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Scott
Creel
Professor of Ecology
302 Lewis Hall
Department of Ecology
Montana State University
Bozeman MT 59717
Email: screel@montana.edu
Phone: 406-994-7033
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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 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 physiological and genetic data from the lab. My
lab is equipped for extraction and radioimmunoassay or enzyme
immunoassays of steroid hormones, DNA extraction, and
PCR.
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My current research mainly 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 grouping.
(2) To determine the costs 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 South Rift Valley of
Kenya. With Dr. Jonah Western of the African Conservation
Centre and other collaborators, we are examining interactions between
lions, spotted hyenas and their prey, effects on small and medium-sized
carnivores, and interactions
with people and cattle inside and outside of a Community Conservation
Area. This research is testing the
generality of conclusions from the wolf-elk project, and identify
ramifications for ecosystem function, management and conservation in
East Africa.
My students and I
address these goals with a wide variety of methods,
including behavioral observations, demographic monitoring, ground and
aerial
censuses, camera trapping, GPS and VHF
radiotelemetry, RIA/EIA of fecal steroids to measure
pregnancy rates and stress responses, measurements of
abiotic factors such as weather and snow conditions, and assessments of
diet quality by chemical and microhistological methods.
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Teaching:
- Conservation
Biology, BIOL 447 & BIOL 521, Fall
semester
- Principles of Ecology, BIOL
303, Spring semester. I am currently
revising this course, but it will cover aspects of demography/life
history, single-species population dynamics, competition, predation,
disease, community ecology, behavioral ecology and evolutionary ecology.
- Animal
Physiology, Biol 411, Fall semester (not currently involved,
though I taught part of this course for 10 years)
- I also teach a one credit graduate seminar (BIOL 500) in
Fall of most years. The emphasis is on discussion of
papers, grant proposals and presentations of our own work.
Recent Publications:
Creel
S, Winnie JA & Christianson D (in press) Glucocorticoid stress
hormones and the effect of predation risk on elk reproduction. Proceedings of the
National Academy of Sciences.
Christianson D & Creel S (in press) A nutritionally mediated
risk effect of wolves on elk. Ecology.
Creel S (in press) Studying predator-prey interactions from the
perspective of the predator. Ch 328 in The Encycolpedia of Animal
Behavior. Eds M Breed & J moore. Elsevier.
Creel S & Creel M (in press) Density-dependence and climate
effects in Rocky Mountain elk: an application of regression with
instrumental variables for population time series with sampling
error. Journal
of Animal Ecology
Creel
S & Christianson D (in press) Wolf presence and increased
willow consumption by Yellowstone elk: implications for trophic
cascades. Ecology.
Christianson D & Creel S. (in press)
Using chlorophyll in feces to
describe the relationship between primary production and
consumption. Ecological
Applications.
(PDF) Christianson
D
& Creel S. 2009 Effects
of grass and browse consumption on winter mass dynamics of elk Oecologia 158: 603 - 613.
(PDF)
Creel S &
Christianson D 2008. Relationships between direct
predation and risk effects. Trends in Ecology
& Evolution 23:
194-201.
(PDF)
Creel
S, Winnie JA,
Christianson D & Liley S 2008.
Time and space in general models of antipredator response: tests with
wolves and elk. Animal Behavior
76: 1139-1146
(PDF)
Christianson D & Creel S. 2008. Risk effects in
elk: sex-specific response in grazing and browsing due to predation
risk from wolves. Behavioral Ecology 19: 1258 - 1266.
Costello C, Creel S, Kalinowski ST, Vu, NV & Quigley HB 2008.
Sex-biased natal dispersal and inbreeding avoidance in American black
bears as revealed by spatial genetic analyses. Molecular Ecology 17: 4713-4723
Wagner AP,
Frank
LG, Creel S 2008. Spatial grouping in behaviourally solitary
striped hyenas (Hyaena
hyaena) Animal Behaviour 75: 1131-1142. PDF
(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, Christianson D, Liley S & Winnie J 2007.
Effects of predation risk on reproductive physiology and
demography in elk. Science 315: 960.
Also see Supplemental
Online Material for this paper with methods, detailed assay
validation data and additional results.
(PDF)
Wagner AP Frank LG, Creel S, Coscia EM 2007.
Transient genital
abnormalities in striped hyenas (Hyaena hyaena) Hormones and Behavior 51: 626-632,
(PDF)
Wagner AP, Creel S, Frank LG & Kalinowski S.
2007. Patterns of relatedness and parentage in an asocial,
polyandrous striped hyena population. Molecular Ecology
16: 4356 - 4369.
Creel S 2007. Helogale parvula. In: Kingdon, J.S. &
Hoffmann, M. (Eds). The Mammals of Africa. Vol 5. Academic Press,
Amsterdam.
Nelson, JL, Cypher BL, Creel S & Bjurlin C 2007.
Effects of landscape modification on competition between endangered kit
foxes and coyotes. Journal of Wildlife
Management 71: 1467-1475.
(PDF)
Christianson D & Creel S 2007. A review of environmental factors
affecting winter elk diets. Journal of
Wildlife Management 71: 164-176.
(PDF)
Winnie, J & Creel, S 2007. Sex-specific behavioral responses
of elk to spatial and temporal variation in the threat of wolf
predation. Animal Behaviour.
71: 215
- 225.
(PDF)
Wagner AP, Creel S & Kalinowski ST 2006. Maximum likelihood
estimation of relatedness and relationship using microsatellite loci
with null alleles. Heredity
97: 336-345
(PDF)
Winnie, J, Christianson D, Maxwell B & Creel, S 2006.
Elk decision-making rules are simplified in the presence of
wolves. Behavioral Ecology and
Sociobiology 61: 277 - 289.
(PDF)
Kalinowski ST, ML Taper, S Creel 2006. Using DNA from
non-invasive samples to census populations: an evidential approach
tolerant of genotyping errors. Conservation
Genetics 7: 319-329.
(PDF)
Creel S 2006. Recovery of the Florida panther - genetic rescue,
demographic rescue, or both? Animal
Conservation. 9: 125-126.
Bergman E, Garrott R, Creel S, Borkowski J, Jaffe R, Watson F
2006. Assessment of prey vulnerability through analysis of wolf
movements and kill sites. Ecological
Applications 16: 273-284.
(PDF)
Creel S 2006. In the minds of animals. Nature
439: 662-663.
(PDF)
Dalerum F, Creel S & Hall S 2006. Behavioural and endocrine
correlates of reproductive failure in social aggregations of captive
wolverines. Journal of Zoology 269: 527 - 536.
(PDF)
Creel S, Winnie JA, Maxwell B, Hamlin K & Creel M 2005. Elk
alter habitat selection as an antipredator response to wolves. Ecology 86:3387-3397.
(PDF)
Creel S & Winnie J 2005. Responses of elk
herd size to fine-scale spatial and temporal variation
in the risk of predation by wolves. Animal
Behaviour 69:1181-1189.
(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.
Students:
Here are some excellent general
notes about preparing for the graduate qualifying exam, from
Colleen Cassady
St Clair at the Univesity of Alberta. Though they pertain to the
exam
at a different department & school, they apply well here.
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.
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 (PhD): 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.
Angela Brennan (PhD): Angela, co-advised by Dr. Paul Cross of
USGS, is developing a project to examine the ecology and epidemiology
of brucellosis in elk on Wyoming
feedgrounds and the Yellowstone Ecosystem.
Tyler Creech (MS): Tyler co-advised by Dr. Paul Cross of
USGS, is also developing a project to examine the
ecology and epidemiology of brucellosis in elk on
Wyoming feedgrounds and the Yellowstone Ecosystem.
Tiffany Holland (MS): Tiffany, co-advised by Dr. Marcel
Huisjer of WTI, is examining factors that contribute to vehicle
collisions with deer on US 93, specifically testing for reductions
after mitigation measures.
Recent graduate students are:
Dave Christianson (PhD):
Dave is
examined changes in elk foraging
behavior in response to the presence of wolves, and the impacts of
these changes on elk diets, nutrition, and demography. He went on to an
EU Fellowship with Anne Loison's research group at the
Université de Savoie, to
examine interactions between predation and resource limitation on
chamois nutrition and population dynamics.
Cecily Costello (PhD): Cecily
studied black bear social organization and
space use as they relate to population genetics and patterns
of relatedness, using data from two populations in NM. Wildlife
Conservation Society/Hornocker Wildlife Institute grants. Cecily
went on to research with the Interagency Grizzly Bear Study
Team on the genetic structure of N. Rockies populations of grizzly
bears and
issues to do with ESA protections.
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.
For questions or
comments: screel@gemini.oscs.montana.edu
Back to the MSU Department
of Ecology.
Perception
test
Ecology
and cell phone coverage - who says ecology doesn't solve important
societal problems?
The
latest in research on squirrel fishing
In case of
terrorist attack...
Oooh...
scaary..
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