Peer-Reviewed Publications (with lab members in bold)

In press, in revision, or in review

Bidder, O., T. Connor, J. Morales, G. Rickbeil, J. Merkle, R. Fuda, J. Rogerson, B. Scurlock, W. Edwards, E.  Cole, D. McWhirter, A. Courtemanch, S. Dewey, M. Kauffman, D. MacNulty, J. du Toit, Johan, D. Stahler, A. D. Middleton. In review. Forage senescence and disease status influence elk pregnancy across the Greater Yellowstone Ecosystem.


Serota, M.W., K.J. Barker, L.C. Gigliotti, S.M.L. Maher, A.L. Shawler, G.R. Zuckerman, W. Xu, G. Verta, E. Templin, C.L. Andreozzi, A.D. Middleton. 2023. Incorporating human dimensions is associated with better wildlife translocation outcomes. Nat Commun 14, 2119.

Barker, K.J., E. Cole, A. Courtemanch, S. Dewey, D. Gustine, K. Mills, J. Stephenson, B. Wise, and A.D. Middleton. 2023Large carnivores avoid humans while prioritizing prey acquisition in anthropogenic areas. Journal of Animal Ecology 92:889-900.

Karandikar, H., E. Donadio, J.A. Smith, O.R. Bidder, and A.D. Middleton. 2023. Spatial ecology of the Vicuña (Lama vicugna) in a high Andean protected area. Journal of Mammalogy.

Perrig, P.L., S.A. Lambertucci, E. Donadio, J.A. Smith, A.D. Middleton, and J.N. Pauli2023. Risk effects cascade up to an obligate  scavenger. Ecology 104:e3871.

Xu, W., L.C. Gigliotti, R. Royauté, H. Sawyer, A.D. Middleton. 2023Fencing amplifies individual differences in movement with implications on survival for two migratory ungulates. Journal of Animal Ecology, 92:677-689.

Gigliotti, L. C., M.P. Atwood, E.K. Cole, A. Courtemanch, S. Dewey, J.A. Gude, M. Hurley, M.J. Kauffman, K. Kroetz, B. Leonard, D. MacNulty, E. Maichak, D. McWhirter, T.W. Mong, K. Proffitt, B. Scurlock, D. Stahler, and A.D. Middleton. 2023Multi-level thresholds of residential and agricultural land use for elk avoidance across the Greater Yellowstone Ecosystem. Journal of Applied Ecology, 001-11.


Gigliotti, L.C., W. Xu, G.R. Zuckerman, M.P. Atwood, E.K. Cole, A. Courtemanch, S. Dewey, J.A. Gude, P. Hnilicka, M. Hurley, M.J. Kauffman, K. Kroetz, A. Lawson, B. Leonard, D. MacNulty, E. Maichak, D. McWhirter, T.W. Mong, K. Proffitt, B. Scurlock, D. Stahler, and A.D. Middleton. 2022. Wildlife migrations highlight importance of both private lands and protected areas in the Greater Yellowstone Ecosystem. Biological Conservation 275:109752.

Karandikar, H., M.W. Serota, W.C. Sherman, J.R. Green, G. Verta, C. Kremen, and A.D. Middleton. 2022Dietary patterns of a versatile large carnivore, the puma (Puma concolor). Ecology and Evolution 12:e9002.

Monk, J. D., J. A. Smith, E. Donadio, P. Perrig, J. N. Pauli, O. Bidder, M. Fileni, P Gregorio, O. J. Schmitz, and A. D. Middleton. 2022. Cascading effects of disease in a remote protected area. Ecology Letters 25:1152-1163.

Rodriguez Curras, M., E. Donadio, A.D. Middleton, and J.N. Pauli. 2022. Carnivore niche partitioning in a human landscape. The American Naturalist 199:496-509.


Barker, K. J., W. Xu, A. Van Scoyoc, M. Serota, J. Moravek, A. Shawler, R. Ryan, and A. D. Middleton. 2021. Toward a new framework for restoring lost wildlife migrations. Conservation Letters e12850.

Curras, M., E. Donadío, A.D. Middleton, and J.N. Pauli. 2021. Perceived risk structures the space use of competing carnivores. Behavioral Ecology 32:1380-90.

Kauffman, M. J., F. Cagnacci, S. Chamaille-Jammes, [and 89 others, including A. D. Middleton]. 2021. Mapping out a future for ungulate migrations. Science. 372:566-569.

Kauffman M. J., E. Aikens, S. Esmaeili, P. Kaczensky, A. D. Middleton, K. L. Monteith, T. A. Morrison, T. Mueller, H. Sawyer, and J. Goheen. 2021. Causes, consequences and conservation of ungulate migration. Annual Reviews in Ecology and Systematics 52:453-78.

Perrig, P., S. Lambertucci, P. Alarcón, A. D. Middleton, J. Padró, P. Plaza, G. Blanco. J. A. Sánchez-Zapata, J. Donazar; J. N. Pauli. 2021. Limited sexual segregation in a dimorphic avian scavenger, the Andean condor. Oecologia 196:77-88.

Xu, W., K. Barker, A. Shawler, A. Van Scoyoc, H. Sawyer, T. Mueller, C. Andreozzi, O. Bidder, H. Karandikar, S. Mumme, J. Smith, E. Templin, and A.D. Middleton. 2021. The plasticity of ungulate migrations in a changing world. Ecology e03293.

Xu, W., N. Dejid, V. Herrmann, H. Sawyer, and A.D. Middleton. 2021. Barrier Behaviour Analysis (BaBA) reveals extensive effects of fencing on wide‐ranging ungulates. Journal of Applied Ecology 58:690-698.

Morrison, T., J. Merkle, G. Hopcraft, E.O. Aikens, J.L. Beck, R.B. Boone, A. Courtemanch, S.P. Dwinnell, W.S. Fairbanks, B. Griffith, A.D. Middleton, K. Monteith, B. Oates, H. Sawyer, K. Smith, J. Stabach, K. Taylor, and M. J. Kauffman. 2020. Drivers of site fidelity in ungulates. Journal of Animal Ecology 90:955-966.


Smith, J.A., E. Donadio, O.R. Bidder, J.N. Pauli, M.J. Sheriff, P.L. Perrig, and A.D. Middleton. 2020. Where and when to hunt? Decomposing predation success of an ambush carnivore. Ecology 101:p.e03172.

Middleton, A.D., T. Stoellinger, H. Karandikar, B. Leonard, H. Doremus, and C. Kremen. 2020. Harnessing visitors' enthusiasm for national parks to fund cooperative large‐landscape conservation. Conservation Science and Practice, p.e335.

Aikens, E.O., A. Mysterud, J.A. Merkle, [and 33 others, including A. D. Middleton]. 2020. Wave-like patterns of plant phenology determine ungulate movement tactics. Current Biology 30: 3444-3449.e4.

Perrig, P.L., S.A. Lambertucci, J. Cruz Bernal, P.A.E. Alarcón, P. Plaza, A.D. Middleton, G. Blanco, J.A. Sánchez, Zapata, J.A. Donazar, and J.N. Pauli. 2020. Identifying conservation priority areas for the Andean condor in southern South America. Biological Conservation 243: 108494.

Middleton, A.D., H. Sawyer, J.A. Merkle, M.J. Kauffman, E.K. Cole, S.R. Dewey, J.A. Gude, D.D. Gustine, D.E. McWhirter, K.M. Proffitt, and P.J. White. 2020. Conserving ungulate migrations across the Greater Yellowstone Ecosystem requires transboundary science, policy, and management. Frontiers in Ecology and the Environment 18: 83-91.


Sawyer, H., C. W. Lebeau, T. L. MacDonald, W. Xu, and A.D. Middleton. 2019. All routes are not created equal: An ungulate’s choice of migration route can influence its survival. Journal of Applied Ecology 56: 1860-69.

Rickbeil, G.J.M., J.A. Merkle, G. Anderson, M.P. Atwood, J.P. Beckmann, E.K Cole, A.B. Courtemanch, S. Dewey, D.D. Gustine, M.J. Kauffman, D.E. McWhirter, T. Mong, K. Profitt, D.R Stahler, P.J. White, and A.D. Middleton. 2019. Plasticity in elk migration timing is a response to changing environmental conditions. Global Change Biology 25: 2368-2381.

Smith, J.A., E. Donadio, J.N. Pauli, M.J. Sheriff, and A.D. Middleton. 2019. Integrating temporal refugia into landscapes of fear: prey exploit predator downtimes to forage in risky places. Oecologia 189:883-890.

Smith, J.A., E. Donadio, J.N. Pauli, M.J. Sheriff, A.D. Middleton. 2019. Habitat complexity mediates the predator-prey space race. Ecology 100: e02724.

Gaynor, K.M., J.S. Brown, A.D Middleton, M.E. Power, and J.S. Brashares. 2019. Landscapes of fear: spatial patterns of risk perception and response. Trends in Ecology and Evolution 34:355-368.

Sawyer, H., J. Merkle, A.D. Middleton, S.P.H.  Dwinnell, and K. Monteith. 2019Migratory plasticity is not ubiquitous among large herbivores. Journal of Animal Ecology 88:450-460.


van Eeden, L.M., A. Eklund, J.R.B. Miller, J. V. Lopez-Bao., G. Chapron, M.R. Cejtin, M. S. Crowther, C.R. Dickman, J. Frank, M. Krofel, D.W. Macdonald, J. McManus, T.R. Meyer, A.D. Middleton, T.M. Newsome, W.J. Ripple, E. G. Richie, O.J. Schmitz, K. J. Stoner, M. Tourani, A. Treves. 2018. Carnivore conservation needs evidence-based livestock protection.  PLoS Biology 16:e2005577.

Middleton, A.D., J.A. Merkle, D.E. McWhirter, J.G. Cook, R.C. Cook, P.J. White, M.J. Kauffman. 2018. Green-wave surfing influences fitness in a large migratory herbivore. Oikos 127: 1060-1068.

Lukacs, P., M. Hebblewhite, B. Johnson, H. Johnson, M. Kauffman, M. Mitchell, K. Proffitt, P. Zager, J. Brodie, K. Hersey, A. Holland, M. Hurley, S. McCorquodale, A.D. Middleton, M. Nordhagen, J. Nowak, D. Walsh, P.J. White. 2018. Factors influencing elk recruitment across three ecotypes in the western USA. Journal of Wildlife Management 82: 698-710.

Cotterill, G.G., P.C. Cross, A.D. Middleton, J.D. Rogerson, B.M Scurlock, J.T. Toit, JT. 2018. Hidden cost of disease in a free‐ranging ungulate: brucellosis reduces mid‐winter pregnancy in elk. Ecology and Evolution 8: 10733-10742.


Perrig, P., E. Donadio, A.D. Middleton, and J.N. Pauli. 2016. Predation subsidizes an obligate scavenger in the high Andes. Journal of Animal Ecology 54: 846-853.

Miller, J.R.B., K.J. Stoner, M.R. Cejtin, T.K. Meyer, A.D. Middleton, and O.J. Schmitz. 2016. Tools for human-carnivore coexistence: A quantitative assessment of techniques for reducing large carnivore depredation of livestock. Wildlife Society Bulletin 40:806-815.

Sawyer, H., A.D. Middleton, K. Monteith, M. Hayes, and M.J. Kauffman. 2016. The extra mile: Ungulate migration distance alters use of seasonal range and exposure to anthropogenic risk. Ecosphere 7:e01534.

Merkle, J.A., K.L. Monteith, E.O. Aikens, M.M. Hayes, K.R. Hersey, A.D. Middleton, B.A. Oates, H. Sawyer, B.M. Scurlock, and M.J. Kauffman. 2016. Large herbivores surf waves of green-up in spring. Proceedings of the Royal Society of London B 283:20160456.

Nelson, A.A., M.J. Kauffman, A.D. Middleton, M.D. Jimenez, D.E. McWhirter, and K. Gerow. 2016. Native prey distribution and migration mediates wolf predation on domestic livestock in the Greater Yellowstone Ecosystem. Canadian Journal of Zoology 94:291-299.


Hand, B.K., S. Chen, N. Anderson,  A. Beja-Pereira, P.C. Cross, M. Ebinger, H. Edwards, R.A. Garrott, M.D. Kardos, M. Kauffman, E.L. Landguth, A.D. Middleton, M. Schwartz, B. Scurlock, P.J. White, P. Zager, and G. Luikart. 2014. Limited maternal gene flow among elk populations in the Greater Yellowstone Ecosystem revealed by mitochondrial DNA. Journal of Fish and Wildlife Management 5:124-132.


Middleton, A.D., M.J. Kauffman, D.E. McWhirter, R.C. Cook, J.G. Cook, M.D. Jimenez, S.E. Albeke, H. Sawyer, and P.J. White. 2013. Linking antipredator behavior to prey demography reveals the limited risk effects of an actively hunting large carnivore. Ecology Letters 16:1023-1030.

Middleton, A.D., T.A. Morrison, J.K. Fortin, M.J. Kauffman, C.T. Robbins, K.M. Proffitt, P.J. White, D.E. McWhirter, T.M. Koel, D. Brimeyer, and W.S. Fairbanks. 2013. Grizzly bear predation links the loss of native trout to the demography of migratory elk in Yellowstone. Proceedings of the Royal Society B – Biological Sciences 280:20130870.

Middleton, A.D., M.J. Kauffman, D.E. McWhirter, J.G. Cook, R.C. Cook, A.A. Nelson, M.D. Jimenez, and R.W. Klaver. 2013. Rejoinder: challenge and opportunity in the study of ungulate migration amid environmental change. Ecology 94:1280-86.

Middleton, A.D., M.J. Kauffman, D.E. McWhirter, J.G. Cook, R.C. Cook, A.A. Nelson, M.D. Jimenez, and R.W. Klaver. 2013. Animal migration amid shifting patterns of predation and phenology: Lessons from a Yellowstone elk herd. Ecology 94:1245-56.

Sawyer, H. M.J. Kauffman, A.D. Middleton, T.A. Morrison, R.M. Nielson, and T.B. Wyckoff. 2013. A framework for understanding barrier effects on migratory ungulates. Journal of Applied Ecology 50:68-78.

Christianson, D., R.W. Klaver, A.D. Middleton, and M.J. Kauffman. 2013. Confounded winter and spring conditions on temperate herbivore ranges. Landscape Ecology 28:427-437.


Nelson, A.A., M.J. Kauffman, A.D. Middleton, M.D. Jimenez, D.E. McWhirter, J.J. Barber, and K. Gerow. 2012. Elk migration patterns and human activity influence wolf habitat selection in the Greater Yellowstone Ecosystem. Ecological Applications 22:2293-2307.

Cross, P.C., E.K. Cole, A.P. Dobson, W.H. Edwards, K.L. Hamlin, G. Luikart, A.D. Middleton, B.M. Scurlock, and P.J. White. 2010. Probable causes of increasing elk brucellosis in the Greater Yellowstone Ecosystem. Ecological Applications 20:278-288.

Hohl, A., C. Picard, S.G. Clark, and A.D. Middleton. 2010. Approaches to large-scale conservation: a survey. In Large Scale Conservation in the Common Interest, eds. S.G. Clark, A. Hohl, C. Picard, and D. Newsome. Springer. 191 pp.

Pauli, J.N., J.P. Whiteman, M. Riley, and A.D. Middleton. 2009. Defining “noninvasive” for sampling of vertebrates. Conservation Biology 24:349-352.

Other Publications

Middleton, A.D., T. Stoellinger, D.E. Bennett, T. Brammer, L. Gigliotti, H. Byerly Flint, S. Maher, and B. Leonard. 2022. The role of private lands in conserving Yellowstone's wildlife in the Twenty-First Century. Wyoming Law Review 22:237-302.

Middleton, A.D.  and Brashares, J. 2020. More than twice the size of Texas. The New York Times. (op-ed).

Stoellinger, T., H. J. Albers, A.D. Middleton, J.F. Shogren and R. Bonnie. Where the deer and the antelope play: conserving the big game migrations of the West. 31 Duke Environmental Law and Policy Forum 81-161 (2020).

Middleton, A.D. 2019. Tracking Wolves, pp. 179-187 in "Matthew Barney, Redoubt" by Yale University Art Gallery, Yale University Press, New Haven, CT. 368 pp.

Middleton, A.D. 2018. Protecting America's last great animal migrations. The New York Times (op-ed).

Middleton, A.D. 2017. Sustaining migrations in the modern West. In “Yellowstone Migrations by J. Riis. Pp 119-124.

Middleton, A.D. and L. Allison. 2016. Beyond boundaries in the Greater Yellowstone Ecosystem: scientists and ranchers exploring solutions for the future. Western Landowners Alliance. 28pp.

Middleton, A.D. 2016. Seeing Yellowstone: Science at its limits in a conservation crucible. pp. 36-51 in “Invisible Boundaries: Exploring Yellowstone’s Great Animal Migrations” ed. Karen B. McWhorter. Buffalo Bill Center of the West, Cody, WY. 80 pp.

Middleton, A.D. 2016. “Putting the prey back in predator-prey interactions” (Book review). Quarterly Review of Biology 91: 366.

Middleton, A.D. 2015. “Living with large carnivores: do ecology and conservation biology have the answers? (Book review). Ecology 96: 880-81.

Middleton, A.D. 2014. "Changes in habitat quality and predation shape a Yellowstone elk migration" and "Cascading consequences of the lake trout invasion on Yellowstone's migratory elk?" Yellowstone Science 22:36-37.

Martínez del Rio, C. and A.D. Middleton. 2010. Laws for ecology? (Book review). Ecology 91:1244-1245.