K.
A diminutive pterosaur (Pterosauria: Eudimorphodontidae) from the Greenlandic Triassic..
Bulletin of the Museum of Comparative Zoology. 2001; 156 (1) : 151-170.
|Hutchinson, John R, Gatesy, Stephen M Bipedalism. Encyclopedia of Life Sciences. 2001;|
Skin impressions in Triassic theropod tracks as records of foot movement..
Bulletin of the Museum of Comparative Zoology. 2001; 156 (1) : 137-149.
The evolutionary history of the theropod caudal locomotor module..
2001; : 333-350.
|Hutchinson, J.R. and Gatesy, S.M. Adductors, abductors, and the evolution of archosaur locomotion.. Paleobiology. 2000; 26 (4) : 734-751.|
|MIDDLETON, KEVIN M., GATESY, STEPHEN M. Theropod forelimb design and evolution. Zoological Journal of the Linnean Society. 2000; 128 (2) : 149-187.|
|Farlow, J. O. Theropod Locomotion. Integrative and Comparative Biology. 2000; 40 (4) : 640-663.|
|Gatesy, S.M. Guineafowl hind limb function I: cineradiographic analysis and speed effects.. Journal of Morphology. 1999; 240 : 115-127.|
|Gatesy, S.M. Guineafowl hind limb function II: electromyographic analysis and motor pattern evolution.. Journal of Morphology. 1999; 240 : 127-142.|
|Gatesy SM Guineafowl hind limb function. I: Cineradiographic analysis and speed effects.. Journal of Morphology. 1999; 240 (2) : 115-125.|
|Gatesy SM Guineafowl hind limb function. II: Electromyographic analysis and motor pattern evolution.. Journal of Morphology. 1999; 240 (2) : 127-142.|
|Gatesy, S.M. and Middleton, K.M. Theropod hind limb disparity revisited: a response. . Journal of Vertebrate Paleontology. 1999; 19 (3) : 606.|
|SM Gatesy, KM Middleton, FA Jenkins Jr, NH Shubin Three-dimensional preservation of foot movements in Triassic theropod dinosaurs. J. Geophys. Res.. 1999; 399 (6732) : 141-144.|
|Gatesy, S.M. An electromyographic analysis of hindlimb function in Alligator during terrestrial locomotion. . Journal of Morphology. 1997; 234 (2) : 197-212.|
|Gatesy SM An electromyographic analysis of hindlimb function in Alligator during terrestrial locomotion.. Journal of Morphology. 1997; 234 (2) : 197-212.|
|Gatesy, Stephen M., Middleton, Kevin M. Bipedalism, flight, and the evolution of theropod locomotor diversity. Journal of Vertebrate Paleontology. 1997; 17 (2) : 308-329.|
|Jenkins, Farish A., Gatesy, Stephen M., Shubin, Neil H., Amaral, William W. Haramiyids and Triassic mammalian evolution. J. Geophys. Res.. 1997; 385 (6618) : 715-718.|
|Gatesy SM, Dial KP FROM FROND TO FAN: ARCHAEOPTERYX AND THE EVOLUTION OF SHORT-TAILED BIRDS.. Evolution. 1996; 50 (5) : 2037-2048.|
|Gatesy, S.M. and Dial, K.P. From frond to fan: Archaeopteryx and the evolution of short-tailed birds.. 1996; 50 (5) : 2037-2048.|
|Gatesy SM, Dial KP LOCOMOTOR MODULES AND THE EVOLUTION OF AVIAN FLIGHT.. Evolution. 1996; 50 (1) : 331-340.|
|Gatesy, S.M. and Dial, K.P. Locomotor modules and the evolution of avian flight. . Evolution. 1996; 50 (1) : 331-340.|
Functional evolution of the hind limb and tail from basal theropods to birds..
1995; : 219-234.
Jenkins, F.A. Jr., Shubin, N.H., Amaral, W.W., Gatesy, S.M., Schaff, C.R., Clemmensen, L.B., Downs, W.R., Davidson, A.R., Bonde, N. and Osbaeck, F.
Late Triassic continental vertebrates and depositional environments of the Fleming Fjord Formation, Jameson Land, East Greenland..
Meddelelser om Gronland, Geoscience. 1994; 32 : 1-25.
|Gatesy, Stephen M. Neuromuscular Diversity in Archosaur Deep Dorsal Thigh Muscles. Brain Behav Evol. 1994; 43 (1) : 1-14.|
|Gatesy SM Neuromuscular diversity in archosaur deep dorsal thigh muscles.. Brain Behav Evol. 1994; 43 (1) : 1-14.|
|Gatesy SM, English AW Evidence for compartmental identity in the development of the rat lateral gastrocnemius muscle.. Developmental dynamics : an official publication of the American Association of Anatomists. 1993; 196 (3) : 174-82.|
Gatesy, S.M. and Dial, K.P.
Tail muscle activity patterns in walking and flying pigeons (Columba livia)..
The Journal of Experimental Biology. 1993; 176 : 55-76.
|Gatesy, S. M., Biewener, A. A. Bipedal locomotion: effects of speed, size and limb posture in birds and humans. J. Zoology. 1991; 224 (1) : 127-147.|
|Gatesy, Stephen M. Hind limb movements of the American alligator ( Alligator mississippiensis ) and postural grades . J. Zoology. 1991; 224 (4) : 577-588.|
|Gatesy, S.M. Hind limb scaling in birds and other theropods: implications for terrestrial locomotion. . Journal of Morphology. 1991; 209 (1) : 83-96.|
|Gatesy, S.M. Caudofemoral musculature and the evolution of theropod locomotion. . Paleobiology. 1990; 16 (2) : 170-186.|
My research is directed at understanding the evolution of the vertebrate locomotor system and the functional consequences of changing morphology as seen in the fossil record. The majority of my work focuses on bipedal locomotion and the evolution of flight in carnivorous dinosaurs, using living birds, footprints, and fossils. Our group is pioneering morphology-based 3-D motion analysis by combining computer models and bi-planar X-ray imaging (so called X-ray Reconstruction of Moving Morphology, or XROMM).
Our lab has focused on the clade Archosauria, a group that includes birds, other dinosaurs, crocodilians, pterosaurs, and additional extinct forms. We are pursuing two main questions. First, how has bipedal locomotion, a highly conserved trait, changed over the last 200 million years of theropod (carnivorous dinosaur) evolution? Second, how was the body plan of a terrestrial theropod transformed to acquire an entirely novel capability-powered flight? Both of these questions involve gaining a better understanding of how living birds use their legs, wings, and tail.
My students and I integrate data from structural, functional, historical, and computer modeling approaches. We use techniques such as electromyography, motion analysis (based on X-ray and light video or film), and force plates to interpret the interaction of the skeletal, muscular, and nervous systems during locomotion. Three-dimensional (3-D) computer animation has become increasingly important for our research over the past ten years. Using animation software (Autodesk's Maya and its predecessor), we create articulated virtual skeletons to quantify complex movement in living archosaurs, reconstruct locomotion in extinct theropods, and explore limb movement from a more theoretical perspective.
National Science Foundation, "Footprint formation and interpreting fossil dinosaur tracks", $315,697, 03/01/15-02/28/18, PI with P. Falkingham (co-PI).
National Science Foundation, “Collaborative Research: ABI Development: Integrated X-Ray Motion Analysis Software and Video Data Management for the Comparative Biomechanics Community”, $906,192, 07/01/13-06/30/16, co-PI.
Salomon Faculty Research Award, "The Origin of Dinosaur Footprint Diversity", Brown Univeristy, $15,000, 2013.
National Science Foundation, "Kinematics and Kinetics of Long-axis Rotation in Avian Bipedal Locomotion", #IOS- 0925077, $422,979, 8/1/09-7/31/12, PI with T. Roberts (co-PI).
W.M. Keck Foundation, "A Proposal to Design and Build a Dynamic 3-D Skeletal Imaging System", $1,962,464, 2/01/07-1/31/10, co-PI.
National Science Foundation, "IDBR: Hardware and Software Development for 3D Visualization of Rapid Skeletal Motion in Vertebrate Animals", #DBI-0552051, $345,486, 6/1/06-5/31-09, co-PI.
National Science Foundation, "SGER: Scientific Rotoscoping: A Morphology-Based Method of Motion Analysis and Visualization", #IOB-0532159, $110,000, 9/1/05-8/31/07, PI.
Brown University, Research Seed Funding Award, "Development and Verification of CTX Imaging for Musculoskeletal Biomechanics Research", $100,000, 1/06-12/06, co-PI.
National Science Foundation, "Reconstructing Theropod Dinosaur Limb Movements Using 3D Computer-Animated Track Simulation", #DBI-9974424, $172,706, 9/15/99-6/30/03, PI.
National Science Foundation Dissertation Improvement Grant, "Evolution and Functional Morphology of the Theropod Foot", #IBN-0073136, $6,012, 7/1/00-1/31/03, acting PI for doctoral student Kevin Middleton.
Salomon Faculty Research Award, "Analysis of Bird Wings by 3-Dimensional Computer Animation", Brown University, $12,000, 1996.
National Science Foundation, "Avian Neuromuscular Diversity: The Evolution of Muscle Morphology, Function, and Development", #IBN-9407367, $183,419, 7/1/94-6/30/99, PI.
A sampling of our work over the past 25 years:
Falkingham, P.L. and Gatesy, S.M. 2014. The birth of a dinosaur footprint: Subsurface 3D motion reconstructionand discrete element simulation reveal track ontogeny. Proceedings of the National Academy of Sciences 111: 18279-18284.
Kambic, R.E., Roberts, T.J. and Gatesy, S.M. 2014. Long-axis rotation: a missing degree of freedom in avian bipedal locomotion. Journal of Experimental Biology 217: 2770-2782.
Baier, D.B. and Gatesy, S.M. 2013. Three-dimensional skeletal kinematics of the shoulder girdle and forelimb in walking Alligator. Journal of Anatomy 223: 462-473.
Gatesy, S.M., Baier, D.B., Jenkins, F.A. and Dial, K.P. 2010. Scientific rotoscoping: a morphology-based method of 3-D motion analysis and visualization. Journal of Experimental Zoology 313A:244–261.
Brainerd, E.L., Baier, D.B., Gatesy, S.M., Hedrick, T.L., Metzger, K.A., Gilbert, S.L. and Crisco, J.J. 2010. X-ray reconstruction of moving morphology (XROMM): precision, accuracy and applications in comparative biomechanics research. Journal of Experimental Zoology 313A:262–279.
Gatesy, S.M., Bäker, M. and Hutchinson, J.R. 2009. Constraint-based exclusion of limb poses for reconstructing theropod dinosaur locomotion. Journal of Vertebrate Paleontology 29(2):535–544.
Baier, D.B., Gatesy, S.M. and Jenkins, F.A., Jr. 2007. A critical ligamentous mechanism in the evolution of avian flight. Nature 445:307-310.
Gatesy, S.M., Middleton, K.M., Jenkins, F.A. Jr. and Shubin, N.H. 1999. Three-dimensional preservation of foot movements in Triassic theropod dinosaurs. Nature 399:141-144.
Jenkins, F.A.J., Jr., Gatesy, S.M., Shubin, N.H. and Amaral, W.W. 1997. Haramiyids and Triassic mammalian evolution. Nature 385:715-718.
Gatesy, S.M. and Dial, K.P. 1996. Locomotor modules and the evolution of avian flight. Evolution. 50(1):331-340.
Gatesy, S.M. 1990. Caudofemoral musculature and the evolution of theropod locomotion. Paleobiology 16(2):170-186.
|Brainerd, Elizabeth||Professor of Biology, Professor of Medical Science|
|Crisco, Joseph||Henry F. Lippitt Professor of Orthopedics, Professor of Engineering (Research)|
|Laidlaw, David||Professor of Computer Science|
|Roberts, Thomas||Vice Chair of the Department of Ecology and Evolutionary Biology, Professor of Ecology and Evolutionary Biology|
Member, Society of Vertebrate Paleontology.
Member, Society for Integrative and Comparative Biology
Member, International Society of Vertebrate Morphologists.
I am trained as a vertebrate morphologist and functional anatomist/paleontologist. Since 1995 I have participated in the gross anatomy course for first year medical students. I also teach Ecology and Evolutionary Biology graduate seminars and supervise undergraduate independent studies and honors theses.