Publications
*Indicates an author who was a student (graduate or undergraduate) when they contributed to the project.
2026
Torque limitations on cantilevering in jumping and gliding snakes
Graham, M.*, H.C. Astley, C.J. Clemente, J.J. Socha. Journal of Experimental Biology 229 (16): jeb251396.
Movie 1 | Download: [pdf]
2025
Pressure-induced circumferential and longitudinal deformations of tracheal tubes in the American cockroach
Webster, M.*, R. Wijeratne*, J.J. Socha, and R. De Vita. Journal of Experimental Biology 228 (23): jeb251415.
| Download: [pdf]
A microfluidic study of transient perfusion through the forewing of dragonfly
Ryu, S., H. Zhang, T. Palone, M.K. Salcedo, and J.J. Socha. Lab on a Chip 25, 3718-3729.
| Download: [pdf]
Two steps forward, one back: kinematic observations of a South American Small-eyed Coralsnake, Micrurus lemniscatus (Linnaeus, 1758), regurgitating a Brown-banded Watersnake, Helicops angulatus (Linnaeus, 1758)
Mortensen, T.L.*, A. Marsh*, J. Anderson, Jr.*, I.T. Moore, B.Z. Socha*, A. Moore*, E. Torres Pacaya, J.J. Socha. Herpetology Notes 18: 293-298.
Movie 1 | Download: [pdf]
Anacondas get vertical on the ground
Socha, J.J., H. Pendar, J.M. Rieser. Nature Physics 1-2.
| Download: [pdf]
2024
Skittering locomotion in the cricket frog Acris crepitans: a form of porpoising
Weiss, T.*, G.B. Gillis, and J.J. Socha. Journal of Experimental Biology 227, jeb249403.
Movie 1 | Movie 2 | Download: [pdf+SI]
Quasi-steady aerodynamic theory under-predicts glide performance in flying snakes
Yeaton, I.J.*, S.D. Ross, and J.J. Socha. Journal of Experimental Biology 227 (19): jeb247989.
| Download: [pdf]
Animal locomotion: Wing-like femoral lobes help orchid mantid nymphs glide
Socha, J.J. and P.C. Khandelwal. Current Biology 34(3) R94-98.
| Download: [pdf]
2023
Dynamic gap crossing in Dendrelaphis, the sister taxon of flying snakes
Graham, M.* and J.J. Socha. Journal of Experimental Biology 226: jeb245094.
Cover image | Download: [pdf]
A year at the forefront of gliding locomotion
Khandelwal, P.C., M.A. Zakaria*, and J.J. Socha. Biology Open. 12 (8): bio059973.
| Download: [pdf]
3D X-ray analysis of Laricobius (Coleoptera: Derodontidae), a specialist predator of Adelges tsugae (Hemiptera: Adelgidae)
Hillen, A.P.*, J.R. Foley, M.K. Salcedo, J.J. Socha, and S.M. Salom. Journal of Insect Science 23(3): 19; 1–7.
| Download: [pdf]
Air sacs are a key adaptive trait of the insect respiratory system
Harrison, J.F., E.K.G. McKenzie, S. Talal, J.J. Socha, M.W. Westneat, and P.G.D. Matthews. Journal of Experimental Biology 226, jeb245712.
| Download: [pdf]
Air-to-land transitions: from wingless animals and plant seeds to shuttlecocks and bio-inspired robots
Ortega-Jimenez, V.M., A. Jusufi, C. Brown, Y. Zeng, S. Kumar, R. Siddall, B. Kim, E. Challita, Z. Pavlik, M. Priess, T. Umhofer, J.-S. Koh, J.J. Socha, R. Dudley, and M.S. Bhamla. Bioinsipiration and Biomimetics 18 (2023) 051001.
| Download: [pdf]
Transient use of hemolymph for hydraulic wing expansion in cicadas
Salcedo, M.K., T.E. Ellis*, A.S. Sáenz*, J.L. Lu*, T. Worrell*, M.L. Madigan, and J.J. Socha. Scientific Reports. 13:6298 (2023).
| Supplementary material | Download: [pdf]
Convergence in Gliding Animals: Morphology, Behavior, and Mechanics
Khandelwhal, P., S.D. Ross, H. Dong, and J.J. Socha. In: Convergent Evolution pp. 391-429 Eds V.L. Bels and A.P. Russell, Springer, New York.
| Download: [pdf]
A biologically accurate model of directional hearing in the parasitoid fly Ormia ochracea
Mikel-Stites*, M.R., M.K. Salcedo, J.J. Socha, P.E. Marek, and A.E. Staples. Bioinspiration and Biomimetics 18 (3) 035007.
| Download: [pdf]
Mallard landing behavior follows a 𝜏̇-constant braking strategy
Whitehead, J.G.*, T. Worrell*, and J.J. Socha. Journal of Experimental Biology 226 (5):jeb244256.
Supplementary material | Download: [pdf]
A living network: complex hemolymph circulation patterns in locust wings
Salcedo, M.K., B.H. Jun*, J.J. Socha, P.P. Vlachos, N.E. Pierce, and S.A. Combes. Communications Biology. 6, 313 (2023).
Movies | Supplementary material | Download: [pdf]
Material properties of skin in a flying snake (Chrysopelea ornata)
Dellinger, S.B.*, R. De Vita, P.P. Vlachos, M.M. Muñoz, and J.J. Socha. Journal of Experimental Zoology A._339: 269–283.
| Download: [pdf]
2022
Computational anlysis of flying-snake-like horizontal undulation during gliding flight
Gong, Y.*, J. Wang, J.J. Socha, and H. Dong. Physics of Fluids. 34: 121907.
| Download: [pdf]
Isometric spiracular scaling in scarab beetles: Implications for diffusive and advective oxygen transport
Wagner*, J.M., C.J. Klok, M.E. Duell*, J.J. Socha, G. Cao, H. Gong*, and J.F. Harrison. 2022. eLife. 11, 82129.
| Download: [pdf]
The impact of sampling frequency on kinetic outcomes
Renner*, K.E., A.T. Peebles*, J.J. Socha, and R.M. Queen. 2022. Journal of Biomechanics. 135 (2022): 111034.
| Download: [pdf]
2021
Dynamic movements facilitate extreme gap crossing in flying snakes
Graham, M.* and J.J. Socha. 2021. The Journal of Experimental Biology, 10.1242/jeb.242923.
| Download: [pdf+SI]
The aerodynamics of flying snake airfoils in tandem configuration
Jafari, F.*, Holden, D., R. La Foy*, P.P. Vlachos, and J.J. Socha. 2021. The Journal of Experimental Biology. 224(14): jeb233635.
Inside JEB | Download: [pdf+SI]
Nonlinear elasticity and damping govern ultrafast dynamics in click beetles
Bolmin, O.*, J.J. Socha, A.C. Dunn, M. Alleyne, and A.A. Wissa. 2021. Proceedings of the National Academy of Sciences of the United States of America. 188(5):118 e2014569118.
Movie S1 | Movie S2 | Supplementary Information | Download: [pdf]
Frequency-specific, valveless flow control in insect-mimetic microfluidic devices
Chatterjee, K.*, P.M. Graybill, J.J. Socha, R.V. Davalos, and A.E. Staples. 2021. Bioinspiration and Biomimetics. 16(3) 036004.
| Movie 1 | Download: [pdf]
2020
Visual acuity in the flying snake, Chrysopelea paradisi
Zamore, S.A., N. Araujo*, and J.J. Socha. 2020. Integrative and Comparative Biology. Invited paper in the 2020 SICB symposium "Long limbless locomotors: The mechanics and biology of elongate, limbless vertebrate locomotion".
| Download: [pdf]
Circulation in insect wings
Salcedo, M.K. and J.J. Socha. 2020. Integrative and Comparative Biology. 10.1093/icb/icaa124. Invited paper in the 2020 SICB symposium "Melding Modeling and Morphology: integrating approaches to understand the evolution of form and function".
| Download: [pdf]
Validity of using 2D video analysis and automated marker tracking to assess continuous sagittal plane running kinematics
Peebles, A.T.*, M.M. Carrol, J.J. Socha, D.O. Schmitt, and R.M. Queen. 2020. Annals of Biomedical Engineering. 10.1007/s10439-020-02569-y.
| View only: [pdf]
Undulation enables gliding in flying snakes
Yeaton, I.J.*, S.D. Ross, G.A. Baumgardner*, J.J. Socha. 2020. Nature Physics. https://doi.org/10.1038/s41567-020-0935-4.
| Nature Video | Nature News & Views | Altmetric media summary | Movies 1-7 | Download: [pdf] [Supplement]
Physiological responses to gravity in an insect
Harrison, J.F., K. Adjerid*, A. Kassi*, C.J. Klok, J.M. VandenBrooks, M.E. Duell*, J.E. Campbell*, S. Talal, E. Alanis*, C. Abdo, H. Pendar, J.J. Socha. 2020. Proceedings of the National Academy of Sciences. doi.org/10.1073/pnas.1915424117.
Altmetric media summary | Movie 1 | Movie 2 | Download: [pdf] [Supplement]
Going the distance: the biomechanics of gap-crossing behaviors
Graham, M.* and J.J. Socha. 2020. Journal of Experimental Zoology A. 333: 60-73. Invited paper for special issue in honor of R. McNeil Alexander. (Published online in 2019)
| Download: [pdf]
2019
Functional compartmentalization in the hemocoel of insects
Pendar, H., J. Aviles*, K. Adjerid*, C. Schoenewald*, and J.J. Socha. 2019. Scientific Reports. 9(1): 6075.
| Download: [pdf]
2018
Patterns of tracheal compression in the thorax of the ground beetle, Platynus decentis
Hochgraf, J.S.*, J.S. Waters*, J.J. Socha. 2018. Yale Journal of Biology and Medicine. 91(4): 409-430.
| Download: [pdf+SI]
How temperature influences the viscosity of hornworm hemolymph
Kenny, M.C.*, M.N. Giarra*, E. Granata*, and J.J. Socha. 2018. Journal of Experimental Biology. 221 (21): jeb186338.
| Download: [pdf+SI | pdf]
Developmental plasticity and stability in the tracheal networks supplying Drosophila flight muscle in response to rearing oxygen level
Harrison, J.F., J.S. Waters, T.A. Biddulph, A. Kovacevic, C.J. Klok, and J.J. Socha. 2018. Journal of Insect Physiology. 106: 189-198.
| Download: [pdf]
Burst mode pumping: A new mechanism of drinking in mosquitoes
Kikuchi, K.*, M.A. Stremler, W.-K. Lee, S. Chatterjee*, O. Mochizuki, and J.J. Socha. 2018. Scientific Reports. 8(1): 4885.
Movie 1 | Movie 2 | Movie 3 | Movie 4 | Movie 5 | Movie 6 | Supplement | Download: [pdf]
NSF video animation | Interview of Socha | APS press release | Virginia Tech press release
2017
Control of gliding in a flying snake-inspired n-chain model
Jafari, F.*, S. Tahmasian, S.D. Ross, and J.J. Socha. 2017. Bioinspiration and Biomimetics. 12(6) 066002.
Movie 1 | Movie 2 | Download: [pdf]
Global dynamics of non-equilibrium gliding in animals
Yeaton, I.J.*, J.J. Socha, and S.D. Ross. 2017. Bioinspiration and Biomimetics. 10(2) 026013.
Noted as a Highlight of 2017 | Download: [pdf]
2016
Bare-hand volume cracker for raw volume data analysis
Laha, B.*, D.A. Bowman, and J.J. Socha. 2016. Frontiers in Robotics and AI. 3: 56.
| Download: [pdf]
Multigenerational effects of rearing atmospheric oxygen level on the tracheal dimensions and diffusing capacities of pupal and adult Drosophila melanogaster
Klok, C.J., A. Kaiser, J.J. Socha, W.-K. Lee, J.F. Harrison. 2016. In Hypoxia: Translation in Progress. Springer US, 285-300. (book chapter)
| Download: [pdf]
Recovering signals in physiological systems with large datasets
Pendar, H., J.J. Socha, and J. Chung. 2016. Biology Open. 5, 1163-1174.
Movie | Supplement | Download: [pdf]
2015
Dogs lap using acceleration-driven open pumping
Gart, S.*, J.J. Socha, P.P. Vlachos, and S. Jung. 2015. Proceedings of the National Academy of Sciences. 112 (52): 15798–15802.
NY Times Science Take | Press release | Journal cover | Download: [pdf]
Structure of tracheae and the functional implications for collapse in the American cockroach
Webster*, M., Socha, J.J., Teresi, L., Nardinocchi, P., De Vita, R. 2015. Bioinspiration & Biomimetics. 10 (2015): 066011,
| Download: [pdf]
Estimation of instantaneous gas exchange in flow-through respirometry systems: A modern revision of Bartholomew's Z-transform method
Pendar*, H. and J.J. Socha. 2015. PLoS ONE.10 (10): e0139508.
Code (direct download) (Github) | Supplement | Download: [pdf]
Tracheal compression in pupae of the beetle Zophobas morio
Pendar*, H., M. Kenny*, and J.J. Socha. 2015. Biology Letters. 11 (6): 20150259.
Science Shot | Cover image | Video abstract | Movies | Dryad data | Supplement | Download: [pdf]
How animals glide: from trajectory to morphology
Socha, J.J., F. Jafari*, Y. Munk, and G. Byrnes. 2015. Canadian Journal of Zoology. 93: 901-924.
| Download: [pdf]
2014
Of snakes and robots
Socha. J.J. 2014. Science. 346 (6206): 160-161.
| Download: [pdf]
A theoretical analysis of gliding in flying snakes
Jafari, F.*, S.D. Ross, P.P. Vlachos, and J.J. Socha. 2014. Bioinspiration and Biomimetics. 9(2): 025014.
| Download: [pdf]
Variation in tensile properties of tracheal tubes in the American cockroach
Becker, W.*, M.R. Webster*, J.J. Socha, and R. De Vita. 2014. Smart Materials and Structures. 23 (5): 057001.
| Download: [pdf]
Effects of VR system fidelity on analzing isosurface visualization of volume datasets
Laha, B.*, D.A. Bowman, and J.J. Socha. 2014. IEEE Transactions on Visualization and Computer Graphics (Proceedings of Virtual Reality 2013). 20 (4): 513-522.
| Download: [pdf]
Lift and wakes of flying snakes
Krishnan, A.* J.J. Socha, P.P. Vlachos, and L.A. Barba. 2014. Physics of Fluids. 26: 03190.
| Download: [pdf]
Aerodynamics of the flying snake, Chrysopelea paradisi: how a bluff-body cross-sectional shape contributes to gliding performance
Holden, D.*, J.J. Socha, N. Cardwell, and P.P. Vlachos. 2014. Journal of Experimental Biology. 217 (3): 382-394.
Video abstract | Journal supplementary video (free via pdf) | Movie 1 | Movie 2 | Movie 3 | Download: [pdf]
2013
Hypoxia-induced compression in the tracheal system of the tobacco hornworm caterpillar, Manduca sexta L
Greenlee, K.J., J.J. Socha, H.B. Eubanks*, G. Thapa, P. Pedersen, W.-K. Lee, and S.D. Kirkton. 2013. Journal of Experimental Biology. 216: 2293-2301.
Journal supplementary video (free via pdf) | Movie 1 | Movie 2 | Download: [pdf]
Dynamics of tracheal compression in the horned passalus beetle
J.S. Waters*, W.-K. Lee, M.W. Westneat, and J.J. Socha. 2013. American Journal of Physiology: Regulatory, Integrative and Comparative Physiology. 304: R621-R627.
Movie S1.mov | Movie S2.mov | Movie S3.mov | Download: [pdf]
How locusts breathe
Harrison, J.F., J.S. Waters*, A.J. Cease, J.M. VandenBrooks, V. Callier, C.J. Klok, K. Shaffer, and J.J. Socha. 2013. Physiology. 28: 18-27.
| Download: [pdf]
Biomechanics of turtle shells: how whole shells fail in compression
Magwene, P.M.* and J.J. Socha*. 2013. Journal of Experimental Zoology A. 319A:86-98. [published online, 2012]
| Download: [pdf]
2011
Gliding flight in Chrysopelea: Turning a snake into a wing
Socha, J.J. 2011. Integrative and Comparative Biology. 51(6): 969–982.
Movie 1 | Movie 2 | Download: [pdf]
Mechanical properties of tracheal tubes in the American cockroach (Periplaneta americana)
Webster, M.*, R. De Vita, J. Twigg*, J.J. Socha. 2011. Journal of Smart Materials and Structures, 20 (2011) 094017.
| Download: [pdf]
2010
Non-equilibrium trajectory dynamics and the kinematics of gliding in a flying snake
Socha, J.J., K. Miklasz*, F. Jafari*, and P.P. Vlachos. 2010. Bioinspiration and Biomimetics, 5 (2010) 045002.
Summary video | Journal supplementary video | Press | Download: [pdf] [corrigendum]
Visceral-locomotory pistoning in crawling caterpillars (Manduca sexta)
Simon, M.A.*, W.A. Woods, Y.V. Serebrenik, S.M. Simon, L.I. van Griethuijsen, J.J. Socha, W.-K. Lee, and B.A. Trimmer. 2010. Current Biology 20: 1-6.
Video abstract | Movie S1 | Movie S2 | Movie S3 | Download: [pdf]
Issues of convection in insect respiration: Insights from synchrotron x-ray imaging and beyond
Socha, J.J., T. Förster*, and K.J. Greenlee. 2010. Respiratory Physiology and Neurobiology, 173S (2010) S65–S73.
| Download: [pdf]
Effects of body cross-sectional shape on flying snake aerodynamics
Miklasz, K.*, M. LaBarbera, X. Chen, and J.J. Socha. 2010. Experimental Mechanics, 50 (9): 1335-1348.
| Download: [pdf]
Canaliculi in the tessellated skeleton of cartilaginous fishes
Dean, M.N.*, J.J. Socha, K.M. Claeson, B.K. Hall, and A.P. Summers. 2010. Journal of Applied Ichthyology 26(2):263-267.
| Download: [pdf]
2009
Synchrotron x-ray visualisation of ice formation in insects during lethal and non-lethal freezing
Sinclair, B.J., A.G. Gibbs, W.-K. Lee, A. Rajamohan*, S.P. Roberts, and J.J. Socha. 2009. PLoS One 2009 4(12):e8259.
Movies | Press release | Download: [pdf]
A plesiosaur containing an ichthyosaur embryo as stomach contents from the Sundance Formation of the Bighorn Basin, Wyoming
O’Keefe, F.R., H.P. Street*, J.P. Cavigelli, J.J. Socha, and R.D. O’Keefe. 2009. Journal of Vertebrate Paleontology 29(4): 1306-1310.
| Download: [pdf]
Direct visualization of hemolymph flow in the heart of a grasshopper (Schistocerca americana)
Lee, W.-K. and J.J. Socha. 2009. BMC Physiology 2009 9:2.
| Download: [pdf]
2008
Correlated patterns of tracheal compression and convective gas exchange in a carabid beetle
Socha, J.J., W.-K. Lee, J.F. Harrison, J.S. Waters*, Fezzaa, K., M.W. Westneat. 2008. Journal of Experimental Biology 211: 3409-3420.
Movies | Cover image | Download: [pdf]
Advances in biological structure, function and physiology using synchrotron x-ray imaging
Westneat, M.W., J.J. Socha, and W.-K. Lee. 2008. Annual Review of Physiology 70: 119-142.
| Download: [pdf]
Use of synchrotron tomography to image naturalistic anatomy in insects
Socha, J.J. and F. De Carlo. 2008. In: Developments in X-Ray Tomography VI: 2008, San Diego, CA, USA: SPIE; 2008: 70780A-70787.
| Download: [pdf]
2007
Increase in tracheal investment with beetle size supports hypothesis of oxygen limitation on insect gigantism
Kaiser, A., C.J. Klok, J.J. Socha, W.-K. Lee, M.C. Quinlan, and J.F. Harrison. 2007. Proceedings of the National Academy of Sciences 104 (32): 13198-13203.
Press release | Commentary | Download: [pdf]
Real-time phase-contrast x-ray imaging: a new technique for the study of animal form and function
Socha, J.J., M.W. Westneat, J.F. Harrison, J.S. Waters*, and W.-K. Lee. 2007. BMC Biology 2007 5:6.
2006
Becoming airborne without legs: the kinematics of take-off in a flying snake, Chrysopelea paradisi
Socha, J.J.* 2006. Journal of Experimental Biology 209 (17): 3358-3369.
Movies | Cover image | Download: [pdf] [pdf with cover]
2005
A three-dimensional kinematic analysis of gliding in a flying snake, Chrysopelea paradisi
Socha, J.J.*, M. LaBarbera, and T. O’Dempsey. 2005. Journal of Experimental Biology 208 (10): 1817-1833.
Movies | Cover image | Download: [pdf] [pdf with cover] [Inside JEB]
Effects of size and behavior on aerial performance of two species of flying snakes (Chrysopelea)
Socha, J.J.* and M. LaBarbera. 2005. Journal of Experimental Biology 208 (10): 1835-1847.
Movies | Cover image | Download: [pdf] [pdf with cover] [Inside JEB]
Chrysopelea ornata, C. paradisi (Flying Snakes). Behavior
Socha, J.J.* and C.A. Sidor*. 2005. Herpetological Review 36(2): 190-191.
| Download: [pdf]
2002
Gliding flight in the paradise tree snake
Socha, J.J.* 2002. Nature 418: 603-604.
| Download: [pdf]