Arthur D. Middleton
Fall 2015
CURRENT POSITION
Associate research scientist
Yale School of Forestry and Environmental Studies
Cell: (307) 460-0880
370 Prospect Street
E-mail: arthur.middleton@yale.edu
New Haven, CT 06511
Web: www.environment.yale.edu/profile/arthur-middleton
EDUCATION
University of Wyoming, Ph.D.
Yale University, M.E.M.
Bowdoin College, B.A.
Ecology
Ecology, ecosystems and biodiversity
English literature, government
2007-12
2005-07
1997-2003
FELLOWSHIPS
Postdoctoral Fellow, Yale Institute for Biospheric Studies
Gaylord Donnelley Postdoctoral Fellow, Yale Institute for Biospheric Studies
Resident Fellow, Buffalo Bill Center of the West, Cody, WY
Doris Duke Conservation Fellow, Yale University
Thomas J. Watson Fellow, Thomas J. Watson Foundation
2014-15
2012-14
2009-10
2006-07
2003-04
SELECTED ACADEMIC PUBLICATIONS
Nelson, A.A., M.J. Kauffman, A.D. Middleton, M.D. Jimenez, D.E. McWhirter, and K. Gerow. In review.
Native prey distribution and migration mediates wolf predation on domestic livestock in the Greater
Yellowstone Ecosystem.
Legler, S.B., A.D. Middleton, M.J. Kauffman, K.G. Gerow, and D.B. Tinker. In review. Plant community
composition and nutritional values on the seasonal ranges of a partially migratory Yellowstone elk
herd: natural variation and agricultural subsidies.
Miller, J.R.B., K.J. Stoner, M.R. Cejtin, T.K. Meyer, A.D. Middleton, and O.J. Schmitz. In review. Tools for
human-carnivore coexistence: A quantitative assessment of techniques for reducing large carnivore
depredation of livestock.
Middleton, A.D., D.K. Skelly, D.M. MacNulty, A.T. Ford, O.J. Schmitz, and M.J. Sheriff. In revision.
Predator-prey encounter rates, lethality, and the challenge of predicting risk effects in diverse
vertebrates.
Middleton, A.D., M. Hebblewhite, L.R. Prugh, A. Treves, M. Ogada, J. Maron, S.G. Clark, D.T. Blumstein,
J.S. Brashares, D. Fortin, N.T. Hobbs, G. Hopcraft, S. Hussain, M.J. Kauffman, M. Letnic, D.J.
Mattson, K.L. Parker, D.W. Smith, C.C. Wilmers, and O.J. Schmitz. In revision. Characterizing
coexistence landscapes to strengthen the links between ecological science and large carnivore
conservation.
Lukacs, P.M., M. Hebblewhite, B.K. Johnson, H. Johnson, M.J. Kauffman, K.M. Proffitt, P. Zager , J. Brodie,
K. Hersey, A.A. Holland, M. Hurley, S. McCorquodale, A.D. Middleton, M.S. Mitchell, M.
Nordhagen, J.J. Noway, D.P. Walsh, P.J. White. In review. Declines in juvenile elk recruitment
across the western United States are associated with recolonizing predators, habitat and climate
conditions.
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. Media
coverage by ScienceNOW, Casper Star-Tribune, Jackson Hole News and
Guide, Missoulian, Bozeman Chronicle, Wildlife Professional.
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. 2013. Limited maternal gene flow among elk populations in the Greater
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Arthur D. Middleton
Fall 2015
Yellowstone Ecosystem revealed by mitochondrial DNA. Journal of Fish and Wildlife
Management.
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. Media coverage by Discover Magazine, Nature
Research Highlights, ScienceNOW, Billings Gazette, Casper Star-Tribune, Jackson Hole News and
Guide, Bozeman Chronicle, Conservation Magazine, Agence France-Presse, Le
Monde, Spiegel, High Country News, Western Confluence.
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. Media coverage by Connecticut Public
Radio (audio), Wyoming Public Radio's Open Spaces (audio), Billings Gazette, Casper StarTribune, Jackson Hole News and Guide, San Francisco Chronicle.
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: Integrating Science, Management, and Policy in the Common
Interest, eds. S.G. Clark, A. Hohl, C. Picard, and D. Newsome. Yale School of Forestry and
Environmental Studies Publication Series. 276 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.
ACADEMIC MANUSCRIPTS IN PREPARATION
Perrig, P., E. Donadio, A.D. Middleton, and J.N. Pauli. Predation subsidizes an obligate scavenger in the high
Andes.
Middleton, A.D., P.J. White, M. Hayes, M.J. Kauffman, P. Atwood, J. Beckmann, D. Brimeyer, E. Cole, A.
Courtemanch, S. Dewey, D. McWhirter, and K. Proffitt. Migration links wilderness to private
agricultural lands on an ecosystem frontier.
Sawyer, H. and A.D. Middleton. Variable migration distances in a single population – implications for
migratory ungulates.
MacNulty, D.R., Kindermann, R., D.R. Stahler, A.D. Middleton, M.J. Kauffman, D.E. McWhirter and D.W.
Smith. Reproduction in Greater Yellowstone elk: a search for wolf-caused risk effects.
NON-ACADEMIC PUBLICATIONS
Middleton, A.D. Living with large carnivores: do ecology and conservation biology have the answers?
Ecology (book review) 96: 880-81.
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Fall 2015
Middleton, A.D. 2015. San Guillermo Expedition 2015. National Geographic Explorer’s Journal (blog series
on puma-vicuna interactions).
Middleton, A.D. 2014. San Guillermo Expedition 2014. National Geographic Explorer’s Journal (blog series
on puma-vicuna interactions).
Middleton, A.D. "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.
Middleton, A.D. 2014. Is the wolf a real American hero? The New York Times (op-ed). p. A21. Media
coverage by Popular Science.
Middleton, A.D. 2012. As wolves return to the West, greens go to court (op-ed). The Wall Street Journal. p.
A11.
Martínez del Rio, C. and A.D. Middleton. 2010. Laws for ecology? (book review). Ecology 91:1244-1245.
SELECTED PRESENTATIONS
Middleton, A.D. Re-discovering the migrations of the Greater Yellowstone Ecosystem. National Geographic.
Invited presentation (“Closer Look”) with J. Riis, Washington, DC, October 2015.
Middleton, A.D. Bridging science and the arts to tell the story of Yellowstone’s wildlife migrations. The
Nature Conservancy, Wyoming Chapter. Invited presentation, board meeting, Cody, Wyoming,
June 2015.
Middleton, A.D. Bridging science and the arts to tell the story of Yellowstone’s wildlife migrations. Ucross
Foundation Grasslands Symposium. Invited presentation, Ucross, Wyoming, May 2015.
Middleton, A.D. Wildlife migration and the future of Yellowstone. Greater Yellowstone Coordinating
Committee. Invited presentation, Cody, Wyoming, April 2015.
Middleton, A.D. Ecology and conservation of animal migration in the Greater Yellowstone Ecosystem.
University of Massachusetts – Amherst, Environmental Conservation Seminar Series. Invited
presentation, Amherst, Massachusetts, April 2015.
Middleton, A.D. and J. Prosek. Invisible boundaries: Wildlife migrations and the future of Yellowstone. Yale
Institute for Biospheric Studies, External Advisory Board, New Haven, CT, April 2015.
Middleton, A.D. Large carnivores and their prey in the Greater Yellowstone Ecosystem: ecology in a
conservation crucible. Vargsymposiet (The Wolf Symposium). Invited foreign speaker, Vålådalen,
Sweden, March 2015.
Middleton, A.D. Conveying Yellowstone’s changing ecological interactions to the public. Vargsymposiet
(The Wolf Symposium). Invited foreign speaker, Vålådalen, Sweden, March 2015.
Middleton, A.D. Ecology and conservation of animal migration in the Greater Yellowstone Ecosystem.
Department of Biology, Queens College at the City College of New York. Invited presentation,
New York, New York, November 2014.
Middleton, A.D. Imagining and re-imagining mountain ecosystems through linked science and storytelling.
National Geographic. Invited presentation (“Closer Look”), Washington, DC, October 2014.
Middleton, A.D. Re-discovering the elk migrations of the Greater Yellowstone Ecosystem: linking science
and outreach. 12th Yellowstone Science Conference. Mammoth, WY, October 2014.
Middleton, A.D. Animal migration as a basis for understanding and conserving western landscapes. U.S. Fish
and Wildlife Service Director’s Office. Invited presentation, Washington, DC, September 2014.
Middleton, A.D. Migration of the Cody elk herd. Upper South Fork Landowners’ Association Annual
Meeting. Invited presentation, Cody, WY, August 2014.
Middleton, A.D. Leaps and bounds: a new era of research and conservation for migratory wildlife in the West
Wildlife Heritage and Hunting Conservation Council (convened by the U.S. Fish and Wildlife
Service). Invited presentation, Cody, WY, June 2014.
Middleton, A.D. The changing ecology of elk migration in the Greater Yellowstone Ecosystem. Invited
seminar, Universite de Sherbrooke, Sherbrooke, Quebec. April 2014.
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Fall 2015
Middleton, A.D. Large carnivores, drought, and the changing ecology of elk migration in Yellowstone.
Invited seminar, Conservation Ecology series, University of Michigan School of Environment and
Natural Resources, Ann Arbor, MI, January 2014.
Middleton, A.D. A tale of two predators. Invited public lecture, Real Artways, Science on Screen series,
Hartford, CT. September 2013.
Middleton, A.D., C. Martinez del Rio, and M.J. Kauffman. Prey body condition: a common influence on
antipredator behavior and the “ecology of fear?” Invited symposium presentation, Annual Meeting
of the Ecological Society of America, Minneapolis, MN, August 2013.
Middleton, A.D. The changing ecology of elk migration in the Greater Yellowstone Ecosystem. Invited public
lecture, University of Wyoming – National Park Service Research Station, Grand Teton National
Park, WY, June 2013.
Middleton, A.D. Can wolves influence elk and plants via fear alone? Wolf-elk interactions in Wyoming’s
Absaroka Mountains. Invited public lecture (co-sponsored by Wyoming Wildlife Federation and
Wyoming Game and Fish Department), Teton County Library, Jackson, WY, June 2013.
Middleton, A.D. Large mammals in landscapes of fear. Yale Institute for Biospheric Studies Seminar Series,
New Haven, CT, February 2013.
Middleton, A.D., M.J. Kauffman, D.E. McWhirter, M.D. Jimenez, R.C. Cook, J.G. Cook, S.E. Albeke, H.
Sawyer, and P.J. White. Spatial and temporal context limit the behavioral and physiological
responses of Yellowstone elk to wolves. Annual Meeting of the Wildlife Society, Portland, OR,
October 2012; GYE Science Conference, Mammoth, WY, October 2012; Annual Meeting of the
Ecological Society of America, Portland, OR, August 2012.
Middleton, A.D. Predation, climate, and elk migration in Yellowstone. Invited seminar, Department of Forest
and Wildlife Ecology, University of Wisconsin-Madison, September 2011.
Middleton, A.D. Changing times in Wyoming elk country: Large carnivores, drought, and elk migration.
Rocky Mountain Elk Foundation, Annual Habitat Council Retreat, Cody, WY, June 2011.
Middleton, A.D. The ecology of elk migration in the Greater Yellowstone Ecosystem. Invited seminar,
Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata, Argentina,
March 2011.
Middleton, A.D. Large carnivore recovery and severe drought reduce the benefits of migration for a
Yellowstone elk herd. Invited presentation, Faculty Seminar Series, Yale School of Forestry and
Environmental Studies, New Haven, CT, October 2010.
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. Large carnivore recovery and severe drought reduce the benefits of migration for a
Yellowstone elk herd. Annual Meetings of the Ecological Society of America, Pittsburgh, PA, and
the American Society of Mammalogists, Laramie, WY. June 2010.
Middleton, A.D. Drought, predation, and migratory Clarks Fork elk. Several invited talks for the Wyoming
Game and Fish Department's Cody All-Region Meeting and Terrestrial Habitat Section Annual
Meeting and the U.S. Fish and Wildlife Service Wolf Recovery Annual Meeting. Cody and Kelly,
WY and West Yellowstone, MT. May-June 2010.
Middleton, A.D. Drought, predation, and migratory Clarks Fork elk. Invited public presentations at the
Buffalo Bill Center of the West in Cody, WY and Teton County Library in Jackson, WY. May
2010.
SELECTED GRANTS AND AWARDS
2015
2015
Fran and Lenox Baker Foundation. “Invisible Boundaries: The Seasonal Journeys of
Yellowstone’s Wildlife” museum exhibition. With Buffalo Bill Center of the West, C.
Preston, K. McWhorter, J. Riis, J. Nichols, J. Prosek. $200,000.
Bole and Klingenstein Foundation. “Elk River” film. $95,200. With J. Riis, J. Nichols, J.
Prosek.
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Arthur D. Middleton
2015
2015
2015
2015
2015
2015
2014
2014
2014
2014
2013
2013
2013
2013
2013
2013
2013
2013
2012
2012
2012
2012
Fall 2015
Theodore C. Rogers. “Invisible Boundaries: The Seasonal Journeys of Yellowstone’s
Wildlife” museum exhibition. With Buffalo Bill Center of the West, C. Preston, K.
McWhorter, J. Riis, J. Nichols, J. Prosek. $30,000.
Duncan Fund. Re-discovering the elk migrations of the Greater Yellowstone Ecosystem: a
project of trans-boundary science and outreach. With J. Riis. $15,000.
Mary Anne and Bill Dingus. Re-discovering the elk migrations of the Greater Yellowstone
Ecosystem: a project of trans-boundary science and outreach. With J. Riis. $15,000.
Wyoming Outfitters and Guides Association. Wiggins Fork elk migration project. $3,000.
Cody Country Outfitters and Guides Association. Wiggins Fork elk migration project.
$3,000.
U.S. Forest Service, Shoshone National Forest. Wiggins Fork elk migration project.
$15,000.
Devonwood Foundation. A landscape of fear in the high Andes: Evaluating puma-vicuna
interactions at San Guillermo National Park, Argentina. $20,000.
Knobloch Family Foundation. Re-discovering the Greater Yellowstone Ecosystem through
its animal migrations. $250,000.
George B. Storer Foundation. Re-discovering the elk migrations of the Greater Yellowstone
Ecosystem: a project of trans-boundary science and outreach. With J. Riis. $60,000.
Fran and Lenox Baker Foundation. Re-discovering the elk migrations of the Greater
Yellowstone Ecosystem: a project of trans-boundary science and outreach. With J. Riis.
$25,000.
National Geographic Society, Expeditions Council. Re-discovering the elk migrations of
the Greater Yellowstone Ecosystem. $22,000. With J. Riis.
Fran and Lenox Baker Foundation. Re-discovering the elk migrations of the Greater
Yellowstone Ecosystem: a project of trans-boundary science and outreach. With J. Riis.
$25,000.
Knobloch Family Foundation. Re-discovering the elk migrations of the Greater
Yellowstone Ecosystem: a project of trans-boundary science and outreach. With J. Riis.
$40,000.
George B. Storer Foundation. Re-discovering the elk migrations of the Greater Yellowstone
Ecosystem: a project of trans-boundary science and outreach. With J. Riis. $40,000.
Camp Monaco Prize (Buffalo Bill Historical Center and University of Wyoming
Biodiversity Institute). Re-discovering the elk migrations of the Greater Yellowstone
Ecosystem: a project of trans-boundary science and outreach. With J. Riis. $100,000.
Devonwood Foundation. A landscape of fear in the high Andes: Evaluating puma-vicuna
interactions at San Guillermo National Park, Argentina. $20,000.
Association of Zoos and Aquariums, Conservation Endowment Fund. Evaluating the
importance of native prey species to the diet of Andean condors in the high Andes of
northwestern Argentina. $19,400. With J. Pauli, P. Perrig, E. Donadio.
National Geographic Society, Committee for Research and Exploration. A landscape of fear
in the high Andes: Evaluating puma-vicuna interactions at San Guillermo National Park,
Argentina. $25,400.
George B. Storer Foundation. Evaluating impacts of disturbance on wildlife. $20,000.
Wyoming Game and Fish Department. Evaluating nutritional condition of mule deer in the
Pinedale Anticline Project Area. With K. Monteith, M. Kauffman, H. Sawyer. $60,000.
Boone and Crockett Club. Fitness consequences of energy development: understanding and
reducing the human footprint on ungulate populations. With K. Monteith, H. Sawyer, M.
Kauffman. $15,500.
Rocky Mountain Elk Foundation. What drives variation in elk pregnancy rates? With J.
Rodgerson, B. Scurlock, M. Kauffman. $10,000.
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Arthur D. Middleton
2012
2012
2011
2008-11
2008-11
2008-11
2009-10
2008-09
2008-09
Fall 2015
Wyoming Governor’s Big Game License Coalition. What drives variation in elk pregnancy
rates? With J. Rodgerson, B. Scurlock, M. Kauffman. $5,000.
Devonwood Foundation. A landscape of fear in the high Andes: Evaluating puma-vicuna
interactions at San Guillermo National Park, Argentina. $20,000.
National Birds of Prey Trust. Evaluating the importance of native prey species to Andean
condors in the high Andes of Argentina. £6,413. With E. Donadio, J. Pauli.
Wyoming Animal Damage Management Board. Absaroka Elk Ecology Project. With M.
Kauffman, D. McWhirter, M. Jimenez. $111,750.
Rocky Mountain Elk Foundation. Absaroka Elk Ecology Project. With M. Kauffman, D.
McWhirter, M. Jimenez. $47,500.
Wyoming Governor’s Big Game License Coalition. Absaroka Elk Ecology Project. With
M. Kauffman, D. McWhirter, M. Jimenez. $14,000.
Wyoming Game and Fish Department. Elk-wolf-habitat relations. With M. Kauffman, D.
McWhirter, M. Jimenez. $80,000.
U.S. Forest Service. Absaroka Elk Ecology Project. $6,000.
University of Wyoming National Park Service Research Station. Absaroka Elk Ecology
Project. $5,000.
PROFESSIONAL AFFILIATIONS
Ecological Society of America, The Wildlife Society
REVIEWER
American Naturalist, Behavioral Ecology and Sociobiology, Biological Conservation, Ecology, Ecological
Applications, Ecosphere, Journal of Animal Ecology, Journal of Wildlife Management, Oikos, PLoS ONE,
Proceedings of the Royal Society B, Western North American Naturalist, Wildlife Biology, University of
Chicago Press, National Geographic Committee for Research and Exploration
OTHER SKILLS AND EXPERIENCE
-Wildlife chemical immobilization, Wyoming Game and Fish Department and Safe-Capture International
-Aviation safety, U.S. Department of Interior
-Coordination of helicopter and ground-trapping operations to capture large mammals
-Master-level falconer with professional experience training >20 raptor species for education and hunting
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