Yellowstone National Park

Gray Wolf (Canis lupus)

Northwestern Gray Wolf (Canis lupus occidentalis)
Northern Rocky Mountain Wolf (Canis lupus irremotus)

 Ecology /Prey

Ecology – Greater Yellowstone Ecosystem

Effect on Quaking Aspen (Populus tremuloides Michx.), Willow (Salix spp.) , Cottonwood (Populus spp.).

Birds – Raven (Corvus corax)

Effected by Global Warming, Mountain Pine Beetle, Fire, Hunting.


Red Deer “Rocky Moutain Elk” (Cervus elaphus),  Elk “Wapiti” (Cervus canadensis),  Bison “American Buffalo” (Bison bison), Mule Deer (Odocoileus hemionus), Moose (Alces alces), Beaver (Castor canadensis)

Non-Prey – Coyote (Canis latrans), grizzly bear, black bear, and cougar

Yellowstone Wolves  / National Park Service
Yellowstone 24-1 Wolf Facts
Yellowstone Science 24-1 Celebrating 20 Years of Wolves
Yellowstone Wolf Project Citizen Science
Wolf Restoration in Yellowstone National Park
Yellowstone Wolf Project Annual Reports

Journal Articles

The Spatial Ecology of Predator-Prey Interactions: A Case Study of Yellowstone Elk, Wolves, and Cougars. Kohl, M.T., 2019.

Estimating Detection for Gray Wolf (Canis lupus) Pups in Yellowstone National Park. Cassidy, B., 2019. University of Montana

Population responses of common ravens to reintroduced gray wolves
LE Walker, JM Marzluff, MC Metz… – Ecology and …, 2018 – Wiley Online Library

Sexually dimorphic aggression indicates male gray wolves specialize in pack defense against conspecific groups
KA Cassidy, LD Mech, DR MacNulty, DR Stahler, DW Smith – Behavioural Processes, 2017 – Elsevier

Wolf dispersal in the Rocky Mountains, Western United States: 1993–2008. Jimenez, M.D. E.E. Bangs, D.K. Boyd, D.W. Smith, S.A. Becker, D.E. Ausband, S.P. Woodruff, E.H. Bradley, J. Holyan, & K. Laudon. 2017. The Journal of Wildlife Management

Behavioral and ecological implications of seasonal variation in the frequency of daytime howling by Yellowstone wolves
R McIntyre, JB Theberge, MT Theberge, DW Smith; – Journal of Mammalogy, 2017

Competition between apex predators? Brown bears decrease wolf kill rate on two continents.Tallian, A., A. Ordiz, M.C. Metz, C. Milleret, C. Wikenros, D.W. Smith, D.R. Stahler, J. Kindberg, D.R. MacNulty, P. Wabakken, & J.E. Swenson. 2017.  Proceedings of the Royal Society

Predator foraging response to a resurgent dangerous prey. Tallian, A., D.W. Smith, D.R. Stahler, M.C. Metz, R.L. Wallen, C. Geremia, J. Ruprecht, C.T. Wyman, & D.R. MacNulty. 2017. Functional Ecology

Selecting habitat to what purpose? The advantage of exploring the habitat–fitness relationship. Uboni, A., D.W. Smith, D.R. Stahler, & J.A.Vucetich. 2017.  Ecosphere

Do gray wolves (Canis lupus) support pack mates during aggressive inter-pack interactions?
KA Cassidy, RT McIntyre – Animal cognition, 2016 – Springer

Implications of Harvest on the Boundaries of Protected Areas for Large Carnivore Viewing Opportunities; Bridget L. Borg, Stephen M. Arthur, Nicholas A. Bromen, Kira A. Cassidy, Rick McIntyre, Douglas W. Smith, Laura R. Prugh; PLOS; Published: April 28, 2016

Negative­-assortative mating for color in wolves. Hedrick, P.W., D.W. Smith, and D.R. Stahler. 2016.  Evolution

Managing wolves in the Yellowstone area: bal­ancing goals across jurisdictional boundaries. Smith, D.W., P.J. White, D.R. Stahler, A. Wydeven, and D.E. Hallac. 2016. Wildlife Soci­ety Bulletin

Group composition effects on aggressive interpack interactions of gray wolves in Yellowstone National Park; Kira A. Cassidy, Daniel R. MacNulty, Daniel R. Stahler, Douglas W. Smith, L. David Mech, , Behavioral Ecology, Volume 26, Issue 5, September-October

Infanticide in wolves: seasonality of mortalities and attacks at dens support evolution of territoriality, Douglas W. Smith, Matthew C. Metz, Kira A. Cassidy, Erin E. Stahler, Richard T. McIntyre, Emily S. Almberg, Daniel R. Stahler,  Journal of Mammalogy, Volume 96, Issue 6, 24 November 2015

Group composition effects on aggressive interpack interactions of gray wolves in Yellowstone National Park
KA Cassidy, DR MacNulty, DR Stahler… – Behavioral …, 2015 –

Density-dependent intraspecific aggression regulates survival in northern Yellowstone wolves (Canis lupus). Cubaynes, S., D.R. MacNulty, D.R. Stahler, K.A. Quimby, D.W. Smith, and T. Coulson. 2014.  Journal of Animal Ecology

Trophic cascades in a multicausal world: Isle Royale and Yellowstone. Annual Review of Ecology, Peterson, R.O., J.A. Vucetich, J.M. Bump, and D.W. Smith. 2014. Evolution, and Systematics

Influence of group size on the success of wolves hunting bison. MacNulty, D.R., A. Tallian, D.R Stahler, and D.W. Smith. 2014.  PLoS ONE

Hetero-zygote advantage in a finite population: black color in wolves. Hedrick, P.W., D.R. Stahler, and D. Dekker. 2014. Journal of Heredity

-Temporal variation in wolf predation dynamics in the multi-prey system of northern Yellowstone National Park
MC Metz, DW Smith, M Hebblewhite… – PeerJ PrePrints, 2016 –

[HTML] The effect of human activities and their associated noise on ungulate behavior
CL Brown, AR Hardy, JR Barber, KM Fristrup… – PloS one, 2012 –

Group composition effects on inter-pack aggressive interactions of gray wolves in Yellowstone National Park
KA Cassidy – 2013 –

Density‐dependent intraspecific aggression regulates survival in northern Yellowstone wolves (Canis lupus)
Sarah Cubaynes, Daniel R. MacNulty, Daniel R. Stahler, Kira A. Quimby, Douglas W. Smith, Tim Coulson – Journal of Animal …, 2014 – Wiley Online Library

Native prey distribution and migration mediates wolf (Canis lupus) predation on domestic livestock in the Greater Yellowstone Ecosystem
AA Nelson, MJ Kauffman… – … Journal of Zoology, 2016 – NRC Research Press

… Modeling Exploration of Risk Factors Associated with Sarcoptic Mange Severity, Susceptibility, and Duration in Yellowstone National Park Grey Wolves (Canis lupus)
S Wu – 2016 –

Social Status, Epigenetic Variation, and Complex Phenotypes: How Dominance Hierarchies Affect Plastic DNA Methylation in the Yellowstone Gray Wolf (Canis lupus)
E McCoy – 2014 –

Indirect effects and traditional trophic cascades: a test involving wolves, coyotes, and pronghorn. Berger KM, Gese EM, Berger J. Ecology. 2008 Mar

Restoring Yellowstone’s aspen with wolves. Ripple WJ, Beschta RL. Biological Conservation. 2007 Sep

Landscape heterogeneity shapes predation in a newly restored predator–prey system. Kauffman, Matthew J., Nathan Varley, Douglas W. Smith, Daniel R. Stahler, Daniel R. MacNulty, and Mark S. Boyce. Ecology letters 10, no. 8 (2007)

Foraging and feeding ecology of the gray wolf (Canis lupus): lessons from Yellowstone National Park, Wyoming, USA. Stahler DR, Smith DW, Guernsey DS. The Journal of nutrition. 2006 Jul

Habitat selection by elk before and after wolf reintroduction in Yellowstone National Park. Mao, Julie S., Mark S. Boyce, Douglas W. Smith, Francis J. Singer, David J. Vales, John M. Vore, and Evelyn H. Merrill. The Journal of Wildlife Management 69, no. 4 (2005)

Trophic facilitation by introduced top predators: grey wolf subsidies to scavengers in Yellowstone National Park. Wilmers CC, Crabtree RL, Smith DW, Murphy KM, Getz WM.  Journal of Animal Ecology. 2003 Nov 1

Yellowstone after wolves; DW Smith, RO Peterson, DB Houston – BioScience, 2003 –

Assessing the impact of wolves on ungulate prey. Eberhardt LL, Garrott RA, Smith DW, White PJ, Peterson RO. Ecological Applications. 2003 Jun

Leadership behavior in relation to dominance and reproductive status in gray wolves, Canis lupus; RO Peterson, AK Jacobs, TD Drummer… – … Journal of Zoology, 2002 – NRC Research Press

Trophic cascades among wolves, elk and aspen on Yellowstone National Park’s northern range. Ripple, William J., Eric J. Larsen, Roy A. Renkin, and Douglas W. Smith.  Biological conservation 102, no. 3 (2001)

The ungulate prey base for wolves in Yellowstone National Park. The Greater Yellowstone Ecosystem: Redefining America’s Wilderness Heritage. Singer FJ. 1991: