Scott Creel

311-A Lewis Hall
Conservation Biology & Ecology Program
Department of Ecology

Montana State University
Bozeman MT 59717
Phone: 406-994-7033


Current Research

General Areas of Interest:  Behavioral ecology, population biology, conservation, behavioral endocrinology, evolutionary ecology.  Virtually all of my research is based on field studies, generally using observational methods, and often following known individuals.  Much of the work in my lab has involved the integration of behavioral and demographic data from the field with physiological and genetic data from the lab.  My lab is equipped for extraction and enzyme immunoassays of steroid and peptide hormones, microhistology and other assays

My current research mainly examines risk effects, or the costs of antipredator responses by prey species.  Recent research with many species has shown that direct killing constitutes only a fraction of the total limiting effect of predators on their prey.  We have recently examined the responses of elk to variation in the risk of predation by wolves, including changes in behavior, group size, habitat selection, feeding ecology, and spatial distributionsWe related these responses to changes in nutrition, physiology, demography and population dynamics. This work won the 2010 Carl Gustav Bernhard Medal from the Royal Swedish Academy of Sciences.

Our primary field research is now with the Zambian Carnivore Programme, headed by Dr. Matt Becker, to examine risk effects and carnivore conservation in three ecosystems (Liuwa, South Luangwa and Kafue National Parks), with several species of predators and prey.  This work aims to measure variation in direct predation rates and risk effects, to identify the ecological factors that drive this variation, and provide data for the conservation and management of predators and prey.  In this work, we are studying African wild dogs, lions, spotted hyenas, cheetahs and leopards, together with their primary prey, which varies among species and ecosystems, but notably includes African buffalo, giraffe, wildebeest, zebra, impala, puku, lechwe and oribi.

My students and I address these questions with a variety of methods, including behavioral observations, demographic & ecological monitoring, ground and aerial censuses, camera trapping, GPS and VHF radiotelemetry, enzyme immunoassay of fecal steroid metabolites to measure pregnancy rates and glucocorticoid stress responses, measurements of abiotic factors such as weather and soil nutrients, and assessments of diet quality by chemical, radioisotope and microhistological methods.

roan and

awd kafue

busanga brothers


  Recent Publications: 

will burrard lucas AWD photoCreel S, M’soka J, Dröge E, Rosenblatt E, Becker M, Matandiko W, Simpamba T. 2016. Assessing the sustainability of African lion trophy hunting, with recommendations for policy. Ecological Applications, in press.

Schuette P, Creel S & Christianson D. 2016. Ungulate distributions in a rangeland with competitors, predators, and pastoralists. Journal of Applied Ecology 
DOI: 10.1111/1365-2664.12610

Rosenblatt, E., Creel, S., Becker, M. S., Merkle, J., Mwape, H., Schuette, P., & Simpamba, T. 2016. Effects of a protection gradient on carnivore density and survival: an example with leopards in the Luangwa valley, Zambia. Ecology and Evolution. DOI: 10.1002/ece3.2155

Scott Creel, Matthew Becker, David Christianson, Egil Dröge, Neil Hammerschlag, Matt W. Hayward, Ullas Karanth, Andrew Loveridge, David W. Macdonald, Wigganson Matandiko, Jassiel M’soka, Dennis Murray, Elias Rosenblatt, Paul Schuette. 2015. Questionable policy for large carnivore hunting.  Science 350:  1473-1475 (PDF)

Creel S. & Creel N.M. 2015.  Opposing effects of group size on reproduction and survival in African wild dogs.  Behavioral Ecology 26: 1414-1422.
Brennan A, Cross P & Creel S.  2015 Managing more than the mean: using quantile regression to uncover relationships with large elk groups and inform management.  Journal of Applied Ecology, 52(6), 1656-1664.

Foley, A.M., Cross, P., Christianson, D, Scurlock, B & Creel S. 2015.  Influences of supplemental feeding on winter elk calf:cow ratios in the Southern Greater Yellowstone Ecosystem.  Journal of Wildlife Management,
79(6), 887-897.

Nelson  J.L,  Creel S & Cypher B.L. 2015. Fecal glucocorticoid levels of endangered San Joaquin kit foxes (Vulpes macrotis mutica) in natural and urban habitats.Western North American Naturalist 75(1): 52–57.
Durant, S.M., Becker, M.S., Creel, S., Bashir, S., Dickman, A.J., Beudels-Jamar, R.C., Lichtenfeld, L., Hilborn, R., Wall, J., Wittemyer, G., Badamjav L., Blake, S., Boitani, L., Breitenmoser, C., Broekhuis, F., Christianson, D., Cozzi, G., Davenport, T.R.B., Deutsch, J., Devillers, P., Dollar, L., Dolrenry, S., Douglas-Hamilton, I., Dröge, E., FitzHerbert, E., Foley, C., Hazzah, L., Hopcraft, J.G.C., Ikanda, D., Jacobson, A., Joubert, D., Kelly, M.J., Milanzi, J., Mitchell, N., M’Soka, J., Msuha, M., Mweetwa, T., Nyahongo, J., Rosenblatt, E., Schuette, P., Sillero-Zubiri, C., Sinclair, A.R.E., Stanley-Price, M. R., Zimmermann, A., Pettorelli, N. 2015.  Developing fencing policies for dryland ecosystems.  Journal of Applied Ecology

(PDF) Christianson D & Creel S 2015.  Photosynthetic pigments estimate diet quality in elk (Cervus elaphus) feces and forage.  Canadian Journal of Zoology  93: 51–59

(PDF) Rosenblatt E., Becker M, Creel S, Droge E, Mweetwa T, Schuette P, Watson F, Merkle, J, Mwape H 2014. Detecting declines of apex carnivores and evaluating their causes: an example with Zambian lions.  Biological Conservation 180: 176-186.

(PDFChristianson D & Creel S  2014. Ecosystem scale declines in elk recruitment and population growth with wolf colonization: a before-after-control-impact approach. PLoS ONE 9(7): e102330. doi:10.1371/journal.pone.010233

(PDF) Creel S, Schuette P & Christianson D 2014.  Effects of predation risk on group size, vigilance and foraging behavior in an African ungulate community.  Behavioral Ecology 25 (4): 773-784

Brennan, A., P. C. Cross, M. D. Higgs, W. H. Edwards, B. M. Scurlock, and S. Creel. 2014. A multi-scale assessment of animal aggregation patterns to understand increasing pathogen seroprevalence. Ecosphere 5(10):138.

Benavides JA Cross PC, Luikart G & Creel S 2014  Limitations to estimating bacterial cross-species transmission using genetic and genomic markers: inferences from simulation modelling.  Evolutionary Ecology, advanced online doi: 10.1111/eva.12173.

(PDF) Creel S, Winnie JA, Christianson D. 2013 Underestimating the frequency, strength and cost of anti-predator responses with data from GPS collars: an example with wolves and elk. Ecology and Evolution 3: 5189-5200.

Coleman TA, Schwartz CC, Gunther, KA & Creel S. 2013 Influence of overnight recreation on grizzly bear movement and behavior in Yellowstone National Park. Ursus 24: 101-110.

(PDF) Creel, S, Christianson D & Schuette P 2013. Glucocorticoid stress responses of lions in relation to group composition, human land use and proximity to people. Conservation Physiology 1: 1-11   doi: 10.1093/conphys/cot021

(PDF) Creel, S., Becker, M.S., Durant, S.M., M’Soka, J., Matandiko, W., Dickman, A.J., Christianson, D.,  Dröge, E., Mweetwa, T., Pettorelli, N., Rosenblatt, E., Schuette, P., Woodroffe, R., Bashir, S., Beudels-Jamar, R.C., Blake, S., Borner, M., Breitenmoser, C., Cozzi, G., Davenport, T.R.B., Deutsch, J., Dollar, L., Dolrenry, S., Douglas-Hamilton, I., Foley, C., Hazzah, L., Henschel, P., Hilborn, R., Hopcraft, G., Ikanda, D., Jacobson, A., Joubert, B., Joubert, D., Kelly, M.S., Lichtenfeld, L., Mace, G.M., Milanzi, J., Mitchell, N., Msuha, M., Nyahongo, J., Pimm, S., Purchase, G., Schenck, C., Sillero-Zubiri, C., Sinclair, Songorwa, A., A.R.E., Stanley-Price, M., Tehou, J., A., Trout, C., Wall, J., Wittemeyer, G., Zimmermann, A., 2013. Conserving large populations of lions - the argument for fences has holes. Ecology Letters 16: 1413 - e3,  doi: 10.1111/ele.12145

(PDF) Schuette P,  Creel S & Christianson D  2013. Coexistence of African lions, livestock, and people in a landscape with variable human land use and seasonal movements. Biological Conservation 157: 148-154.

(PDF) Cross, PC, Creech, TG, Ebinger MR, Manlove K, Irvine K, Henningsen J, Rogerson J, Scurlock BM, Creel S. 2013.  Female elk contacts are neither frequency nor density dependent. Ecology 94:2076–2086 

(PDF) Coleman, TH, Schwartz, CC, Gunther, KA, Creel, S. 2013.  Grizzly bear and human interaction in Yellowstone National Park: An evaluation of bear management areas. Journal of Wildlife Management 77: 1311-1320.   

Brennan A, Cross P, Ausband D, Barbknecht A, Creel S. 2013.  Testing automated howling devices in a wintertime wolf survey.  Wildlife Society Bulletin 37: 389 - 393.

AWD book cover

Creel S & Rosenblatt E 2013 Using pedigree reconstruction to estimate population size: genotypes are more than individually unique marks. Ecology and Evolution 3: 1294-1304 (open access at

(PDF) Creel S, Dantzer B, Goymann W & Rubsentein D 2013.  The ecology of stress: effects of the social environment.  Functional Ecology 27:66-80. (invited review).
Brennan A, Cross PC, Higgs M, Beckmann JP, Klaver RW, Scurlock BM & Creel S 2013 Inferential consequences of modeling rather than measuring snow accumulation in studies of animal ecology. Ecological Applications, 23:643-653.

(PDF) Creel, S 2013.  Helogale parvula.  In: Mammals of Africa, Volume V: Carnivores, Pangolins, Equids and Rhinoceroses. Kingdon J, Happold D, Butynski T, Hoffmann M, Happold M, Kalina J (eds). pp 368-373,  Bloomsbury: London.

(PDF) Schuette P, Wagner A, Wagner M & Creel S 2013. Occupancy patterns and niche partitioning within a diverse carnivore community exposed to anthropogenic pressures.  Biological Conservation 158:301-312.

Marsden, C.... Creel, S et al. 2012. Spatial and temporal patterns of neutral and adaptive genetic variation in the endangered African wild dog (Lycaon pictus) Molecular Ecology 21:1379-1393.

Cross PC, Creech TG, Ebinger MR, Heisey D,  Irvine K & Creel S. 2012.  Wildlife contact analysis: emerging methods, questions and challenges. Behavioral Ecology and Sociobiology

Forristal  VE, Creel S, Taper ML, Scurlock BM, Cross PC  2012.  Effects of supplemental feeding and aggregation on fecal glucocorticoid metabolite concentrations in elk. Journal of Wildlife Management 76:694-702.

Ezenwa V, Ekernas LS, Creel S 2012. Unravelling complex associations between testosterone and parasite infection in the wild. Functional Ecology 26: 123-133.

Creech, T., P.C. Cross, S. Creel, E. Maichak, & B. Scurlock 2012. Effects of low-density feeding on elk-fetus contact rates on Wyoming feedgrounds. Journal of Wildlife Management 76:877-886.

(PDF) Creel S 2011. Toward a predictive theory of risk effects: hypotheses for prey attributes and compensatory mortality.  Ecology 92: 2190–2195

(PDF) Creel S, Christianson D and Winnie JA  2011.  A survey of the effects of wolf predation risk on pregnancy rates and calf recruitment in elk.  Ecological Applications 21: 2847–2853

Abbott, P...Creel S et al 2011  Inclusive fitness theory and eusociality.  Nature 471: E1-E4.

awd cover Sawaya MA, Ruth TK, Creel S, Rotella JJ, Quigley HB, Stetz JB, Kalinowski ST (2011)  Evaluation of noninvasive genetic sampling methods for cougars in Yellowstone National Park  Journal of Wildlife Management 75: 612–622.

Griffin K, Hebblewhite M, Robinson H, Zager P, Barber-Meyer S, Christianson D, Creel S, Harris N, Hurley M, Jackson D ( 2011) Neonatal mortality of elk driven by climate, predator phenology and predator diversity. Journal of Animal Ecology  80: 1246-1257

 (PDF) Creel S & Rotella J 2010 Meta-analysis of relationships between human offtake, total mortality and population dynamics of gray wolves (Canis lupus). PLoS One 5(9): e12918.  (available open-access at doi:10.1371/journal.pone.0012918)

(PDF) Christianson D & Creel S 2010  A nutritionally mediated risk effect of wolves on elk.  Ecology 91:1184-1191

Creel S. 2010.  Interactions between wolves and elk in the Yellowstone Ecosystem, pp. 65 - 80 In Knowing Yellowstone: Science in America's First National Park, J. Johnson (ed) Taylor Trade Publishing, Boulder CO.

Creel S 2010. Studying predator-prey interactions from the perspective of the predator.  Ch 328 in The Encyclopedia of Animal Behavior.  Eds M Breed & J moore. Elsevier.

PDF files for papers from 2001-2009:

2007     2006 
2005     2004 
2003     2002 

Graduate Students:
Current graduate students    Former graduate students    I encourage my graduate students to participate fully in developing their own research questions and to pursue independent funding.  Here are some brief notes on the process of writing a paper, and here are some ideas about preparing for the qualifying exam.

Older Research:

From 1984-1986, I studied beh
avioral 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 Na
ncy 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 these papers.

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 these papers, and in Creel S & Creel N 2002. The African wild dog: behavior, ecology and conservation.  Princeton 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 t
hese papers.