Burnett, N.P., Keliher, E., and Combes, S.A. (2022).  Wind and route choice affect performance of bees flying above versus within a cluttered obstacle fieldPLoS ONE 17(3): e0265911. DOI:10.1371/journal.pone.0265911.


Ravi, S., Siesenop, T., Bertrand, O., Li, L., Doussot, C., Warren, W.H., Combes, S.A., and Egelhaaf, M. (2020).  Bumblebees perceive the spatial layout of their environment in relation to their body size and form to minimize inflight collisionsProceedings of the National Academy of Sciences USA

Burnett, N.P., Badger, M.A., and Combes, S.A. (2020).  Wind and obstacle motion affect honeybee flight strategies in cluttered environmentsJournal of Experimental Biology jeb.222471.

Crall, J.D., Brokaw, J., Gagliardi, S.F., Mendenhall, J.D., Pierce, N.E., and Combes, S.A. (2020).  Wind drives temporal variation in pollinator visitation in a fragmented tropical forestBiology Letters 16: 20200103.

Combes, S.A.*, Gagliardi, S.F.*, Switzer, C., and Dillon, M.E. (2020).  Kinematic flexibility allows bumblebees to increase energetic efficiency when carrying heavy loadsScience Advances 6(6): eaay3115.  [*co-first authors]

Ravi, S., Noda, R., Gagliardi, S., Kolomenskiy, D., Combes, S., Liu, H., Biewener, A.A., and Konow, N. (2020).  Modulation of flight muscle recruitment and wing rotation enables hummingbirds to mitigate aerial roll perturbationsCurrent Biology 30(2): 187-195.


Burnett, N.P. and Combes, S.A. (2019).  Post-doc interviews in the life sciences: An often-overlooked process that is susceptible to biasIntegrative Organismal Biology 1(1): obz027.

Switzer, C.M., Russell, A.L., Papaj, D.R., Combes, S.A., and Hopkins, R. (2019).  Sonicating bees demonstrate flexible pollen extraction without instrumental learningCurrent Zoology 65(4): 425–436.


Crall, J.D., Switzer, C.M., Oppenheimer, R.L., Ford Versypt, A.N., Dey, B., Brown, A., Eyster, M., Guérin, C., Pierce, N.E., Combes, S.A., and de Bivort, B.L. (2018).  Neonicotinoid exposure disrupts bumblebee nest behavior, social networks, and thermoregulationScience 362(6415): 683-686.

Ortega-Jiménez, V.M. and Combes, S.A. (2018).  Living in a trash can: turbulent convective flows impair Drosophila flight performanceJournal of the Royal Society Interface 15(147): 20180636.

Crall, J.D., Gravish, N., Mountcastle, A.M., Kocher, S.D., Oppenheimer, R.L., Pierce, N.E., and Combes, S.A. (2018).  Spatial fidelity of workers predicts collective response to disturbance in a social insectNature Communications 9:1201.

Switzer, C.M., Combes, S.A., and Hopkins, R. (2018).  Dispensing Pollen Via catapult: Explosive Pollen Release in Mountain Laurel (Kalmia latifolia)American Naturalist 191(6): 767-776.

Calisi, R.M., …… Combes, S.A., et al. (2018).  Opinion: How to tackle the conference-childcare conundrumProceedings of the National Academy of Sciences USA 115(12): 2845-2849.


Peters, J.M., Gravish, N., and Combes, S.A. (2017).  Wings as impellers: honey bees co-opt flight system to induce nest ventilation and disperse pheromonesJournal of Experimental Biology 220(12): 2203-2209.

Crall, J.D., Chang, J.J., Oppenheimer, R.L., and Combes, S.A. (2017).  Foraging in an unsteady world: bumblebee flight performance in field-realistic turbulence. Interface Focus 7(1): 20160086.


Mistick, E.A., Mountcastle, A.M., and Combes, S.A. (2016).  Wing flexibility improves bumblebee flight stability.  Journal of Experimental Biology 219(21): 3384-3390.

Chang, J.J., Crall, J.D. and Combes, S.A. (2016).  Wind alters landing dynamics in bumblebees.  Journal of Experimental Biology 219(18): 2819-2822.

Switzer, C.M. and Combes, S.A. (2016).  Bumblebee sonication behavior changes with plant species and environmental conditions.  Apidologie 48(2): 223-233.

Switzer, C.M. and Combes, S.A. (2016).  Bombus impatiens (Hymenoptera: Apidae) display reduced pollen foraging behavior when marked with bee tags vs. paintJournal of Mellitology 62: 1-13.

Switzer, C.M. and Combes, S.A. (2016).  The neonicotinoid pesticide, imidacloprid, affects Bombus imatiens (bumblebee) sonication behavior when consumed at doses below the LD50Ecotoxicology 25(6): 1150-1159.

Mountcastle, A.M., Alexander, T.M., Switzer, C. M., and Combes, S.A. (2016).  Wing wear reduces bumblebee flight performance in a dynamic obstacle courseBiology Letters 12(6): 20160294.

Switzer, C.M., Hogendoorn, K., Ravi, S., and Combes, S.A. (2016).  Shakers and head bangers: differences in sonication behavior between Australian Amegilla murrayensis (blue-banded bees) and North American Bombus impatiens (bumblebees)Arthropod-Plant Interactions 10(1): 1-8.


Crall, J.D., Gravish, N., Mountcastle, A.M., and Combes, S.A. (2015).  BEEtag: A Low-Cost, Image-Based Tracking System for the Study of Animal Behavior and LocomotionPLoS One 10(9): e0136487.

Gravish, N., Peters, J.M., Combes, S.A., and Wood, R.J. (2015).  Collective flow enhancement by tandem flapping wingsPhysical Review Letters 115(18): 188101.

Mountcastle, A.M., Ravi, S., and Combes, S.A. (2015).  Nectar vs. pollen loading affects the tradeoff between flight stability and maneuverability in bumblebees. Proceedings of the National Academy of Sciences USA 112(33): 10527-10532.

Crall, J.D., Ravi, S., Mountcastle, A.M., and Combes, S.A. (2015).  Bumblebee flight performance in cluttered environments: effects of obstacle orientation, body size and accelerationJournal of Experimental Biology 218(17): 2728-2737.

Long, J.H. Jr., Combes, S., Nawroth, J., Hale, M., Lauder, G., Swartz, S., Quinn, R., and Chiel, H. (2015).  How Does Soft Robotics Drive Research in Animal Locomotion?  Soft Robotics 1(3): 161-168.

Ravi, S., Crall, J.D., McNeilly, L., Gagliardi, S.F., Biewener, A.A., and Combes, S.A. (2015).  Hummingbird flight stability and control in freestream turbulent windsJournal of Experimental Biology 218: 1444-1452.

Hedrick, T.L., Miller, L.A., and Combes, S.A. (2015).  Recent developments in the study of insect flightCanadian Journal of Zoology 93: 925-943.


Mountcastle, A.M. and Combes, S.A. (2014).  Biomechanical strategies for mitigating collision damage in insect wings: structural design versus embedded elastic materialsJournal of Experimental Biology 217: 1108-1115.

Combes, S.A. (2014).  Neuroscience: Dragonflies predict and plan their hunts.  Invited News & Views article summary.  Nature 517: 279-280.

Gravish, N., Chen, Y., Combes, S.A. and Wood, R.J. (2014).  High-throughput study of flapping wing aerodynamics for biological and robotic applications. In Intelligent Robots and Systems (IROS 2014), 2014 IEEE/RSJ International Conference, pp. 3397-3403.

Gravish, N., Combes, S. and Wood, R. (2014).  A bio-inspired wing driver for the study of insect-scale flight aerodynamicsLecture Notes in Computer Science 8608: 396-398.

Gerick, A.A., Munshaw, R.G., Palen, W.J., Combes, S.A. and O’Regan, S.M. (2014).  Thermal physiology and species distribution models reveal climate vulnerability of temperate amphibiansJournal of Biogeography 41: 713-723.


Combes, S.A., Salcedo, M.K., Pandit, M.M. and Iwasaki, J.M. (2013).  Capture success and efficiency of dragonflies pursuing different types of preyIntegrative and Comparative Biology 53(5): 787-798.

Ravi, S., Crall, J., Fisher, A. and Combes, S.A. (2013).  Rolling with the flow: bumblebees flying in unsteady wakesJournal of Experimental Biology 216: 4299-4309.

Mountcastle, A.M. and Combes, S.A. (2013).  Wing flexibility enhances load-lifting capacity in bumblebeesProceedings of the Royal Society B 280: 20130531.


Combes, S.A., Rundle, D.E., Iwsaki, J.M. and Crall, J.D. (2012).  Linking biomechanics and ecology through predator-prey interactions: Flight performance of dragonflies and their preyJournal of Experimental Biology 215: 903-913.

Donoughe, S., Crall, J.D., Merz, R.A. and Combes, S.A. (2011).  Resilin in dragonfly and damselfly wings and its implications for wing flexibilityJournal of Morphology 272(12): 1409-1421.

Combes, S.A. (2010).  Materials, structure, and dynamics of insect wings as bioinspiration for MAVs.  In Encyclopedia of Aerospace Engineering, Vol. 7 (Vehicle Design). John Wiley & Sons, West Sussex, UK. 10 pp.

Combes, S.A., Crall, J.D. and Mukherjee, S. (2010).  Dynamics of animal movement in an ecological context: dragonfly wing damage reduces flight performance and predation successBiology Letters 6(3): 426-429.

Combes, S.A. and Dudley, R. (2009).  Turbulence-driven instabilities limit insect flight performanceProceedings of the National Academy of Sciences USA 106(22): 9105-9108.

Shang, J.K., Combes, S.A., Finio, B.M. and Wood, R.J. (2009).  Artificial insect wings of diverse morphology for flapping-wing micro air vehiclesBioinspiration & Biomimetics 4(3): 036002, 6 pp.

Before 2009

Combes, S.A. and Daniel, T.L. (2005).  Flexural stiffness in insect wings: Effects of wing venation and stiffness distribution on passive wing bending. American Entomologist, Spring 2005: 42-44.

Combes, S.A. and Daniel, T.L. (2003).  Flexural stiffness in insect wings. I. Scaling and the influence of wing venation.  Journal of Experimental Biology 206(17): 2979-2987.

 Combes, S.A. and Daniel, T.L. (2003).  Flexural stiffness in insect wings. II. Spatial distribution and dynamic wing bendingJournal of Experimental Biology 206(17): 2989-2997.

Combes, S.A. and Daniel, T.L. (2003).  Into thin air: Contributions of aerodynamic and inertial-elastic forces to wing bending in the hawkmoth Manduca sexta.  Journal of Experimental Biology 206(17): 2999-3006.

Daniel, T.L. and Combes, S.A. (2002).  Flexing wings and fins: bending by inertial or fluid-dynamic forces?  Integrative and Comparative Biology 42(5): 1044-1049.

Combes, S.A. and Daniel, T.L. (2001).  Shape, flapping and flexion: Wing and fin design for forward flightJournal of Experimental Biology 204(12): 2073- 2085.


Combes, S.A. (2005).  Are we putting our fish in hot water? Global warming and the world’s fisheries. Illustrated brochure, 16 pp. Gland, Switzerland: World Wildlife Fund.

Combes, S.A. (2004).  How do vaccines work? A primer in the art of customizing the immune system.  Northwest Science and Technology, Autumn 2004: 14.

Combes, S.A., Prentice, M.L., Hansen, L. and Rosentrater, L. (2003).  Going, going, gone! Climate change and global glacier decline.  Illustrated brochure, 6 pp. Gland, Switzerland: World Wildlife Fund.

Combes, S.A. (2003).  A slough story: science and student mentoring in the brackish waters of an artificial tidal channel.  Northwest Science and Technology, Autumn 2003: 28-32.

Combes, S.A. (2003).  Protecting freshwater ecosystems in the face of global climate change.  In Buying Time: A User’s Manual for Building Resistance and Resilience to Climate Change in Natural Systems (eds. L.J. Hansen, J.L. Biringer and J.R. Hoffman), pp. 203-242. Gland, Switzerland: World Wildlife Fund.

Combes, S.A. (2001).  Bugs, slugs, ‘n chips: downloading biologyNorthwest Science and Technology, Spring 2001, 32-36.

Combes, S.A. (2001).  Environmental threats and conservation. In Brazil – Amazon and Pantanal: The Ecotravellers’ wildlife guide (by L. Beletsky), pp. 39-50. London, England: Academic Press.

Combes, S.A. (2000).  A (non)sticky situation: how geckos climb up walls and why we should care.  Northwest Science and Technology, Autumn 2000: 8-11.

Combes, S.A. and Goldman, E.B. (2000).  Refuge for reflection: UW’s Friday Harbor laboratories.  Northwest Science and Technology, Spring 2000: 34-37.

Combes, S.A. (1999).  SIRTI propels ideas from the lab to the marketplace.  Northwest Science and Technology, Autumn 1999: 26-27.

Combes, S.A. (1999).  Tricking the Body’s Gatekeepers.  Northwest Science and Technology, Autumn 1999: 44.