The Wolf Intelligencer


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.


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

Other Predators Coyote (Canis latrans), grizzly bear, black bear (Ursus americanus), cougar (Puma concolor)

Beautiful Birds of Yellowstone!
Trumpeter Swan (Cygnus buccinator), Common Raven (Corvus corax), Black-billed Magpie (Pica hudsonia), American Crow (Corvus brachyrhynchos), Canada goose (Branta canadensis), Red-necked Grebe (Podiceps grisegena), Dusky Grouse (Dendragapus obscurus), Red-breasted Nuthatch (Sitta canadensis)

More Birds

American Dipper (Cinclus mexicanus), Calliope Hummingbird (Selasphorus calliope), Black-bellied Plover (Pluvialis squatarola), Sanderling (Calidris alba), American White Pelican (Pelecanus erythrorhynchos), Osprey (Pandion haliaetus), Golden Eagle (Aquila chrysaetos), Great Horned Owl (Bubo virginianus), Lewis’s Woodpecker (Melanerpes lewis), American Kestrel (Falco sparverius), Peregrine Falcon (Falco peregrinus), Mountain Chickadee (Poecile gambeli), Black-rosy Finch (Leucosticte atrata), Red-winged Blackbird (Agelaius phoeniceus), Yellow Warbler (Setophaga petechia)

Yellowstone Wolves  / National Park Service
Yellowstone 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

The Wolf Intelligencer Book Library – Yellowstone

Gray Wolf Videos National Park Service
Yellowstone National Park Telemetry: Episode 5, The Value of One Wolf. Kira Cassidy
Sound Library – Yellowstone National Park


Wolf and Wildlife News from Yellowstone

Journal Articles

Key observations of flexed-leg urination in the free-ranging Gray Wolf (Canis lupus) D Mech, R McIntyre – The Canadian Field-Naturalist, 2022


Flexed-leg urination (FLU) in female Gray Wolves (Canis lupus) has been little studied in the wild. Captive females in packs do not exhibit FLU unless they are both mature and dominant to an associate female, but these characteristics have not been confirmed in free-ranging wolves. We present observations of wolves in Yellowstone National Park that accord with those of wolves in captivity, extend our knowledge of FLU in Gray Wolf, pose additional questions about it, and suggest new areas of study to better understand it.

Intended and unintended consequences of wolf restoration to Yellowstone and Isle Royale National Parks. Smith DW, Peterson RO.Conservation Science and Practice. 2021 Apr

How climate impacts the composition of wolf killed‐elk in northern Yellowstone National Park. Wilmers CC, Metz MC, Stahler DR, Kohl MT, Geremia C, Smith DW. Journal of Animal Ecology. 2020 Mar

Heritability of inter‐pack aggression in a wild pedigreed population of North American gray wolves. vonHoldt BM, DeCandia AL, Heppenheimer E, Janowitz‐Koch I, Shi R, Zhou H, German CA, Brzeski KE, Cassidy KA, Stahler DR, Sinsheimer JS. Molecular Ecology. 2020 Jan

Do prey select for vacant hunting domains to minimize a multi‐predator threat?. Kohl MT, Ruth TK, Metz MC, Stahler DR, Smith DW, White PJ, MacNulty DR. Ecology letters. 2019 Nov

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.,. University of Montana 2019

Wetland drying linked to variations in snowmelt runoff across Grand Teton and Yellowstone national parks. Ray AM, Sepulveda AJ, Irvine KM, Wilmoth SK, Thoma DP, Patla DA. Science of the Total Environment. 2019 May

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

A Novel Approach to Assessing Abundance and Behavior in Summer Populations of Little Brown Myotis in Yellowstone National Park. Waag, A.G., 2018.

Wolves for Yellowstone: dynamics in time and space. Boyce MS. Journal of Mammalogy. 2018 Sep

Historical Landscape Change in Yellowstone National Park: Demonstrating the Value of Intensive Field Observation and Repeat Photography. Meyer JL, Youngs Y. Geographical Review. 2018 Jul

How perceptions about naturalness affect science in Yellowstone National Park. Keigley RB. Rangeland ecology & management. 2018 May

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

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. The Journal of Wildlife Management 2017

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.  Proceedings of the Royal Society 2017

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. Functional Ecology 2017

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

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

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: 2016 Apr

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

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.. Wildlife Soci­ety Bulletin 2016 Sept

Compassionate Conservation for Yellowstone’s Wolves. Povilitis T. Natural areas journal. 2016 Jul

Park visitor support for cross-boundary protection of Yellowstone wolves. Povilitis T. Hum. Dimens. Wildlife. 2016 Jan

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

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

Riparian vegetation recovery in Yellowstone: the first two decades after wolf reintroduction. Beschta RL, Ripple WJ. Biological Conservation. 2016 Jun

-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

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, Sep

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..  Journal of Animal Ecology 2014

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. . Evolution, and Systematics 2014

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

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

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

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

Is science in danger of sanctifying the wolf?. Mech LD. Biological Conservation. 2012 Jun

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

Seasonal patterns of predation for gray wolves in the multi‐prey system of Yellowstone Metz MC, Smith DW, Vucetich JA, Stahler DR, Peterson RO. National Park. Journal of Animal Ecology. 2012 May

Scavenging: how carnivores and carrion structure communities. Wilson EE, Wolkovich EM. Trends in Ecology & Evolution. 2011 Mar

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

The genealogy and genetic viability of reintroduced Yellowstone grey wolves. Vonholdt BM, Stahler DR, Smith DW, Earl DA, Pollinger JP, Wayne RK. Molecular ecology. 2008 Jan

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

Evaluating prey switching in wolf–ungulate systems.  Garrott RA, Bruggeman JE, Becker MS, Kalinowski ST, White PJ. Ecological Applications. 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 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

Influence of harvest, climate and wolf predation on Yellowstone elk, 1961‐2004. Vucetich JA, Smith DW, Stahler DR. Oikos. 2005 Nov

Habitat selection by elk before and after wolf reintroduction in Yellowstone National Park. Mao JS, Boyce MS, Smith DW, Singer FJ, Vales DJ, Vore JM, Merrill EH. The Journal of Wildlife Management. 2005 Oct

Yellowstone’s ungulates after wolves–expectations, realizations, and predictions. White PJ, Garrott RA. Biological Conservation. 2005 Sep

Adaptive management for reintroductions: updating a wolf recovery model for Yellowstone National Park. Varley N, Boyce MS. Ecological Modelling. 2006 Mar

Winter prey selection and estimation of wolf kill rates in Yellowstone National Park, 1995–2000. Smith DW, Drummer TD, Murphy KM, Guernsey DS, Evans SB. The Journal of Wildlife Management. 2004 Jan

Large-carnivore response to recreational big-game hunting along the Yellowstone National Park and Absaroka-Beartooth Wilderness boundary. Ruth TK, Smith DW, Haroldson MA, Buotte PC, Schwartz CC, Quigley HB, Cherry S, Murphy KM, Tyers D, Frey K. Wildlife Society Bulletin. 2003 Dec

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

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

Wolves, elk, and bison: reestablishing the” landscape of fear” in Yellowstone National Park, USA. Laundré JW, Hernández L, Altendorf KB. Canadian Journal of Zoology. 2001 Aug

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 2001

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

The public and wolf reintroduction in Yellowstone National Park. Bath AJ. Society & Natural Resources. 1989 Jan

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