Wildlife Society Bulletin, attraction, habituation, Richard L. Knight, wildlife professionals, Island Press, stimuli, avoidance, Colorado State University, wildlife management, Doug Whittaker, Journal of Wildlife Management, New York, New York, consequences, Stable URL, pp, J. Whittaker, extraneous variables, Wintering Bald Eagles, Daniel J. Grout, basic concepts, response, Stanley A. Temple, Klopfer, experimental settings, culturally transmitted, Anchorage, Alaska, Knight, human persecution, research tradition, wildlife researchers, Pennsylvania State University, European settlement, response to stimuli, Understanding wildlife, North American Wildlife, Bear Research and Management, Oregon State University Press, Piscataway, New Jersey, D S. A. TEMPLE19, Saskatchewan, Canada, Northwest Territories Department of Renewable Resources, Plenum Press, Wildlife Society, Washington D.C., Washington, D.C., University of Chicago Press, Cambridge University Press, professionals, Whittaker, responses, Winchester Press, J. M. Gerard, Belknap Press, Stackpole Books
IN MY OPINION
Understanding wildlife responses
Doug Whittaker and Richard L. Knight
A herd of 40 elk (Cervus canadensis) barely look begin with attention to definitions of concepts and
up from grazing in a Yellowstone meadow as a dozen extend to the development of new models for de-
motorhomes crowd a nearby highway. A 450-kg scribing, predicting, and evaluating responses. Hop-Brown Bear
(Ursus arctos) flees his fishing spot on an ing to stimulate discussion and the development of
Alaskan stream at the sight of an 80-kg man. Canada improved models, we review the basic concepts and
geese (Branta canadensis) in a city park walk di- identify 3 issues that wildlife professionals should
rectly toward people in search of hand-outs. Differ- consider when working in the area of human-wild-
ent settings, different species, and different re- life interactions.
sponses to humans or their environments, but each
suggests the same underlying question: Why do wildlife exhibit such diverse behavior around people?
The general answer is that wildlife have developed Although wildlife responses to humans vary, it is
situation-specific responses because some combina- possible to broadly describe 3 classes of wildlife re-
tion of learning and genetics have made them suc- sponses as attraction, habituation, and avoidance
cessful. As the human-wildlife interface changes, (Knight and Cole 1991). In general, an animal can
however, some of these responses may become detri- find human-provided stimuli reinforcing (leading to
mental for wildlife, or lead to conflicts
with people. In order to manage these
situations, wildlife professionals should
differentiate between responses, iden-
tify their causes and consequences, and
evaluate those consequences. While
several research traditions provide con-
cepts useful for this effort, variation ex-
ists in the way these concepts are un-
derstood and applied by researchers,
managers, and the public. Some of
these concepts have also become value-
laden (always an issue when humans
study phenomena connected to them-
selves), and this can become an obstacle
We think there is a need to recon-
sider how wildlife management describes and makes life responses to humans. This should
Habituated brown bears at Katmai National Park, Alaska provide high-quality viewing opportunities, but habituation may make bears easier targets for poachers or hunters if they leave protected areas
. Photo from Brooks River. Photo by J.Whittaker.
Address for Doug Whittaker: Human Dimensions of natural resources
, Colorado State University
, Fort Collins
, CO 80523, USA. Present address for Doug Whittaker: P.O. Box 2036 Nome, Alaska 99762, USA. Address for Richard L. Knight: Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins, CO 80523, USA. Key words: attraction, avoidance, habituation, human-wildlife interface, wildlife responses
Wildlife Society Bulletin 1998,26(2):312-317
In my opinion Whittaker and Knight 313
attraction), aversive (leading to avoidance), or neutral Sensitization is an allied concept that is also com-
(leading to habituation; Gilbert 1989). Most wildlife monly misused. "Sensitization" describes increased
professionals have a general understanding of these response to stimuli, and is the opposite of habitua-
terms, but the specific meanings they attach to these tion (Domjam and Burkhard 1986, Immelmann and
terms are influenced by their training. Some of the Beer 1989). Researchers have suggested that sensiti-
terms (particularly habituation) were developed from zation and habituation may occur simultaneously;
precise concepts in behavioral psychology
and ethol- only the net effect is observed (Groves and Thomp-
ogy, but they are also used by wildlife professionals in son 1970). This dual-process theory explains phe-
broader contexts. While recognizing the value of pre- nomena such as recovery, in which habituation is re-
cise terminology, we believe these concepts should placed by a highly sensitized state with a slight
be widely understood across backgrounds. In this change in the stimulus.
spirit, a brief review of terms follows.
"Avoidance" is the opposite of attraction, an aver-
"Attraction" in wildlife management is defined as sion to negative consequencesassociated with a stim-
the strengthening of an animal's behavior because of ulus (Knight and Cole 1991). Some behaviorists dis-positive reinforcement
, and implies movement to- tinguish avoidance from escape; the former refers to
ward the stimuli (Knight and Cole 1991). Developed actions that prevent aversive consequences, whereas
within the associative-learning tradition of behavioral the latter is a reaction to aversive consequences
psychology (Klopfer 1974, Domjan and Burkhard (Domjam and Burkhard 1986).
1986,Manning and Dawkins 1992),wildlife attraction Avoidance can apply to a range of stimuli, includ-
is often presumed to be about food-conditioning but ing those in which the stimulus is aversive (uncondi-
is equally applicable to behaviors that attract wildlife tioned response) and those in which the stimulus is
to shelter or security. Examples of attraction include associated with aversive consequences (conditioned
Clark's nutcrackers (Nuayraga columbiana) that fly response). Deer (Odocoileus spp.) can learn to
to picnickers for handouts, cardinals (Cardinalis car- avoid touching an electrified fence, and wolves (Ca-
dinalis) at bird baths or feeders, or rainbow trout nis lupus) can learn to avoid towns or road systems
(Salmo gairdneri) lured to the cold water of a dam because they associate them with human persecu-
tailwater (or the fly on the end of an angler's line). tion (Thurber et al. 1994).
Habituation Of the 3 terms, "habituation"is the most often misapplied and is commonly confused with attraction. Unlike attraction, which involves a reinforcing stimuli, habituation is a waning of response to a repeated, neutral stimuli (Humphrey 1930, Thorpe 1956, EiblEibelsfeldt 1970). This formal definition, used with precision in behavioral psychology and ethology, is less carefully used by some wildlife professionals. Habituation, for example, has little to do with the words "habitual"or "habit,"even though they share a common root. A bear may be in the habit of going to a garbage dump to look for food, but this is attraction, not habituation (Gilbert 1989). Being habituated to human food would imply that the bear ignores it. Wildlife are capable of becoming habituated to people, human-made environments, and most any human stimuli. The wildlife in the Galapagos Islands epitomize animals habituated to the presence of people, but habituation is also an apt description for crows (Corvus spp.) ignoring a scarecrow, or a red fox (Vulpes vulpes) ignoring the human activity of a suburban area.
Issue 1: Wildlife responses and causality Attraction, habituation, and avoidance are used to describe behavior in response to a known stimuli. In Canada geese and other waterfowl may become attractedto urban parks and handouts, which may create unhealthy dependencies on artificial food sources. Photo from Anchorage, Alaska. Photo by J. Whittaker.
314 Wildlife Society Bulletin 1998, 26(2):312-317
behavioral psychology, where these terms were first developed, this is often the case: research explores how animals (usually rock doves [Colunzba livia] and rats [Ratus spp.]) respond to stimuli (usually shocks or food reinforcement) in experimental settings where extraneous variables are tightly controlled (Domjan and Burkhard 1986). In this research tradition, the exploration of causality is central. In the European ethology and wildlife management traditions, however, emphasis on observations in natural settings
make causality far less certain (Klopfer 1974). In these settings, it is difficult to divorce stimuli from their context; this suggests a methodological shift from an experimental to a correlational perspective that explores more variables. This is particularly relevant as we try to understand the underlying processes that cause wildlife responses. Although many wildlife researchers consider attraction, habituation, and avoidance to be functions of learning (Klopfer 1974, Newton 1979, Poole 1981, Buitron 1983, Fraser 1984, Knight and Temple 1986, Knight and Cole 1991, Manning and Dawkins 1992), others suggest genetic roots (Klopfer 1974, Knight and Temple 1995). Hailman's (1967) concept of "learning an instinct" suggests how genetic and learned components may be intertwined and could have particular relevance for understanding avoidance responses. For example, bighorn sheep (Ovis canadensis) and mountain goats (Orenmnos americanus), withdraw to cliffs in response to sudden, loud noises such as rockfalls (Geist 1971, 1978). When gunshots invoke a similar response, it suggests a genetic component being reinforced through learning. In another example, studies of red-tailed hawks (Buteo jamaicensis) suggest that associations with human persecution may have become genetically coded in hawks because hawk populations in areas with longer histories of European settlement
(which had firearms) were less aggressive and more oriented toward avoidance than those in areas with more recent settlement by Europeans (Knight et al. 1989). It is also possible that wildlife responses to humans may be culturally transmitted across generations. Thurber et al. (1994) suggested that wolves teach their young to avoid certain human environments because of associations with persecution, and DouglasHamilton and Douglas-Hamilton (1975) suggested that the behavior of a band of unusually nocturnal and aggressive elephants (Loxodonta africana) could be traced to a persecution event in 1919-too long ago to be part of the living elephants' experience, but too recent to have become genetically encoded in their behavior. Considerable research has
explored the cultural transmission of behavior in higher animals, particularly primates (McGrew et al. 1979, Bonner 1980, Goodall 1986), and most of this research suggests more extensive cultural transmission than previously believed. The likelihood of genetic and culturally transmitted influences on wildlife responses suggests the need for richer models than those developed in the learning and behaviorist traditions. While differentiating these components will be challenging, the information could be useful. Management actions that take advantage of genetic predispositions or culturally transmitted responses are likely to be more successful than those that depend on learned behavior during the lifetimes of individual animals (e.g., aversive conditioning techniques that mimic stimuli that a species finds naturally aversive). More research is needed to explore the links between attraction, habituation, and avoidance. Evidence exists, for example, that attraction can lead to habituation. Primate researchers often gain access to their subjects by providing food attractants during the initial stages of field observation; over time the animals learn to ignore human presence and the attraction can be removed (Goodall 1986, Lofstrom and Stallings 1973). Wildlife professionals also have suggested that habituation may lead to attraction, particularly among bears (McCullough 1982, Herrero 1985,Jope 1985, Olson et al. 1997). In learning to ignore people, habituated wildlife have greater opportunities to find attraction stimuli in human environments. Because humans generally attempt to avoid attracting potentially dangerous animals, understanding these types of links may be critical. Issue 2: Response events or response tendencies? In the behaviorist and ethology traditions, attraction, habituation, and avoidance are descriptions of behavioral events or processes-specific stimuli and specific responses. These events are also carefully placed in context with descriptions of the conditions under which the animal was attracted to the stimulus. Wildlife professionals, however, sometimes label an animal, a sub-group, or an entire population based on limited responses from a few animals. Observing an animal that does not withdraw from stimuli, ethologists might describe the behavior as habituation; in contrast, some wildlife professionals might extend this conclusion and describe the animal as habituated. This kind of conceptual extension has undeniable utility for comparing the response tendencies of indi-
In my opinion Whittaker and Knight 315
vidual animals or groups of animals. But it also invites confusion because the labels go beyond observed events. The event is concrete; the tendency to behave is more abstract and uncertain. Our purpose is not to discourage this kind of labeling, but to encourage clarity about whether the label refers to past behavior or predicted future tendencies. Labeling animals or populations may also direct attention away from context variables. The concepts of attraction, habituation, and avoidance do not suggest that wildlife learn the same responses for all situations. For animals with greater cognitive abilities and longer rearing periods, new settings or additional stimuli may invoke considerably different responses (Orians 1981, Gilbert 1989, Manning and Dawkins 1992). Models that recognize greater complexity and subtlety in wildlife responses are needed. Nest-defense studies, for example, have suggested that birds distinguish different types of predators, different individuals, and different postures of predators when choosing their defense response (Knight and Temple 1986,Knight et al. 1987). Wildlife also behave differently in different locations and during different activities, and the learned outcomes of all these interactions affect subsequent interactions (Gilbert 1989). Success in understanding wildlife responses requires careful measurement of these contextual variables. The focus should not be simply, "Are these animals habituated?" More interesting questions include, "Underwhat conditionsdoes habituated behavior exist?"and "Howlong does habituation take to appear?" and "How will additional stimuli affect responses?" Researchers might also consider treating wildlife responses as continuous rather than dichotomous variables. For example, an animal's behavior is not simply habituated or nonhabituated, but a matter of degree. Wildlife responses occur in context and in 7, ,! Moose may become attracted to the landscaping in suburban environment
s, which can create conflicts with humans. Subdivision in Wasilla, Alaska, near Anchorage. Photo by J.Whittaker.
differing magnitudes in different contexts. This is commonly understood by field biologists who develop decision-rules to identify whether an animal is exhibiting habituation, attraction, or avoidance. But these decision rules may fail to capture important levels of these responses. Attraction, habituation, and avoidance are broad, latent, or intangible dispositions that we measure through outward responses. While 1 yes-or-no variable may offer initial understanding, finer-grained analyses that feature scaled variables are also possible and should be developed.
Issue 3: Evaluating wildlife responses
Attraction, habituation, or avoidance responses are
not intrinsically good or bad. Value judgments, how-
ever, are commonly attached to these terms, and can
be an obstacle to effective management. The conse-
quences of wildlife responses are not always immedi-
ate, direct, or obvious; history is replete with exam-
ples of misguided judgments about the consequences
of an animal's behavior (Steinbeck and Ricketts 1941).
In addition, even when we understand these conse-
quences, disagreement may arise about the accept-
ability of them. Responses harmful to 1 species may
be beneficial to another; it is a philosophical, rather
than a scientific position to suggest which should pre-
vail (Fleischman 1969). Value judgments about "what
should be" are an essential component of resource de-
cisions, but premature integration with information
about "what is" can lead to misapplications (Shelby
and Heberlein 1986, Bekoff and Jamieson 1991).
Cataloging the consequences of wildlife responses
is beyond the scope of this paper. Some examples,
however, may suggest why value judgments should
be cautiously applied:
- Bird feeding mav attract and boost ~ o ~ u l a -
tions such as chickadees (Parus atricapillus)
or bald eagles (Haliaeetus leucocephalus;
Brittingham and Temple 1988, Knight and
Anderson 1990) and is widely accepted. But
bird feeding may also have subtle long-term
consequences by favoring certain species and
contributingto an overall decline in diversity
(George 1982). Feeders may also make birds
dependent on artificial food sources, alter the
quality of their diet, and change intraspecific
interactions in an area.
Attraction of brown bears to garbage dumps
is typically discouraged because it increases
risks of injury to people or bears (Herrero
1985,Gilbert 1989,Albert and Bowyer 1991).
316 Wildlife Society Bulletin 1998, 26(2):312-31 7
Some researchers, however, have suggested that human-provided food centers (e.g., elk carcasses) may help support a threatened population (Craighead et al. 1995). The ability to habituate to humans is considered an essential survival skill for many urban wildlife species, particularly song birds (Burger and Gochfeld 1991). Disturbances from approaching humans would affect many birds' ability to feed, breed, and rest if they were unable to habituate. Bald eagles on salmon rivers with heavy recreational use face the same problem (Knight and Knight 1984). This same habituation response, however, may make these birds susceptibleto human persecution. Brown bears that become habituated to humans at salmon streams provide people with the opportunity to view bears and allow bears to fish without disruption. Habituation responses may also create "sanctuaries" for subadults and family groups, because large boars are more likely to avoid some of these viewing areas (Olson et al. 1997). Other consequences of habituated behavior in bears, however, may be less positive. Bears that ignore people at bear-viewing areas are easier targets for hunters or poachers outside protected areas. Habituated bears are also more likely to encounter attraction stimuli in the human environment (Hererro 1985; R. Squibb, Summary of bear management issues, unpubl. rep., Katmai Natl. Park and Preserve Headquarters, King Salmon,Alas., 1991). Avoidance responses are beneficial to persecuted species for which a human encounter can mean death, yet these same responses may prevent the animals from using otherwise suitable environments, or cause elevated and damaging levels of stress. Some animals, such as black bears (Ursus arnericanus), become adept at using areas of intense human use while still avoiding people. From a park manager's point of view this may be a "model" response, but it still prevents the bears from using their entire home range (McCutchen 1989). Extreme avoidance may be an effective survival strategy for an isolated subpopulation (e.g., Grumbine's 119921 "ghost bears" of the North Cascades), but it may also have detrimental long-term effects on their genetic viability, because they may be unable to use narrow corridors that would connect them to other subpopulations. Although judgments about what is "right" or "nat-
ural" in these examples may seem easy, not all wildlife professionals, interest groups
, or the public share the same values. These are not strictly scientific issues; they require social and political input, the focus of human dimensions research (Vaske et al. 1995). Conclusion Wildlife management is often concerned with how humans will coexist with wild animals, but there are many variations on the coexistence ideal. In an urban setting like Anchorage,Alaska, the ideal might include brown bear populations that demonstrate avoidance behavior; in a protected area like Alaska's Katmai National Park, habituation responses may be preferred. Either situation allows coexistence, but each has different consequences for bears and for people. Improved management requires better information about how human actions affect wildlife responses, as well as clarity about which coexistence ideal is appropriate for the area. In research, the need is for models that explore the complexity of wildlife responses and relate those responses to human management regimes. In management, the need is for explicit statements about the desired coexistence ideal and the development of standards that define acceptable consequenceS. Literature
cited ALBERTD,. M., AND R. T. BOWER.1991. Factors related to grizzly bear-human interactions in Denali National Park. Wildlife Society Bulletin 19:339-349. BEKOFMF,., AND D. JAMIESON19. 91. Reflective ethology, applied philosophy
, and the moral status of animals. Pages 1-32 in P. P. G. Bateson and P. H. Klopfer, editors. Perspectives in Ethology: Human Understanding and Animal Awareness, 9. Plenum Press, New York, New York. BONNEJR. T, . 1980. The evolution of culture in animals. Princeton University
Press, Princeton, New Jersey. B R I ~ T I N GMH.~CM.,,AND S. A. TEMPLE1.988. Impacts of supplemental feeding on survival rates of black-capped chickadees. Ecology 69:581-589. B ~ I T R ODN. ,1983. Variability in the responses of black-billed magpies to natural predators. Behaviour 78:209-236. BURGEJR.,,AND hi.GOCHFEI.D19.91. Human distance and birds: tolerance and response distances of resident and migrant species in India. environmental conservation
18:158-165. CUIGHEAJD. J, ., J. S. SUMNEARND, J. A. MITCHEL1L9.95. The grizzly bears of Yellowstone: their ecology in the Yellowstone ecosystem, 1959-1992. Island Press, Washington, D.C.
DOMJAMN,., AND B. BURKHAR1D98. 6. The principles of learning and behavior. Second edition. Brooks/Cole Publishing Company, Monterey, California. D o ~ c ; l ~ s - H m i r ~I..~, AoN~D, 0. DOUGIAS-HAI\~II:1T9O7N5.. Among the elephants. Collins and Harville, London, England. EIBL-EIBESPEI.I.D1T97, 0. Ethology, the biology of behavior. Holt, Rinehart, and Winston, New York, New York. FLEISCHMPA. N1,969. Conservation: the biological fallacy. Landscape 2:23-26.
In my opinion Whittaker and Knight 317
FRASEJR. ,D. 1984. The impact of humans on bald eagle populations-a review. Pages 68-94 in J. M. Gerard and T. M. Ingram, editors. The bald eagle in Canada. Proceedings of Bald Eagle Days, Winnipeg, Saskatchewan, Canada
. GEISTV,. 1971. Mountain sheep: a study in behavior and evolution. University of Chicago
Press, Chicago, Ill
inois. GEISTV,. 1978. Behavior. Pages 283-296 in J. L. Schmidt and D. L. Gilbert, editors. Big Game of North America: Ecology and Management. Stackpole Books
, Harrisburg, Pennsylvania. GEORGJE. ,L. 1982. Urban Wildlife: a problem analysis for environmental forestry research. Agricultural Experiment Station, The Pennsylvania State University
, State College. GILBERBT.,K. 1989. Behavioral plasticity and bear-human conflicts. Pages 1-8 in Bear-People Conflicts. Proceedings of a Symposium on Management Strategies. Northwest Territories Department of Renewable Resources, Whitehorse, Canada. GOODALJ.L1, 986. The chimpanzees of Gombe: patterns of behavior. Belknap Press of Harvard University
Press, Cambridge, Massachusetts. GROVEPS., M., AND R. F. THOMPSON19. 70. Habituation: a dual process theory. PsychologicalReview 77:419-450. GRUMBIRN.EE,. 1992. Ghost bears: exploring the biodiversity crisis. Island Press, Washington, D.C. HAILMAJ.NP, . 1967. The ontology of an instinct. Behaviour 15:l-159. HERRERSO. ,1985. Bear attacks: their causes and avoidance. Winchester Press, Piscataway, New Jersey. HUMPHRGE.Y,1930. Le Chatelier's rule and the problem of habituation and dehabituation in Helix albdabris. Psychological Forsch 13:113-127. IMMELMAKN.,NAN, D C. BEER.1989. A dictionary of ethology. Harvard University Press, Cambridge, Massachusetts. JOPEK, . L. 1985. Implications of grizzly bear habituation to hikers. Wildlife Society Bulletin 1332-37. KLOPFEPR.,H. 1974. An introduction to animal behavior: ethology's first century. Second edition. Prentice-Hall Inc., Englewood C l i s , New Jersey. KNIGHTR,. L., AND D. P. ANDERSE1N9.90. Effects of supplemental feeding on an avian scavenging guild. Wildlife SocietyBulletin 18:388-394. KNIGHRT.,L., D. P. ANDERSMEN. J,. BECHARADND, N. V. MARR1. 989. Geographic variation in nest-defence behaviour of the redtailed hawk (Buteojamaicensis). Ibis 131:22-26. KNIGHTR,. L., AND D. N. COLE.1991. Effects of recreational activity on wildlife in wildlands. Transactions of the North American
Wildlife and Natural Resource Conference 56:238-247. KNIGHRT.,L., D.J. GROUTAN, D S. A. TEMPLE19. 87. Nest defensebehavior of the American crow in urban and rural areas
. Condor 89:175- 177. KNIGHTR., L., AND S. K. KNIGHT.1984. Responses of wintering bald eagles to boating activity. Journal of Wildlife Management
48:999- 1004. KNIGHT,R. L., AND S. A. TEMPLE19. 86. Why does intensity of avian nest defense increase during the avian cycle? The Auk 103:318-327. KNIGHTR,. L., AND S. A. TEMPLE19. 95. Origin of wildlife responses to recreationists. Pages 81- 91 in R.L. Knight and K. J. Gutzwiller, editors. Wildlife and Recreationists: Coexistence though Management and Research. Island Press, Washington D.C.
LOFSTROIM., A, ND S. STALUNG1S9.73. Aggression in Chimpanzees: Has it become more common? Pages 217-233 in P. P. G. Bateson and P. H. Klopfer, editors. Perspectives in Ethology: Human Understanding and Animal Awareness, 9. Plenum Press, New York, New York.
MANNINAG.,,AND M. S. DAWNS.1992. An introduction to animal behavior. Fourth edition. Cambridge University Press, New York, New York. MCCULL~UDG. HR., 1982. Behavior, bears, and humans. Wildlife Society Bulletin 10:27-33. MCCUTCHEHN.E,. 1989. Cryptic behavior of black bears in Rocky Mountain National Park. InternationalConference on Bear Research and Management 8:65-72. MCGREWW, . C., C. E. TUTLINA,ND P. J. BALDWIN1.979. Chimpanzees, tools, and termites: Cross-cultural comparisons of Senegal, Tanzania, and Rio Mundi. Man 14:185-214. NEWTONI,. 1979. Population ecology of raptors. Buteo Books, Vemillion, South Dakota. OLSONT,. L., B. K. GILBERATN,D R. C. SQUIBB1.997. The effects of increasing human activity on brown bear use of an Alaskan River. Biological Conservation 82:95-99. ORIANGS,. H. 1981. Foraging behavior and the evolution of discriminatory abilities. Pages 389-406 in A. C. Kamil and T. D. Sargent, editors. Foraging behavior: ecological, ethological, and psychological approaches. Garland Press, New York, New York. POOLEA,. 1981. The effects of human disturbance on osprey reproductive success. Colonial Waterbirds 4:20-27. SHELBBY., AND T. A. HEBERLEI1N9.86. Carrying capacity in recreation settings. Oregon State University
Press, Corvallis. STEINBEJC.,KA,ND E. F. RICKEITS. 1941. Sea of Cortez: A Leisurely journal of travel and research. The Viking Press, New York, New York. THORPWE,. H. 1956. Learning and instinct in animals. Methuen and Company, New York, New York. THURBJE.RM, R. 0 . PEERSOTN.,D. DRUMMEANRD, S. A. THOMASMA. 1994. Gray wolf response to refuge boundaries and roads in Alaska. Wildlife Society Bulletin 22:61-68. VASKEJ., J., DECKEDR., J., AND MANFREDMO. J, . 1995. Human dimensions of wildlife management: an integrated framework for coexistence. Pages 33-50 in R. L. Knight and K. J. Gutzwiller, editors. Wildlife and recreationists: coexistence through management and research. Island Press, Washington D.C. Doug Whittaker (photo) is pursuing a Ph.D. in human dimensions of natural resources at Colorado State University. His research interests focus
on river management issues and the human dimensions of wildlife. Richard (Rick) 1. Knight is professor of wildlife ecology at Colorado State University. His Ph.D. is from the University of Wisconsin
0049821 http://www.jstor.org LINKED CITATIONS - Page 1 of 2 - You have printed the following article: Understanding Wildlife Responses to Humans Doug Whittaker; Richard L. Knight Wildlife Society Bulletin, Vol. 26, No. 2. (Summer, 1998), pp. 312-317. Stable URL: http://links.jstor.org/sici?sici=0091-7648%28199822%2926%3A2%3C312%3AUWRTH%3E2.0.CO%3B2-8 This article references the following linked citations. If you are trying to access articles from an off-campus location, you may be required to first logon via your library web site to access JSTOR. Please visit your library's website or contact a librarian to learn about options for remote access to JSTOR. Literature cited Factors Related to Grizzly Bear: Human Interactions in Denali National Park David M. Albert; R. Terry Bowyer Wildlife Society Bulletin, Vol. 19, No. 3. (Autumn, 1991), pp. 339-349. Stable URL: http://links.jstor.org/sici?sici=0091-7648%28199123%2919%3A3%3C339%3AFRTGBH%3E2.0.CO%3B2-I Impacts of Supplemental Feeding on Survival Rates of Black-Capped Chickadees Margaret Clark Brittingham; Stanley A. Temple Ecology, Vol. 69, No. 3. (Jun., 1988), pp. 581-589. Stable URL: http://links.jstor.org/sici?sici=0012-9658%28198806%2969%3A3%3C581%3AIOSFOS%3E2.0.CO%3B2-1 Implications of Grizzly Bear Habituation to Hikers Katherine L. Jope Wildlife Society Bulletin, Vol. 13, No. 1. (Spring, 1985), pp. 32-37. Stable URL: http://links.jstor.org/sici?sici=0091-7648%28198521%2913%3A1%3C32%3AIOGBHT%3E2.0.CO%3B2-1 Effects of Supplemental Feeding on an Avian Scavenging Guild Richard L. Knight; David P. Anderson Wildlife Society Bulletin, Vol. 18, No. 4. (Winter, 1990), pp. 388-394. Stable URL: http://links.jstor.org/sici?sici=0091-7648%28199024%2918%3A4%3C388%3AEOSFOA%3E2.0.CO%3B2-Q
0049822 http://www.jstor.org LINKED CITATIONS - Page 2 of 2 - Nest-Defense Behavior of the American Crow in Urban and Rural Areas Richard L. Knight; Daniel J. Grout; Stanley A. Temple The Condor, Vol. 89, No. 1. (Feb., 1987), pp. 175-177. Stable URL: http://links.jstor.org/sici?sici=0010-5422%28198702%2989%3A1%3C175%3ANBOTAC%3E2.0.CO%3B2-Y Responses of Wintering Bald Eagles to Boating Activity Richard L. Knight; Susan K. Knight The Journal of Wildlife Management, Vol. 48, No. 3. (Jul., 1984), pp. 999-1004. Stable URL: http://links.jstor.org/sici?sici=0022-541X%28198407%2948%3A3%3C999%3AROWBET%3E2.0.CO%3B2-8 Behavior, Bears, and Humans Dale R. McCullough Wildlife Society Bulletin, Vol. 10, No. 1. (Spring, 1982), pp. 27-33. Stable URL: http://links.jstor.org/sici?sici=0091-7648%28198221%2910%3A1%3C27%3ABBAH%3E2.0.CO%3B2-1 Chimpanzees, Tools, and Termites: Cross-Cultural Comparisons of Senegal, Tanzania, and Rio Muni W. C. McGrew; C. E. G. Tutin; P. J. Baldwin Man, New Series, Vol. 14, No. 2. (Jun., 1979), pp. 185-214. Stable URL: http://links.jstor.org/sici?sici=0025-1496%28197906%292%3A14%3A2%3C185%3ACTATCC%3E2.0.CO%3B2-B The Effects of Human Disturbance on Osprey Reproductive Success Alan Poole Colonial Waterbirds, Vol. 4. (1981), pp. 20-27. Stable URL: http://links.jstor.org/sici?sici=0738-6028%281981%294%3C20%3ATEOHDO%3E2.0.CO%3B2-Q Gray Wolf Response to Refuge Boundaries and Roads in Alaska Joanne M. Thurber; Rolf O. Peterson; Thomas D
. Drummer; Scott A. Thomasma Wildlife Society Bulletin, Vol. 22, No. 1. (Spring, 1994), pp. 61-68. Stable URL: http://links.jstor.org/sici?sici=0091-7648%28199421%2922%3A1%3C61%3AGWRTRB%3E2.0.CO%3B2-O