Scott Creel

General Areas of Interest: Population biology, behavioral ecology, conservation, behavioral endocrinology, evolutionary ecology. Virtually all of my research is based on field studies, generally using observational rather than experimental methods, and often following known individuals. Much of the work in my lab involves the integration of behavioral and demographic data from the field with endocrine and genetic data from the lab. My lab is equipped for extraction and radioimmunoassay and enzyme-linked immunosorbent assays of steroid hormones, DNA extraction, and PCR.


	My current research primarily examines responses of prey to the risk of predation. One part of this work examines the responses of elk to variation in the risk of predation by wolves. This work is based in four subsites within the Gallatin Canyon, bordering Yellowstone National Park on federal, state and private land. The goals of the project are: (1) To quantify the responses of elk to variation in predation risk. These include responses in behavior, feeding ecology, distribution, and patterns of aggregation. (2) To determine thecosts of these responses, physiologically and demographically (3) To determine the impact of these costs on the population dynamics of both elk and wolves. A second part of this work is on the Shompole & Olkiramatian Maasai Group Ranches in the Southern Rift Valley of Kenya. With Dr. Jonah Western of the African Conservation Centre, we are examining behavioral aspects of predator-ungulate dynamics and interactions with people and cattle inside and outside of a conservancy recently established by local initiative. This research will test the generality of conclusions from the wolf-elk project, and identify ramifications for ecosystem function, management and conservation in the wooded savannas of East Africa. My students and I are addressing these goals with a wide variety of methods, including behavioral observations, demographic monitoring, ground and aerial censuses,snow and dirt tracking, camera trapping, GPS and VHF radiotelemetry, ELISA of fecal steroids to measure pregnancy rates and stress responses, measurements of urinary allantoin:creatinine ratios to assess foraging success, measurements of abiotic factors such as weather and snow conditions, and assessments of forage quality by chemical and microhistological methods.

Wagner AP, Frank LG, Creel S 2008. Spatial grouping in behaviourally solitary striped hyenas (Hyaena hyaena) Animal Behaviour 75: 1131-1142. doi:10.1016/j.anbehav.2007.08.025

(PDF) Liley S & Creel S 2008. What best explains vigilance in elk: characteristics of prey, predators, or the environment? Behavioral Ecology 19: 245-254

(PDF) Creel S 2005. Dominance, aggression and glucocorticoid levels in social carnivores. Journal of Mammalogy 86:255-264.

Macdonald DW, Mills MGL & Creel S 2004. Canid Society. In: Biology and Conservation of Wild Canids, Ed by DW Macdonald and C Sillero-Zubiri, Oxford University Press, pp. 85-106

Lukins WJ, Creel S, Erbes B & Spong G 2004. An assessment of the Tobacco Root mountain range in Southwestern Montana as a linkage zone for grizzly bears. Northwest Science 78: 168-172.

(PDF) Creel, S, McNutt, JW & Mills, MGL 2004 Demography and Population Dynamics of African Wild Dogs in Three Critical Populations. In Biology and Conservation of Wild Canids, ed by DW Macdonald & C Sillero-Zubiri, Oxford University Press, pp. 337-350.

(PDF) Sands JL & Creel 2004. Social dominance, aggression and fecal glucocorticoid levels in a wild population of wolves, Canis lupus. Animal Behaviour 67: 387-396

(PDF) Spong G & Creel S. 2004. Effects of kinship on territorial conflicts among groups of lions, Panthera leo. Behavioral Ecology and Sociobiology 55:325-331

(PDF) Creel S, Spong G, Sands JL, Rotella J, Ziegle J, Joe L, Murphy KM & Smith D 2003. Population size estimation in Yellowstone wolves with error-prone noninvasive microsatellite genotypes. Molecular Ecology 12:2003-2009.

Creel S & Sands, JL 2003. Is social stress a consequence of subordination or a cost of dominance? In Animal Social Complexity, ed by F. de Waal and P. Tyack, pp. 153-179, Harvard University Press.

Creel S & Creel NM 2002. The African Wild Dog: Behavior, Ecology and Conservation. Princeton University Press, Princeton.

(PDF) Creel, S. Fox, JE, Hardy, A, Sands, J, Garrott, R, and Peterson, R. 2002. Snowmobile activity and glucocorticoid stress responses in wild wolves and elk. Conservation Biology 16:809-814
(Review of this work: Withgott, J. 2002. Science 296:1784-1785)

(PDF) Spong G, Creel S, Stone J & Bjorklund M. 2002 Genetic structure of a population of lions (Panthera leo): implications for the evolution of sociality. Journal of Evolutionary Biology 15(6):945-953.

(PDF) Creel S 2001. Social dominance and stress hormones. Trends in Ecology and Evolution 16: 491-497

Spong, G & Creel, S 2001. Deriving dispersal distances from genetic data. Proceedings of the Royal Society of London: Biological Sciences 268:2571-2574

Creel S 2001. Cooperative hunting and sociality in African wild dogs (Lycaon pictus). In: Model Systems in Behavioral Ecology Ed. by L. Dugatkin, pp. 466-490, Princeton University Press, Princeton.

Creel S, Spong G & Creel NM 2001. Interspecific competition and the population biology of extinction-prone carnivores. In Conservation of Carnivores. Ed by J. Gittleman, D. Macdonald, S. Funk and R Wayne, pp. 35-60. Cambridge University Press.

Girman DJ, Vila C, Geffen E, Creel S, Mills MGL, McNutt JW, Ginsberg J, Kat P & Wayne RK 2001. Patterns of population subdivision, gene flow, and genetic variability in the African wild dog, Lycaon pictus. Molecular Ecology. 10: 1703-1723

Creel S 2001. Four factors modifying the impact of competition on carnivore population dynamics, as illustrated by African wild dogs, Lycaon pictus. Conservation Biology 15: 74-79.

I encourage my graduate students to participate fully in developing their research questions and to pursue independent funding.

Paul Schuette (PhD): NSF GRA. Paul is studying interactions between top carnivores (lions, spotted hyenas, leopards), their competitors, their prey and humans, working on the Olkiramation and Shompole Maasai group ranches. Paul is cosupervised by Dr. Jonah Western of the African Conservation Centre.
Cecily Costello (PhD): Wildlife Conservation Society/Hornocker Wildlife Institute grants. Cecily is studying black bear social organization and space use as they relate to population genetics and patterns of relatedness, using data from two populations in NM.
Dave Christianson (PhD): NSF GK-12 Fellowship, NSF EPSCoR Fellowship, NSF GRA. Dave is examining changes in elk foraging behavior in response to the presence of wolves, and the impacts of these changes on elk diets, nutrition, and demography.
Leslie Frattaroli (MS): Leslie, co-advised by Dr. Chuck Schwartz, is using downloadable GPS collars to visit foraging sites of black bears soon after their use, to assess recreation impacts on habitat use and to assess the affects of grizzly bear range expansion on black bears. Her field work is with Steve Cain in Grand Teton National Park.
Tyler Coleman (MS): Tyler, co-advised by Dr. Chuck Schwartz, is conducting an evaluation of the effectiveness of Yellowstone National Park's grizzly bear management policies, focussing on the closure or restriction of use in bear management areas. His field work is with Kerry Gunther of Yellowstone National Park.

Stewart Liley (MS): Stewart used model selection methods to test the relative strength of predator, prey and environmental characteristics in predicting antipredator responses of elk to the presence of wolves. Surprisingly little prior work has attempted to determine the relative importance of these three types of variables in determining the strength of antipredator responses. Dangerous places? The size or proximity of predator groups? Characteristics of the prey group itself? Answers are in his paper in Behavioral Ecology. Stewart is now the head elk biologist for the state of New Mexico.
John Winnie (PhD): John studied the effects of predation risk from wolves on elk behavior, grouping patterns and spatial distributions, producing wide ranging data that revealed strong responses by elk in almost every aspect of their behavior that we considered. He went on to a postdoc with Wayne Getz and Paul Cross, studying habitat selection by African buffalo, and then to work for WCS on the conservation of argali (Marco Polo sheep) in the Wakhan corridor of Afghanistan.
Aaron Wagner (PhD): Aaron studied striped hyenas on the Laikipia Plateau of Kenya, including aspects of behavior, ecology, endocrinology and genetics. He went on to a postdoc to continue his work on striped hyenas in Kenya, with Kay Holekamp at Michigan State University (thereby keeping his career wholly within institutions called MSU).
Julia Nelson (MS), who studied the impacts of coyotes on the use of space and stress physiology of endangered San Joaquin kit foxes, working with Dr. Brian Cypher. She went on to do some world travelling, study languages and work in Belize for the Peace Corps.
Goran Spong (PhD), who studied population genetics and social evolution in African lions (in the Selous Game Reserve). Goran went on to a postdoc at Cambridge university with Dr. Tim Clutton Brock, studying meerkats, and then to a faculty position at the University of Umea.
Jennifer Sands (MS), who studied interactions between aggression, social status and glucocorticoid stress hormones in wolves (in Yellowstone National Park). Jennifer went on to become a secondary science teacher in Boulder.
Amanda Hardy (MS), who studied the impacts of winter recreation on elk and bison (in YNP). Amanda went to a job as a wildlife ecologist for the Western Transportation Institute, and then a PhD at CSU.

Other Research:

From 1984-1986, I studied behavioral and physiological mechanisms that mediate the effects of early experience on milk production in Holstein dairy cattle. I am consequently one of a relatively small set of people who know how to make cows urinate on demand. Really.

From 1987-1990, my wife Nancy and I studied evolutionary, behavioral and physiological aspects of cooperative breeding in dwarf mongooses (Helogale parvula) in Serengeti National Park, working with Dr. Peter Waser and Dr Jon Rood, for my PhD at Purdue. This work involved using demographic and molecular genetic data to calculate inclusive fitness costs and benefits, and using behavioral and endocrine data to identify the mechanisms responsible for reproductive suppression in socially subordinate adults.

Some of the primary results of this research are found in:

Creel & Creel 1991. Energetics, reproductive suppression and obligate communal breeding in carnivores. Behavioral Ecology and Sociobiology 28:263-270.
Creel et al. 1991. Spontaneous lactation is an adaptive result of pseudopregnancy. Nature 351:660-662.
Creel & Waser 1991. Failures of reproductive suppression in dwarf mongooses (Helogale parvula): accident or adaptation? Behavioral Ecology 2:7-15.
Creel et al 1992. Behavioural and endocrine mechanisms of reproductive suppression in Serengeti darf mongooses. Animal Behaviour 43: 231-245.
Creel & Waser 1994 Inclusive fitness and reproductive strategies in dwarf mongooses. Behavioral Ecology 5:339-348.
Creel S 2001. Social dominance and stress hormones. Trends in Ecology and Evolution 16: 491-497

From 1991-1996, Nancy and I studied African wild dogs (Lycaon pictus) in the Selous Game Reserve. At roughly 80,000 square kilometers, the Selous is one of the largest protected areas in the world, but its ecology is still little-studied. This project focused initially on simply assessing the size of the wild dog population in Selous (a formidable task in itself), and progressed to identifying the ecological factors that cause wild dogs to be endangered, attaining uniformly low densities in comparison to other large carnivores that are well-protected by Tanzania's system of parks and reserves. In this regard, interspecific competition plays a major role in limiting wild dog numbers and distributions. We also used demographic data to make quantitative assessments of extinction risk, and collected a substantial data set on prey selection, predator-prey interactions and the costs/benefits of cooperative hunting. Finally we examined social evolution and behavioral and endocrine mechanisms of reproductive suppression in wild dogs, in a manner similar to our earlier work with dwarf mongooses.

Some of the major results of this work are found in:

Creel & Creel 1995. Communal hunting and pack size in African wild dogs, Lycaon pictus. Animal Behaviour 50:1325-1339.
Creel & Creel 1996. Limitation of African wild dogs by competition with larger carnivores. Conservation Biology 10:526-538.
Creel et al. 1996. Social stress and dominance. Nature 379: 212. (also see Morell, V 1996. Life at the top: animals pay the high price of dominance. Science 271: 292.)
Creel et al. 1997. Rank and reproduction in cooperatively breeding African wild dogs: behavioral and endocrine correlates. Behavioral Ecology 8:298-306.
Vucetich & Creel 1997. Ecological interactions, social organization and extinction risk in African wild dogs. Conservation Biology 13:1172-1182.
Creel & Creel 2002. The African wild dog: behavior, ecology and conservation. Princeton University Press.
Creel et al. 2004. African wild dogs: demography and population dynamics of wild dogs in three crticial populations. In: Biology and Conservation of Wild Canids, ed. by D.W. Macdonald & C Sillero-Zubiri, Oxford University Press.

Incidental to these studies, I've done some collaborative research on the behavioral ecology and evolution of lions, leopards, banded mongooses and slender mongooses, some of which is in the following:

Creel S & Creel NM 1997. Lion density and population structure in the Selous Game Reserve: evaluation of tourist hunting quotas and offtake. African Journal of Ecology 35:83-93
Spong G, Hellborg L & Creel S 2000. Sex ratio of leopards taken in trophy hunting: genetic data from Tanzania. Conservation Genetics 1: 169-171.
Spong G, Creel S, Stone J & Bjorklund M. 2002 Genetic structure of a population of lions (Panthera leo): implications for the evolution of sociality. Journal of Evolutionary Biology 15(6):945-953.
Spong G & Creel S 2004. Effects of kinship on territorial conflicts among groups of lions, Panthera leo. Behavioral Ecology and Sociobiology. 55: 325-331
Waser PM, Elliott L & Creel SR 1995. Habitat variation and mongoose demography. In: Serengeti II: Dynamics, conservation and management of an ecosystem. pp. 421-447, Sinclair ARE & Arcese P (eds). University of Chicago Press, Chicago.