Flight Lab Personnel

Bret Tobalske

Bret Tobalske

Professor, Director Field Research Station

Home Department: Division of Biological Sciences
Office: Health Sciences 208 and Field Research Station
Email: bret.tobalske@mso.umt.edu
Office Hours:

Monday and Wednesday, 10:00 - 12:00, or by appointment

Personal Website

Current Position

Associate Professor, Division of Biological Sciences

Director, Field Research Station at Fort Missoula


BIOE 403 - Vertebrate Design and Evolution

BIOB 391 - Comparative Biomechanics

BIOB 507 - Organismal Function


Ph.D. (1994) University of Montana, Organismal Biology and Ecology

M.A. (1991) University of Montana, Zoology

B.S. (1988) Southern Illinois University at Carbondale

Research Interests

Comparative Biomechanics, Functional Morphology, Muscle Physiology

Central to the biology of birds, bats, and insects: flight requires more power output than any other form of animal locomotion.   Ultimately, I seek to understand how flight has shaped the ecology and evolution of birds.   I use a variety of  techniques to explore the mechanisms of bird flight in both the laboratory and in the field.  Much of my recent work uses a laser-based system to measure patterns of air flow about the wings and in the wake of flying birds.  I am working to relate these flow dynamics to the in vivo muscle work and power output produced during flight.

Although my primary focus is on bird flight, I am also deeply interested in a wide array of topics in comparative biomechanics and functional morphology. I am open to advising graduate students who wish to study taxa other than birds and other forms of animal performance including swimming and feeding.  I also have an abiding interest in the conservation biology of cavity-nesting birds.

Field of Study

Comparative Biomechanics

Functional Morphology


DeSimone, J.G., Tobalske, B.W. and Breuner, C.W. (2021).  Physiology and behavior under

food limitation support an escape, not preparative, response in the nomadic pine siskin (Spinus pinus).  J. Exp. Biol. 224, jeb238774. doi:10.1242/jeb.238774.


Ton, R., Boyce, A.J., Mitchell, A.E., Mouton, J.C., Gobbo, N.R., Blake, W. and Tobalske, B.W.

(in press).  Galliformes show lighter hearts and lungs, smaller tracheas, but similar skeletal mass than other gamebirds.  Wilson. Bull.


León, B.M. Tobalske, B.W., Sassi, N.B., Garant, R., Powers, D.R. and Harlander-Matauschek,

A. (2021).  Domestic egg-laying hens Gallus gallus domesticus, do not modulate flapping flight performance in response to wing condition.  Roy. Soc. Open Sci. 8: 210196. https://doi.org/10.1098/rsos.210196


Lapsansky, A. B., Zatz, D., and Tobalske, B. W. (2020). Alcids ‘fly’at efficient Strouhal

numbers in both air and water but vary stroke velocity and angle. Elife, 9, e55774, 1-17.


Mouton, J. C., Tobalske, B. W., Wright, N. A., and Martin, T. E. (2020). Risk of predation on

offspring reduces parental provisioning, but not flight performance or survival across early life stages. Functional Ecology. DOI: 10.1111/1365-2435.13650 2020:00:1-11


Niese, R. L., Clark, C. J., and Tobalske, B. W. (2020). Specialized feathers produce sonations

during flight in Columbina ground doves. Integrative and Comparative Biology. 2020:1-33.


Shishido, C. M., Woods, H. A., Tobalske, B. W., Lane, S. J., & Moran, A. L. (2020). Body Size

of Temperate Sea Spiders: No Evidence of Oxygen-Temperature Limitations. The Biological Bulletin, 239(1), 51-61.


Stager, M., Senner, N. R., Tobalske, B. W., & Cheviron, Z. A. (2020). Body temperature

maintenance acclimates in a winter-tenacious songbird. Journal of Experimental Biology. Journal of Experimental Biology 2020 223: jeb221853, 1-11. doi: 10.1242/jeb.221853


van Oorschot, B. K., Choroszucha, R., and Tobalske, B. (2020). Passive aeroelastic deflection of

avian primary feathers. Bioinspiration & Biomimetics. Doi: 10.1088/1748-3190/ab97fd 2020:1-12.


Addis, B. R., Tobalske, B. W., Davenport, J. M., and Lowe, W. H. (2019). A distance–

performance trade‐off in the phenotypic basis of dispersal. Ecology and Evolution. 10.1002/ece3.5583 


Moran, A. L., Harasewych, M. G., Miller, B. A., Woods, H. A., Tobalske, B. W., and Marko, P.

B. (2019). Extraordinarily long development of the Antarctic gastropod Antarctodomus thielei (Neogastropoda: Buccinoidea). Journal of Molluscan Studies. eyz015, https://doi.org/10.1093/mollus/eyz015




Shishido, C. M., Woods, H. A., Lane, S. J., Toh, M. W. A., Tobalske, B. W., & Moran, A. L.

(2019). Polar gigantism and the oxygen–temperature hypothesis: a test of upper thermal limits to body size in Antarctic pycnogonids. Proceedings of the Royal Society B, 286(1900), 20190124.


Lapsansky, A. B., & Tobalske, B. W. (2019). Upstroke-based acceleration and head stabilization

are the norm for the wing-propelled swimming of alcid seabirds. Journal of Experimental Biology, jeb-201285.


Lapsansky, A. B., Igo, J. A., & Tobalske, B. W. (2019). Zebra finch (Taeniopygia guttata) shift

toward aerodynamically efficient flight kinematics in response to an artificial load. Biology open, 8(6), bio042572.


Shishido, C. M., Woods, H. A., Lane, S. J., Toh, M. W. A., Tobalske, B. W., and Moran, A. L.

(2019). Polar gigantism and the oxygen–temperature hypothesis: a test of upper thermal limits to body size in Antarctic pycnogonids. Proceedings of the Royal Society B, 286(1900), 20190124.


Moran, A. L., Woods, H. A., Shishido, C. M., Lane, S. J., and Tobalske, B. W. (2018). Predatory

behavior of giant Antarctic sea spiders (Colossendeis) in nearshore environments. Invertebrate biology, 137(2), 116-123.


Martin, T.E., Tobalske, B., Riordan, M.M., Case, S.B. and Dial, K.P. (2018) Age and

performance at fledging are a cause and consequence of juvenile mortality between life stages.  Sci. Adv. 2018;4: eaar1988


Groom, D.J.E., Toledo, M.C.B., Powers, D.R., Tobalske, B.W. and Welch, K.C. Jr. (2018), Integrating morphology and kinematics in the scaling of hummingbird hovering metabolic rate and efficiency.  Proc. Roy. Soc. B. 285, 20172011.


Crandell, K.E., Smith, A.F., Crino, O.L., and Tobalske, B.W. (2018). Coping with compliance during take-off and landing in the diamond dove (Geopelia cuneata).  PLoS One, 13, e0199662.


Lane, S.J, Moran, A.L., Shishido, C.M., Tobalske, B.W. and Woods, H.A. (2018). Cuticular gas exchange by Antarctic sea spiders. J. Exp. Biol. 221, jeb177568.  doi:10.1242/jeb.177568


Lane, S.J., Tobalske, B.W., Moran, A.L., Shishido, C.M., and Woods, H.A. (2018). Costs of epibionts on Antarctic sea spiders.  Marine Biol. 165, 137 https://doi.org/10.1007/s00227-018-3389-9


Moran, A.L., Woods, A.H., Shishido, C.M., Lane, S.J. and Tobalske, B.W. (2018). Predatory behavior of giant Antarctic sea spiders (Colossendeis) in nearshore environments.  Invert. Biol. 127, 116-123.


LeBlanc, C., Tobalske, B., Szkotnicki, B., Harlander-Matauschek, A. (2018). Locomotor behavior of chickens anticipating incline walking.  Front. Vet. Sci. 4:233. doi: 10.3389/fvets.2017.00233


Wang, Y., Tobalske, B.W., Cheng, B. and Deng, X. (2018).  Gravitation-enabled forward acceleration during flap-bounding flight in birds.  J. Bionic. Eng. 15, 505-515.


Lane, S.J., Shishido, C.M., Moran, A.L., Tobalske, B.W., Arango, C.P. and Woods, H.A. (2017). Upper limits to body size imposed by respiratory – structural trade-offs in Antarctic pycnogonids.  Proc. Roy. Soc. B., 284, 20171779.


Powers, D.R. Langland, K.M., Wethington, S.M., Powers, S.M., Graham, C.H., and Tobalske, B.W. (2017). Hovering in the heat: effects of environmental temperature on heat regulation in foraging hummingbirds.  Roy. Soc. Open Sci. 4, 171056.


Crino, O. L., Klaassen van Oorschot, B., Crandell, K. E., Breuner, C. W., & Tobalske, B. W. (2017). Flight performance in the altricial zebra finch: Developmental effects and reproductive consequences. Ecology and evolution, 7(7), 2316-2326.


Klaassen van Oorschot, B., Tang, H. K., & Tobalske, B. W. (2017). Phylogenetics and ecomorphology of emarginate primary feathers. Journal of Morphology.

Woods, Lane, Shishido, Tobalske, Arango, Moran.   (2017). Respiratory gut peristalsis by sea spiders.  Current Biology. 27 (13): R638 DOI: 10.1016/j.cub.2017.05.062

Tobalske, Jackson, Dial. (2017). Ontogeny of flight capacity and pectoralis function in a precocial ground bird (Alectoris chukar).  Integrative and Comparative Biology.


LeBlanc, C. Tobalske, B., Bowley, S. and Harlander-Matauschek, A. (2017).  Development of

locomotion over inclined surfaces in laying hens.  Animal: 2017: 1 12. doi:10.1017/S1751731117001896


Cheng, B., Tobalske, B.W., Powers, D.R., Hedrick, T.L., Wethington, S.M. Chiu, G.T.C. and

Deng, X. (2016).  Flight mechanics and control of escape manoeuvres in hummingbirds I. flight kinematics.  Journal of Experimental Biology, jeb-137539.


Cheng, B., Tobalske, B.W., Powers, D.R., Hedrick, T.L, Wang, Y., Wethington, S.M., Chiu,

G.T.C. and Deng, X. (2016). Flight mechanics and control of escape manoeuvres in hummingbirds II. Aerodynamic force production, flight control and performance limitations. Journal of Experimental Biology, jeb-137570.


Klaassen van Oorshot, B., Mistick, E.A. and Tobalske, B.W. (2016).  Aerodynamic

consequences of wing morphing during emulated take-off and gliding flight in birds.  Journal of Experimental Biology, 219: 3146-3154; doi: 10.1242/jeb.136721


Kozak, M., Tobalske, B., Martins, C., Bowley, S., Wuerbel, H., & Harlander-Matauschek, A.

(2016). Use of space by domestic chicks housed in complex aviaries. Applied Animal Behaviour Science. 181, 115-121.



Lane, S. J., Shishido, C. M., Moran, A. L., Tobalske, B. W., & Woods, H. A. (2016). No Effects

and No Control of Epibionts in Two Species of Temperate Pycnogonids. Biological Bulletin, 230(2), 165-173.


LeBlanc S, Tobalske B, Quinton M, Springthorpe D, Szkotnicki B, Wuerbel H, et al. (2016)

Physical health problems and environmental challenges influence balancing behavior in

laying hens. PLoS ONE 11(4): e0153477. doi:10.1371/journal.pone.0153477


Niese, R. L., & Tobalske, B. W. (2016). Specialized primary feathers produce tonal

sounds during flight in rock pigeons (Columba livia). Journal of Experimental Biology 219, 2173-2181 doi:10.1242/jeb.131649


Powers, D. R., Tobalske, B. W., Wilson, J. K., Woods, H. A., & Corder, K. R. (2015). Heat

dissipation during hovering and forward flight in hummingbirds. Royal Society Open Science, 2(12), 150598.


Ren, Y., Dong, H., Deng, X., & Tobalske, B. (2016). Turning on a dime: Asymmetric vortex

formation in hummingbird maneuvering flight.  Physical Review Fluids, 1, 050511.


Song, J., Tobalske, B. W., Powers, D. R., Hedrick, T. L., & Luo, H. (2016). Three-dimensional

simulation for fast forward flight of a calliope hummingbird. Royal Society Open Science, 3(6), 160230.


Tobalske, B.W. (2016).  Avian Flight. in Fitzpatrick, Koenig, Lovette and Winkler (Eds.), 

Handbook of Bird Biology, 3rd edition.  Blackwell, Oxford.


Tobalske, B.W. (2016). Evolution of avian flight: muscles and constraints on performance. 

Philosphical Transactions of the Royal Society, B. 371: 20150383


Warrick, D.R., Hedrick, T.L., Biewener, A.A. Crandell, K.E. and Tobalske, B.W. (2016).

Foraging at the edge of the world: low-altitude, high-speed maneuvering in barn

swallows.  Philosphical Transactions of the Royal Society, B 371 20150391.


Crandell, K.E. and Tobalske, B.W. (2015). Kinematics and aerodynamics of avian

upstrokes during slow flight. Journal of Experimental Biology, 218, 2518-2527.


Harlander-Matauschek, A., Rodenburg, T.B., Sandilands, V., Tobalske, B.W. and

Toscano, M.J. (2015) Causes of keel bone damage and their solutions in laying hens.  World’s Poultry Science Journal, 71, 461-472.


Song, J., Luo, H., Tobalske, B.W., Hedrick, T.L. (2015). Analysis of cruise flight of the

Calliope hummingbird, American Institute of Aeronautics and Astronautics,



McCullough, E.M., Tobalske, B.W. and Emlen, D.J. (2014).  Structural adaptations to

diverse fighting styles in sexually selected weapons. Proceedings of the National Academy of Sciences doi/10.1073/pnas.1409585111


Heers, A.M., Dial, K.P. and Tobalske, B.W. (2014).From baby birds to feathered dinosaurs:

incipient wings and the evolution of flight. Paleobiology, 40, 459-476.


Provini, P., Tobalske, B.W., Crandell, K.E. and Abourachid, A. (2014).  Transition from wing to

leg forces during landing in birds.  Journal of Experimental Biology, 217, 2659-2666.


McCullough, E.L. and Tobalske, B.W. (2013).  Elaborate horns in a giant rhinoceros beetle incur

negligible aerodynamic costs.  Proceedings of the Royal Society B, 10.1098/rspb.2013.0197. 


D’Agostino, F., Dawson, H. and Tobalske, B.W.  (2013). Motion studies, in Bowen, Keene and

Ng (Eds.) Electronic Visualization in Arts and Culture.  Springer, London.


Powers, D. R., Getsinger, P.W., Tobalske, B.W., Werthington, S.M., Powers, S.D. andWarrick, D.R. (2012). Respiratory evaporative water loss during hovering and forward flight in hummingbirds. Comparative Biochemistry and Physiology, A.161, 279-285.


Warrick, D.R. Hedrick, T., Fernandez, M.J., Tobalske, B. and Biewener, A. (2012). Hummingbird flight. Current Biology, 22, R472-R477.


Dial, T.R., Heers, A.M. and Tobalske, B.W. (2012). Ontogeny of aerodynamics inmallards: comparative performance and developmental implications. Journal of

Experimental Biology. 215, 3693-3702.


Provini, P., Tobalske, B.W., Crandell, K.E. and Abourachid, A. (2012). Transition from

leg to wing forces during take-off in birds. J. Exp. Biol,



Hedrick, T.L, Tobalske, B.W., Ros, I.G., Warrick, D.R. and Biewener, A.A. (2011). Morphological and kinematic basis of the hummingbird flight stroke: scaling of flight muscle transmission ratio. Proceedings of the Royal Society of London, B.doi:10.1098/rspb.2011.2238


Heers, A.M., Tobalske, B.W. and Dial, K.P. (2011). Ontogeny of lift and drag production in ground birds. J. Exp. Biol., 214, 717-725.


Jackson, B.E., Tobalske, B.W. and Dial, K.P. (2011). The broad range of contractile behaviour

of the avian pectoralis: functional and evolutionary implications. J. Exp. Biol., 214, 2354-2361.


Crandell, K.E. and Tobalske, B.W. (2011). Aerodynamics of tip-reversal upstroke in a revolving pigeon wing. J. Exp. Biol., 214, 1867-1873.


Tobalske, B.W., Warrick, D.R., Jackson, B.E. and Dial, K.P. (2011) Morphological correlates of flapping flight. Chapter in Edited Volume, Editor(s): Gareth Dyke and Gary Kaiser Evolution of Modern Birds,Wiley-Blackwell, Hoboken, NJ.

Tobalske, B.W., Hearn, J.S. and Warrick, D.R. (2010). Aerodynamics of intermittent boundsin flying birds. (2010). Editor(s): Graham K. Taylor, Michael S. Triantafyllou and Cameron Tropea, Collection: Animal Locomotion, The Physics of Flying, The Hydrodynamics of Swimming, Springer-Verlag, Berlin


Tobalske, B.W., Biewener, A.A., Warrick, D.R., Hedrick, T.L. and Powers, D.R. (2010) Effects of flight speed upon muscle activity in hummingbirds, J. Exp. Biol.,  213, 2515-2523. doi: 10.1242/jeb.043844


Tobalske, B.W. (2010). Hovering and intermittent flight in birds. Bionspiration &Biomimetics. 5, 1-10.


Fernanda D'Agostino, Harry Dawson and Bret W. Tobalske (2010). Motion Studies: an Art and Science Collaboration", Electronic Visualization and the Arts. 2010, 303-310.


Tobalske, B.W., Hearn, J.W.D. and Warrick, D.R. (2009). Aerodynamics of intermittent bounds in flying birds. Experiments in Fluids, 46, 963-973. DOI 10.1007/s00348-009-0614-9.


Warrick, D.R., Tobalske, B.W. and Powers, D.R. (2009). Lift production in the hovering hummingbird. Proceedings of the Royal Society of London, B. 276, 3747-3752.


Tobalske, B.W. and Biewener, A.A. (2008).  Contractile properties of the pigeon supracoracoideus during different modes of flight. J. Exp. Biol. 211, 170-179.


Tobalske, B.W. (2007).  Biomechanics of bird flight.  J. Exp. Biol.210, 3135-3146.


Tobalske, B.W. Warrick, D.R., Clark, C.J. Powers, D.R., Hedrick, T.L. Hyder, G.A., Biewener, A.A. (2007).  Three-dimensional kinematics of hummingbird flight.  J. Exp. Biol. 210, 2368-2382.


Warrick, D.R., Tobalske, B.W. Powers, D.R. and Dickinson, M.H. (2007).  The aerodynamics of hummingbird flight.  Am. Inst. Aero. Astro. 2007-41, 1-5


Tobalske, B.W. and Dial, K.P.  (2007). Aerodynamics of wing-assisted incline running in birds.  J. Exp. Biol., 210, 1742-1751. 


Tobalske, B.W., Puccinelli, L.A. and Sheridan, D.C.  (2005).  Contractile activity of the pectoralis in the zebra finch according to mode and velocity of flap-bounding flight. J. Exp. Biol. 208, 2895-2901.


Warrick, D.R., Tobalske, B.W. and Powers, D.R.  (2005).  Aerodynamics of hovering in  hummingbirds. Nature.  435, 1094-1097.


Tobalske, B.W., Altshuler, D. and Powers, D.  (2004).  Take-off mechanics in hummingbirds (Trochilidae).  J. Exp. Biol.  207, 1345-1352.

Tobalske, B.W., T.L. Hedrick, and A.A. Biewener. (2003).  Wing kinematics of avian flight across speeds.  J. Avian Biol. 34, 177-184.


Tobalske, B.W., T.L. Hedrick, K.P. Dial, and A.A. Biewener. (2003).  Comparative power curves in bird flight.  Nature.  421, 363-366.


Hedrick, T.L., B.W. Tobalske, and A.A. Biewener (2003).  How cockatiels (Nymphicus hollandicus) modulate pectoralis power output across flight speeds.  J. Exp. Biol. 206, 1363-1378.


Hedrick, T.L., B.W. Tobalske, and A.A. Biewener.  (2002). Estimates of circulation and gait change based on a three-dimensional kinematic analysis of flight in cockatiels (Nymphicus hollandicus) and ringed turtle-doves (Streptopelia risoria).  J. Exp. Biol. 205, 1389-1409.


Tobalske, B.W. (2001).  Morphology, velocity, and intermittent flight in birds. 

Amer. Zool. 41, 177-187.


Warrick, D.R., B.W. Tobalske, A.A. Biewener, and K.P. Dial.  (2001).  Sonomicrometry and

kinematic estimates of the mechanical power of bird flight. Bull. Mus. Comp.Zool.           

156, 257-268.


Tobalske, B.W.  (2000).  Biomechanics and physiology of gait selection in flying birds. Physiol. Biochem. Zool. 73, 736-750.


Tobalske, B.W. and K.P. Dial.  (2000). Effects of body size on takeoff flight performance in the Phasianidae (Aves).  J. Exp. Biol. 203, 3319-3332.


Tobalske, C., and B.W. Tobalske.  (1999).  Using atlas data to model the distribution of woodpecker species in the Jura, France. Condor 101, 472-483.


Tobalske, B.W., W. Peacock, and K.P. Dial.  (1999).  Kinematics of flap-bounding flight in the zebra finch over a wide range of speeds.  J. Exp. Biol. 202, 1725-1739.


Biewener, A. A., W.R. Corning, and B.W. Tobalske. (1998). In vivo pectoralis muscle

force-length behavior during level flight in pigeons (Columba livia). J. Exp. Biol. 201, 3293-3307.


Tobalske, B.W., N.E. Olson, and K.P. Dial.  (1997).  Flight of the black-billed magpie:variation in wing kinematics, neuromuscular control, and muscle composition. J. Exp. Zool. 279, 313-329.


Tobalske, B.W.  (1997).  Lewis' Woodpecker (Melanerpes lewis).  In   The Birds of North America, No. 284 (A. Poole and F. Gill, eds.).  The Academy of Natural Sciences, Philadelphia, PA, and The American Ornithologists’ Union, Washington, D.C.


Dial, K.P., A.A. Biewener, B.W. Tobalske, and D.R. Warrick.  (1997).  Mechanical power

output of bird flight.  Nature 390, 67-70.

Tobalske, B.W. and K.P. Dial.  (1996).  Flight kinematics of the Black-billed Magpie and Rock Dove over a wide range of speeds.  J. Exp. Biol. 199, 263-280.


Tobalske, B.W.  (1996).  Scaling of muscle composition, wing morphology, and intermittent flight behavior in woodpeckers.  Auk  113, 151-177.


Tobalske, B.W.  (1995).  Neuromuscular control and kinematics of intermittent flight in the European Starling (Sturnus vulgaris). J. Exp. Biol. 198, 1259-1273.


Tobalske, C. and B.W. Tobalske.  (1995).  Avifaune et activites humaines: Une étude de la distribution des pics dans le Jura avec un SIG.  Actes du 35Éme Colloque Interégional d'Ornithologie, Lyon, France. 154-174.


Tobalske, B.W., R.C. Shearer, and R.L. Hutto.  (1995).  Maintaining bird diversity in Western Larch/Douglas-fir Forests.  In (W.C. Schmidt and K.J. McDonald, eds.).    Ecology and management of Larix forests:  a look ahead.  Proceedings of an      international symposium; 1992 October 5-9; Whitefish, MT, U.S.A. Gen. Tech. Rep.                   INT-GTR-319.  Ogden, UT: U.S. Department of Agriculture.  Forest Service,                                            Intermountain Research Station.  521 p.


Tobalske, B.W. and K.P. Dial.  (1994).  Neuromuscular control and kinematics of             intermittent

flight in budgerigars (Melopsittacus undulatus).  J. Exp. Biol. 187, 1-18.


Tobalske, B.W.  (1992).  Evaluating habitat suitability using relative abundance and fledging success of red-naped sapsuckers.  Condor 94, 550-553.


Tobalske, B.W., R.C. Shearer, and R.L. Hutto.  (1991).  Bird populations in logged and

unlogged western larch/Douglas-fir forest in northwestern Montana.  Res. Pap. INT-442.  Ogden, UT: U.S. Department of Agriculture.  Forest Service, Intermountain Research           Station.  12 p.


Home Department

Division of Biological Sciences

Area of Expertise

Comparative biomechanics: anatomy, physiology, and fluid dynamics of animal locomotion in the air and water

Teaching Experience

University of Montana (2008-Present)

University of Portland (1999-2008)

Allegheny College (1997-1998)

International Experience

Fulbright Fellowship (1995), France

Sabbatical (2007), France


American Ornithologists' Union

Cooper Ornithological Society

Sigma Xi

Society for Experimental Biology

Society for Integrative and Comparative Biology


Music: Playing, Composing, Recording

Outdoors:  Bicycling, SCUBA, Camping, Hiking