Welcome to the Aeromechanics & Evolutionary Morphology Lab

ecology and evolutionary biology, brown university


About the Lab

At the broadest level, our research program seeks insight into the patterned interrelationship of structure and function in the biological world. We hope to better understand the evolution of animal architecture and biological materials, and how they are influenced by the physical world.

A primary goal of our research is to explore the mechanistic basis of flight in bats. This will not only teach us how bats fly, but also shed light on the origins and diversification of flight within the bat lineage. It helps us interpret the similarities and differences in the evolution of flight among bats, birds, pterosaurs, and insects, and elucidates facets of the ecology and physiology of flying animals. Flying animals share characteristics dictated by the constraints of physics, and our research program seeks to delineate the aspects of the structure, physiology, and mechanics of bats and their wings that distinctively shape bat flight.

Lab News


What We Do

Animals move through their environments in myriad ways, and locomotion influences all aspects of natural history. These patterns of movement depend on intricate systems of organs and tissues that are amenable to structural and mechanical analyses. The study of locomotor systems can provide an ideal context in which to explore structure/function relationships and the evolution of morphological design. In our research into the locomotion of bats, we have three central foci of study.

Structure & Motion of Bat Wings

Biologists have long viewed the flapping wings of flying vertebrates as analogous to the stationary, rigid airfoils of fixed-wing aircraft. But small, slowly moving flying animals experience viscosity effects far greater than even the smallest of aircraft. At this scale, flow over foils becomes turbulent, unsteady, and unpredictable. Basic parameters such as wing aspect ratio, angle of attack, and camber can influence flow patterns and aerodynamic forces in dramatically different ways than in faster flows.

Our lab integrates biological and physical studies of natural and naturalistic flight in living bats, studies of robotic bat wings and simpler physical models, and computational simulations. The methods we have developed to document complex wing motions have helped us show that dynamically changing 3D wing topology is the rule, even in simple straight-line flight. Even more dynamic wing motions are employed in turning and other maneuvers. Our investigations of the material properties of bat wing tissues show that bat wing bones vary greatly in mineral content, so range from highly mineralized and very stiff near the body to nearly cartilaginous and highly compliant at the wing tips. Bat wing skin is also unique, balancing the extreme mechanical demands of flight with the energetic benefits of reducing weight. We have found that the gross architecture of the wing skin’s collagen-elastin network allows a single wing to encompass an extraordinary range of mechanical characteristics.

Experimental Fluid Dynamics

Adapting techniques from experimental fluid dynamics, we can study wakes made by bats to better understand how these animals produce the forces employed in their distinctive flight. We carry out wind tunnel studies of bat wakes, coupled with detailed kinematics at high temporal and spatial resolution. We have found that the wing movements employed by bats generate characteristic wakes that have similarities with and differences from those of birds and insects. Wake structure can also differ almost as much among bat species as between a bird and a bat of comparable mass.

Our physical modeling experiments capture important aspects of the bat flight apparatus in simplified, abstracted form. Unlike the stiff wings of birds and insects, bats and gliding mammals employ airfoils made of stretchy or compliant material. Our pioneering work in compliant airfoils has demonstrated their remarkable capacity to generate lift at zero and very high angles of attack. We have found that the physical basis for this phenomenon lies in part in the self-cambering ability of compliant airfoils, which facilitates persistence of attached flow in conditions that would cause rigid airfoils to stall.

Our most sophisticated physical models are bat-like robots that capture many aspects of realistic bat flight with high fidelity, but allow us to independently modulate characteristics of the wingbeat in a manner that is impossible in living animals. These experiments help us study force production and flow dynamics, and give us controlled conditions under which we can tease apart the effect of motion and materials on aerodynamics and energetics.


A few years ago, my colleague and friend Tom Kunz paid a visit to me from Boston University. He sat in my office, looked directly at me and announced, grinning, “There are fields called terrestrial ecology and marine ecology. It’s time to have Aeroecology.” Tom challenged me to help him define what the recognition of such a discipline might mean to those who study animal flight. The physical environment of the aerosphere is both complex and dynamic, and poses many challenges to the locomotor systems of flying animals. For example, airflows are altered and modulated by motion over and around natural and human-engineered structures, and turbulence is introduced by technologies such as aircraft and wind farms. An aeroecological approach can help better understand mechanics, energetics, sensing of aerial flows, and motor control of flight. From this perspective, we can begin to analyze group flight behaviors of bats, as when immense colonies exit caves for evening foraging. In the long run, we aim to link study flight behavior in nature to the more carefully controlled studies of flight mechanics and energetics we carry out in the lab.

Research in the Aeromechanics and Evolutionary Morphology Lab is supported by the Air Force Office of Scientific Research, the National Science Foundation, and Brown University

Some material on this website supported by NSF IOS-1145549 to SMS. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.


Bat Team

Researchers past and present:

  • Sharon Swartz
    Principal Investigator

  • Erika Tavares
    Lab Manager

    Erika joined the lab in 2009. She enjoys helping with the various research projects as well as managing the everyday operation of the lab.

  • David Boerma
    Graduate Student

    The natural environments in which animals live can be complex and disruptive, and I am fascinated by how locomotor systems function in this context. My research explores how morphology, ecology, and evolutionary history influence control over body orientation in flying bats, especially during acrobatic landing maneuvers and recovery from aerial stumbles.

  • Jeremy Rehm
    Graduate Student

    My research interests fundamentally focus on mechanisms of membrane wing shape control in bats, either by active or passive mechanisms (e.g. muscular action, aerodynamic forces, and/or membrane architecture). How do these structures function together in order to produce the controlled flight we see in bats? And how could this have evolved from a gliding ancestor? Currently my research has focused primarily on the effect of intramembranous muscles on the overall shape of the wing during flight and their subsequent effects on aerodynamic forces.

  • Andrea Rummel
    Graduate Student

    Bats find themselves in a unique thermal environment during flight. Since they are primarily active at night, their wings are subjected to substantial radiative and conductive heat loss, even in warm ambient conditions, but their wing muscles are so poorly insulated as to be clearly visible through the skin. I’m interested in how bats maintain flight performance in the cold from a biomechanical and physiological perspective, and more broadly how muscles of the wing are used during flight.

  • Avi Subramanian

    I am a junior concentrating in Geology-Biology, with a focus on vertebrate paleontology, especially pertaining to archosaurs. I am currently researching the structural and functional aspects of the muscles and collagen-elastin fiber network in bat wing membrane skin, using phylogenetic and computational techniques to look for evolutionary patterns. Through this and other projects I encounter in the lab, I hope to develop skills and techniques in the biological sciences that I can take with me into graduate school and beyond.

  • Molly Magid
    Undergraduate, I-Team

    I am a sophomore concentrating in Biology and Science & Society. I am particularly interested in the evolution of morphology and hown the physiology and behavior of living systems interact with one another. I also enjoy finding ways to communicate science to the public in a clear, novel, and engaging way.

  • Diana Perkins
    Undergraduate, I-Team

    Diana is a Mechanical Engineer who is particularly excited about robotics. She is interested studying the ways that the specifics of bat flight can be applied to the field of biorobotics moving forward.

  • Lawrence Wang

Collaborating Lab

Much of our research is conducted in close collaboration with Kenny Breuer, professor of Engineering at Brown.

Breuer Lab members:

  • Kenny Breuer
    Principal Investigator

    Kenny Breuer is professor of Engineering at Brown, and interested in all things fluid mechanics including animal flight, bacterial motility, fluid-structure interactions, micron-scale fluid mechanics and energy harvesting from fluid flows.
    Breuer Lab Website

  • Hamid Vejdani
    Postdoctoral Research Associate

    Hamid is interested in understanding the dynamics of bat flight from the perspective of robotics. He earned his PhD in Robotics & Mechanical Engineering from Oregon State University. In his research he analyzes the data from bats in the framework of mathematical models to figure out how bats (as dynamical systems) achieve the magnificent maneuverability and agility that are unique to them.

  • Jillian Kiser
    Graduate Student

    Jillian works on the unsteady aerodynamics of active membrane wings. By developing an actuated membrane wing, which can change camber as a function of applied voltage, we hope to mimic the effects of the network of muscles embedded in a bat's wing membrane.

  • Gali Alon
    Graduate Student

    Gali is interested in understanding and modeling the fluid-structure interactions associated with bat-inspired, flapping membrane wings. Using simplified theoretical models, she seeks to identify the key parameters controlling the aerodynamic performance of such wings, with applications to flapping-wing micro air vehicles.

  • Matteo DiLuca
    Graduate Student

    Bats are extremely skilled flyers and are capable of robust flights through turbulent atmospheric conditions. My research combines information from predictive and inertial sensors to explore how bats are able to quickly detect and respond to flow disturbances.

  • Katie Wu

    I have spent the past summer improving a design for a biologically-inspired robotic bat wing with three degrees of freedom. In the future, I hope to increase that to four degrees of freedom, as well as use it to measure the lift and drag forces generated by different flapping patterns.

Lab Alums

Former lab members doing cool things.

what we're reading

We're a Literary Bunch


Works Published:

Journal Articles

*undergraduate co-authors
  • McCracken, G. F., Safi, K., Kunz, T. H., Dechmann, D. K. N., Swartz S. M., Wikelski, M. 2016. Airplane tracking documents the fastest flight speeds recorded for bats. Royal Society Open Science. doi:10.1098/rsos.160398
  • Hubel, T. Y., N. I. Hristov, S. M. Swartz, & K. S. Breuer. 2016. Wake structure and kinematics in two insectivorous bats. Philosophical Transactions of the Royal Society B 371:20150385. doi:10.1098/rstb.2015.0385
  • Bahlman, J. W., R. M. Price-Waldman*, H. W. Lippe*, K. S. Breuer, and S. M. Swartz. 2016. Simplifying a wing: diversity and functional consequences of digital joint reduction in bat wings. Journal of Anatomy 229 (1):114-127. doi:10.1111/joa.12457
  • Swartz, S. M. 2015. Advances in animal flight studies: Introduction. Canadian Journal of Zoology 93(12):v-vi. doi: 10.1139/cjz-2015-0243
  • Swartz, S. M. and N. Konow. 2015. Advances in the study of bat flight: the wing and the wind. Canadian Journal of Zoology 93(12):977-990. doi:10.1139/cjz-2015-0117
  • Bergou, A. J., S. M. Swartz, H. Vejdani, D. K. Riskin, L. Reimnitz, G. Taubin, K. S. Breuer. 2015. Falling with Style: Bats Perform Complex Aerial Rotations by Adjusting Wing Inertia. PLoS Biology 13(11):e1002297. doi:10.1371/journal.pbio.1002297
  • Sample, C. S., A. K. Xu, S. M. Swartz, and L. J. Gibson. 2015. Nanomechanical properties of wing membrane layers in the house cricket (Acheta domesticus Linnaeus). Journal of Insect Physiology 74:10-15. doi:10.1016/j.jinsphys.2015.01.013
  • Cheney, J. A., N. Konow, A. Bearnot, & S. M. Swartz. 2015. A wrinkle in flight: the role of elastin fibres in the mechanical behaviour of bat wing membranes. Journal of the Royal Society Interface 12:20141286. doi:10.1098/rsif.2014.1286
  • Konow, N., J. A. Cheney, T. J. Roberts, J. R. S. Waldman, S. M. Swartz. 2015. Spring or string: does tendon elastic action influence wing muscle mechanics in bat flight? Proceedings of the Royal Society B 282:20151832. doi:10.1098/rspb.2015.1832
  • Skulborstad, A. J., S. M. Swartz, and N. C. Goulbourne. 2015. Biaxial mechanical characterization of bat wing skin. Bioinspiration and Biomimetics 10:036004. doi:10.1088/1748-3190/10/3/036004
  • Long, J. H., Jr., S. A. Combes, J. Nawroth, M. Hale, G. V. Lauder, S. M. Swartz, R. D. Quinn, and H. Chiel. 2014. How does soft robotics drive research in animal locomotion? Soft Robotics, 1:161-168. doi:10.1089/soro.2014.1502
  • Chen, P., S. Joshi, S. M. Swartz, K. S. Breuer, J. W. Bahlman, and G. W. Reich. 2014. Bat-inspired flapping flight. AIAA/ASME/AHS Adaptive Structures 2014:1120,1-18. doi:10.2514/6.2014-1120
  • Cheney, J. A., N. Konow, K. M. Middleton, K. S. Breuer, T. J. Roberts, E. L. Giblin, and S. M. Swartz. 2014. Membrane muscle function in the compliant wings of bats. Bioinspiration and Biomimetics 9:025008. doi:10.1088/1748-3182/9/2/025008
  • Bahlman, J. W., S. M. Swartz, and K. S. Breuer. 2014. How wing kinematics affect power requirements and aerodynamic force production in a robotic bat wing. Bioinspiration and Biomimetics 9:025007. doi:10.1088/1748-3182/9/2/025007
  • Cheney, J. A., D. Ton*, N. Konow, D. K. Riskin, K. S. Breuer, and S. M. Swartz. 2014. Hindlimb motion during steady flight of the lesser dog-faced fruit bat, Cynopterus brachyotis. PLOS One 9:e98093. doi:10.1371/journal.pone.0098093
  • Von Busse, R., R. M. Waldman, S. M. Swartz, C. C. Voigt, and K. S. Breuer. 2014. The aerodynamic cost of flight in bats comparing theory with measurement. Journal of the Royal Society Interface 11:2014147. doi: 10.1098/​rsif.2014.0147
  • Skulborstad, A., Y. Wang, J. Davidson, S. M. Swartz, and N. C. Goulbourne. 2013. Polarized image correlation for large deformation fiber kinematics. Experimental Mechanics. 53:1-9. doi: 10.1007/s11340-013-9751-4
  • Harper, C. J., S. M. Swartz and E. L. Brainerd. 2013. Specialized bat tongue is a hemodynamic nectar mop. Proceedings of the National Academy of Sciences. 110:8852-8857. doi/10.1073/pnas.1222726110
  • von Busse, R., S. M. Swartz, and C. C. Voigt. 2013. Flight metabolism in relation to speed in Chiroptera: Testing the U-shape paradigm in the short-tailed fruit bat, Carollia perspicillata. Journal of Experimental Biology. 216:2073-2080. doi:10.1242/jeb.081760
  • Bahlman, J. W., D. K. Riskin, K. S. Breuer, and S. M. Swartz. 2013. Glide performance and aerodynamics of non-equilibrium glides in northern flying squirrels (Glaucomys sabrinus). Journal of the Royal Society Interface 10:20120794. doi.org/10.1098/rsif.2012.0794
  • Curet, O.M., S. M. Swartz, and K. S. Breuer. 2013. An aeroelastic instability provides a possible basis for the transition from gliding to flapping flight. Journal of the Royal Society Interface 10:20120940. doi.org/10.1098/rsif.2012.0940
  • Bahlman, J. W., S. M. Swartz, and K. S. Breuer. 2013. Design and characterization of a multi-articulated robotic bat wing. Bioinspiration and Biomimetics. 8: 016009. doi:10.1088/1748-3182/8/1/016009
  • Wang, Y., Son, S., Swartz, S. M., Goulbourne, N. C. 2012. A mixed Von Mises distribution for modeling soft biological tissues with two distributed fiber properties. International Journal of Solids and Structures 49: 2914-2923. 10.1016/j.ijsolstr.2012.04.004
  • Iriarte-Díaz, J., Riskin, D. K., Breuer, K. S. and Swartz, S. M. 2012. Kinematic plasticity during flight in fruit bats: individual variability in response to loading. PLoS One 7:e36665. doi:10.1371/journal.pone.0036665
  • Hubel, T. Y., N. I. Hristov, S. M. Swartz, and K. S. Breuer. 2012. Changes in kinematics and aerodynamics over a range of speeds in Tadarida brasiliensis, the Brazilian free-tailed bat. Journal of the Royal Society Interface 9:1120-1130. doi:10.1098/rsif.2011.0838
  • Riskin, D. K., A. J. Bergou, K. S. Breuer, and S. M. Swartz. 2012. Upstroke wing flexion and the inertial cost of bat flight. Proceedings of the Royal Society B – Biological Sciences 279:2945-2950. doi: 0.1098/rspb.2012.0346
  • Bergou, A. J., S. M. Swartz, K. S. Breuer, and G. Taubin. 2011. 3D Reconstruction of bat flight kinematics from sparse multiple views. IEEE International Conference on Computer Vision Theory. doi:10.1109/ICCVW.2011.6130443
  • Willis, D. J., J. W. Bahlman, K. S. Breuer, and S. M. Swartz. 2011. Energetically optimal short range gliding trajectories for gliding animals. AIAA Journal 49:2650-2657
  • Iriarte-Díaz, J., D. K. Riskin, D. J. Willis, K S. Breuer, and S. M. Swartz. 2011. Whole-body kinematics of a fruit bat reveal the influence of wing inertia on body accelerations. Journal of Experimental Biology 214:1546-1553. doi:10.1242/jeb.037804
  • MacAyeal, L. C.*, D. K. Riskin, S. M. Swartz, and K. S. Breuer. 2011. Vertical flight performance and load carrying in lesser dog-faced fruit bats (Cynopterus brachyotis). Journal of Experimental Biology 214:786-793. doi:10.1242/jeb.050195
  • Riskin, D. K., J. Iriarte- Díaz, K. M. Middleton, K. S. Breuer, and S. M. Swartz. 2010. The effect of body size on the wing movements of pteropodid bats, with insights into thrust and lift production. Journal of Experimental Biology 213:4110-4122. doi: 10.1242/jeb.043091
  • Hubel, T. Y., D. K. Riskin, S. M. Swartz, and K. S. Breuer. 2010. Wake structure and wing kinematics: the flight of the lesser dog-faced fruit bat, Cynopterus brachyotis. Journal of Experimental Biology 213: 3427-3440. doi:10.1242/jeb.043257
  • Chen, J., D. K. Riskin, T. Y. Hubel, D. J. Willis, A. Song, H. Liu, K. S. Breuer, S. M. Swartz, and D. H. Laidlaw. 2010. Exploration of bat wing morphology through a strip method and visualization. Special Interest Group on Graphics and Interactive Techniques (SIGGRAPH).
  • Middleton, K. M., B. D. Goldstein*, P. R. Guduru, J. F. Waters, S. A. Kelly, S. M. Swartz and T. Garland, Jr. 2010. Variation in within-bone stiffness measured by nanoindentation in mice bred for high levels of voluntary wheel running. Journal of Anatomy 216:121-131.
  • Chen, J., M. Kostandov, I. V. Pivkin, D. K. Riskin, S. M. Swartz, and D. H. Laidlaw. 2009. Visual analysis of dimensionality reduction in an interactive virtual environment for exploring bat flight kinematics. Joint Virtual Reality Conference of EGVE-ICAT-EUROVR.
  • Riskin, D. K., J. W. Bahlman, T. Y. Hubel, J. M. Ratcliffe, T. H. Kunz, and S. M. Swartz. 2009. Bats go head-under-heels: The biomechanics of landing on a ceiling. Journal of Experimental Biology. 212:945-953.
  • Hubel, T. Y., N. I. Hristov, S. M. Swartz, and K. S. Breuer. 2009. Time-resolved wake structure and kinematics of bat flight. Experiments in Fluids 46:933-943 DOI 10.1007/s00348-009-0624-7
  • Keefe, D. F., D. Acevedo, D., J. Miles, F. Drury, S. M. Swartz, and D. H. Laidlaw. 2008. Scientific sketching for collaborative VR visualization design. IEEE Transactions on Visualization and Computer Graphics 14, 835-847.
  • Waldman, R. M., A. Song, D. K. Riskin, S. M. Swartz, and K. S. Breuer. 2008. Aerodynamic behavior of compliant membranes as related to bat flight. American Institute of Aeronautics and Astronautics Journal: AIAA no. 2008-3716.
  • Riskin, D. K., D. J. Willis, T. L. Hedrick, J. Iriarte-Díaz, M. Kostandov, J. Chen, D. H. Laidlaw, K. S. Breuer, and S. M. Swartz. 2008. Quantifying the complexity of bat wing kinematics. Journal of Theoretical Biology, 254: 604–615. doi: 10.1016/j.jtbi.2008.06.011
  • Iriarte-Díaz, J. and S. M. Swartz. 2008. Kinematics of slow turn maneuvering in the fruit bat Cynopterus brachyotis. Journal of Experimental Biology 211, 3478-3489.
  • Kunz, T. H., S. A. Gauthreaux, Jr, N. I. Hristov, J. W. Horn, G. Jones, E. K. V. Kalko, R. P. Larkin, G. F. McCracken, S. M. Swartz, R. B. Srygley, R. Dudley, J. K. Westbrook, and M. Wikelski. 2008. Aeroecology: probing and modeling the aerosphere. Integrative and Comparative Biology, 8: 1-11.
  • Swartz, S. M. D. J. Willis, and K. S. Breuer. 2008. Aeromechanics in aeroecology: Flight biology in the aerosphere. Integrative and Comparative Biology, 48: 85-98. doi: 10.1093/icb/icn054
  • Song, A, X. Tian, E. Israeli*, R. Galvao*, Bishop, S. Swartz, and Breuer, K. 2008. Aeromechanics of membrane wings with implications for animal flight. AIAA Journal 46:2096-2196. doi: 10.2514/1.36694
  • Middleton, K.M., C.E. Shubin*, D.C. Moore, T. Garland, Jr., P.A. Carter, and S.M. Swartz. 2008. The relative importance of genetics and phenotypic plasticity in dictating bone morphology and mechanics in aged mice: Evidence from an artificial selection experiment. Zoology 111: 135-147.
  • Swartz, S. M. and Middleton, K. M. 2008. Biomechanics of the bat limb skeleton: scaling, material properties and mechanics. Cells Tissues Organs, 187: 59-84. doi: 10.1159/000109964
  • Willis, D. J., E. Israeli, P. Persson, M. Drela, and J. Peraire, S. M. Swartz and K. S. Breuer. 2007. A computational framework for fluid structure interaction in biologically inspired flapping flight. American Institute of Aeronautics and Astronautics Applied Aerodynamics 25: 3803-3809.
  • Willis, D. J., M. Kostandov, D. K. Riskin, J. Peraire, D. H. Laidlaw, S. M. Swartz, K. S. Breuer. 2007. Modeling the flight of a bat: First Place, Informational Graphics, International Visualization Competition. Science 317:1860. doi: 10.1126/science.1133598
  • Swartz, S. M., J. Iriarte-Díaz, D. K. Riskin, A. Song, X. Tian, D. J. Willis, and K.S. Breuer. 2007. Wing structure and the aerodynamic basis of flight in bats. American Institute of Aeronautics and Astronautics Aerospace Sciences 45: 22-26.
  • Chen, J., A. Forsberg, S. M. Swartz, and D. H. Laidlaw. Interactive multiple scale small multiples. IEEE Visualization 2007 Poster Compendium, November 2007.
  • Kostandov, M., I. Pivkin, K. Breuer, S. M. Swartz, and D. H. Laidlaw. 2006. proper orthogonal decomposition and particle image velocimetry in bat flight. IEEE Visualization 2006 Poster Compendium.
  • Tian, X., Iriarte-Díaz, J, Middleton, K, Galvao*, R, Israeli*, E, Roemer*, A, Sullivan*, A, Song, S. M. Swartz and K. S. Breuer. 2006. Direct measurements of the kinematics and dynamics of bat flight. Bioinspiration and Biomimetics. 1:10-18.
  • Pivkin, I., E. Hueso, R. Weinstein*, D. H. Laidlaw, S. Swartz, and G. Karniadakis. 2005. Simulation and Visualization of Air Flow Around Bat Wings During Flight. Proceedings of International Conference on Computational Science, pages 689-694.
  • Sobel, J. S., A. S. Forsberg, D H. Laidlaw, R. C. Zeleznik, D. F. Keefe, I. Pivkin,G. E. Karniadakis, S. M. Swartz, and P. Richardson. 2004. Particle Flurries: Synoptic 3D Pulsatile Flow Visualization. IEEE Computer Graphics and Applications April/May: 2-11.
  • Hueso, E., I. V. Pivkin, S. M. Swartz, D. H. Laidlaw, G. Karniadakis, and K. S. Breuer. 2004. Visualization of Vortices in Simulated Airflow around Bat Wings During Flight. IEEE Visualization 2004 Poster Compendium, October 2004.
  • (non-refereed) Rachel Weinstein*, Igor Pivkin, Sharon Swartz, David H. Laidlaw, George Karniadakis, and K. Breuer. Simulation and visualization of air flow around bat wings during flight. Technical Report CS-02-16, Brown University Computer Science Department, August 2002.
  • Watts, P., E. J. Mitchell*, and S. M. Swartz. 2001. A computational model for estimating mechanics of horizontal flapping flight in bats. Model description and comparison with experimental results. Journal of Experimental Biology. 204: 2873-2898.
  • Swartz, S. M., A. Parker*, and C. Huo*. 1997. Theoretical and empirical scaling patterns and topological homology in bone trabeculae. Journal of Experimental Biology, 201:573-590.
  • Swartz, S. M. 1997. Allometric patterning in the limb skeleton of bats: Implications for the mechanics and energetics of powered flight. Journal of Morphology, 234:277-294.
  • Papadimitriou, H. M. *, S. M. Swartz, and T. H. Kunz. 1996. Ontogenetic and anatomic variation in mineralization of the wing skeleton of the Mexican free-tailed bat, Tadarida brasiliensis. Journal of Zoology, London, 240:411-426.
  • Swartz, S. M., M. D. Groves*, H. D. Kim* and W. R. Walsh. 1996. Mechanical properties of bat wing membrane skin: aerodynamic and mechanical functions. Journal of Zoology, London, 239:357-378.
  • Halgrimmsson, B.* and S. M. Swartz. 1995. Morphological adaptation in the hylobatid ulna: cross-sectional geometry and skeletal loading. Journal of Morphology 224:111-123.
  • Swartz, S. M., M. B. Bennett, and D. R. Carrier. 1992. Wing bone stresses in free flying bats and the evolution of skeletal design for flight. Nature 359:726-729.
  • Anton, S. C*., C. R. Jaslow and S. M. Swartz. 1992. Sutural complexity in artificially deformed human (Homo sapiens) crania. Journal of Morphology 214:321-322.
  • Bertram, J. E. A. and S. M. Swartz. 1991. The “Law of bone transformation”: A case of crying Wolff? Biological Reviews of the Cambridge Philosophical Society 22(3):245-273.
  • Swartz, S. M. 1991. Strain analysis as a tool for functional morphology. American Zoologist 31(4):655-669.
  • Swartz, S. M. 1990. Pendular mechanics and the kinematics and energetics of brachiating locomotion. International Journal of Primatology 10(5):387-418.
  • Swartz, S. M. 1990. Curvature of the limb bones of anthropoid primates: overall allometric patterns and specializations in suspensory species. American Journal of Physical Anthropology 83(4):477-498.
  • Swartz, S. M. 1989. The functional morphology of weight bearing: limb joint surface area allometry in anthropoid primates. Journal of Zoology, London 218:441-460.
  • Swartz, S. M., A. A. Biewener, and J. E. A. Bertram. 1989. Telemetered in vivo strain analysis of locomotor mechanics of brachiating gibbons. Nature 342:270-272.
  • Swartz, S. M. 1987. Skeletal biomechanics and suspensory locomotion: preliminary results of in vivo bone strain analysis of brachiating gibbons. Proceedings of the American Society of Biomechanics 3:151-153.
  • Biewener, A. A., S. M. Swartz and J. E. A. Bertram. 1986. Bone modeling during growth: dynamic strain equilibrium in the chick tibia. Calcified Tissue International 39:390-395.

Book Chapters

*undergraduate co-authors
  • Swartz, S. M., J. Iriarte-Díaz, D. K. Riskin, and K S. Breuer. 2012. A bird? a plane? No, it’s a bat: an introduction to the biomechanics of bat flight. In Evolutionary History of Bats: Fossils, Molecules and Morphology (G. Gunnell and N. B. Simmons, eds). pp. 317-352. Cambridge University Press, Cambridge, UK.
  • Albertani R, T. Y. Hubel, S. M. Swartz, K. S. Breuer, and J. Evers. 2011. In-flight wing-membrane strain measurements on bats. In: Proulx, T. (ed) Experimental and Applied Mechanics. pp. 437-455. Springer, New York.
  • Dumont, E. L. and S. M. Swartz. 2009. Biomechanical approaches and ecological research. In Ecological and Behavioral Methods for the Study of Bats (T. H. Kunz, ed.) pp. 436-458. Johns Hopkins University Press, Baltimore, MD.
  • Swartz, S. M., Bishop, K. L., and Ismael-Aguirre, M. F.* 2005. Bat flight aerodynamics: new insights from three-dimensional kinematic analysis. In Functional and Evolutionary Ecology of Bats (Z. Akbar, G. F. McCracken, and T. H. Kunz, eds) pp. 110-121. Oxford University Press, Oxford, UK.
  • Swartz, S. M., P. Freeman, and E. Stockwell. 2003. Ecomorphology. in Bat Ecology. (T. H. Kunz, ed.) pp. 257-300. The University of Chicago Press, Chicago, IL.
  • Swartz. S. M. 1998. Skin and bones: the mechanical properties of bat wing tissues. in Bats: Phylogeny, Morphology, Echolocation, and Conservation Biology. (T. H. Kunz and P. A. Racey, eds.) Smithsonian Institution Press, Washington, D. C.
  • Swartz, S. M. 1993. The biomechanics of primate limbs. in Postcranial Adaptation in Nonhuman Primates (D. L. Gebo, ed.) pp. 542. Northern Illinois University Press, De Kalb, IL.
  • Swartz, S. M. and A. A. Biewener. 1992. Shape and scaling. in Biomechanics: A Practical Approach. Vol. 2. Structures. (A. A. Biewener, ed.). pp. 20-43. Oxford University Press, Oxford, UK
  • Swartz, H. M. and S. M. Swartz. 1983 Biochemical and Biophysical Applications of Electron Spin Resonance. in Methods of Biochemical Analysis, volume 29, D. Glick, ed. pp. 207-323. John Wiley and Sons, Inc., New York.


Get in touch!

Questions? Ideas? Great bat pictures?

(+1) 401 863-3549 | (+1) 401 863-7544 (fax)
Biomed Center 230, Corner of Meeting and Brown
Dept. of Ecology and Evolutionary Biology, Box G-B206
Brown University, Providence, RI 02912
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