How to Obtain Behavioral and Ecological Data from Free-Ranging, Researcher-Habituated... Bears Author(s): Lynn L. Rogers and Gregory W. Wilker
Transcription
How to Obtain Behavioral and Ecological Data from Free-Ranging, Researcher-Habituated... Bears Author(s): Lynn L. Rogers and Gregory W. Wilker
How to Obtain Behavioral and Ecological Data from Free-Ranging, Researcher-Habituated Black Bears Author(s): Lynn L. Rogers and Gregory W. Wilker Source: Bears: Their Biology and Management, Vol. 8, A Selection of Papers from the Eighth International Conference on Bear Research and Management, Victoria, British Columbia, Canada, February 1989 (1990), pp. 321-327 Published by: International Association of Bear Research and Management Stable URL: http://www.jstor.org/stable/3872935 Accessed: 03/01/2009 20:39 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=iba. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact [email protected]. International Association of Bear Research and Management is collaborating with JSTOR to digitize, preserve and extend access to Bears: Their Biology and Management. http://www.jstor.org HOWTO OBTAINBEHAVIORAL AND ECOLOGICAL DATAFROMFREEBLACKBEARS RANGING,RESEARCHER-HABITUATED LYNN L. ROGERS, U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station, Kawishiwi Field Laboratory, SR 1,Box 7200, Ely, MN 55731 GREGORY W. WILKER,U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station, Kawishiwi Field Laboratory, SR 1, Box 7200, Ely, MN 55731 Abstract: A study was conducted in northeastern Minnesota from 1986-1989 to determine the feasibility of habituating black bears (Ursus americanus) to observers for behavioral and ecological research. Of 18 males and 8 females that repeatedly visited an artificial feeding site in the presence of humans, 3 2-year-old females were radiocollared for further habituation away from the site. The 3 females were repeatedly located, fed, and accompanied at various locations in their territories. After 50-100 hours of this additional contact, each bear accepted human presence and mostly ignored observers that followed 1-10 m behind. The bears were no longer fed in their territories except for the feeding of scat markers for digestion studies. The bears were observed for 24- or 48-hour periods approximately weekly as they matured, reproduced, and raised cubs. While being followed, they foraged, napped, slept through nights, showed REM and non-REM sleep, mated, played, nursed their cubs, captured young animals, maintained territories, marked bear trees, prepared dens, and began hibernation. They relied on natural foods and showed activity and movement patterns similar to daily and annual patterns of 103 non-habituated bears that were radio-tracked previously in approximately the same area. A field computer with an internal clock was programmed to aid in the recording of activities, habitat use, food consumption, and weather. A personal computer was programmed to sort and tabulate the data and calculate I) time spent in each activity, 2) time spent in each cover type, 3) number of bites taken of each food in each cover type, and 4) time spent in each activity and cover type in each weather condition. The computer calculated these time-activity budgets for single 24-hour observation periods or for any combination of observation periods specified. Problems of excessive hunting loss, habitual nuisance behavior, or appreciable observer injury did not occur. Study results were directly useful to forest managers for identification of opportunity areas for habitat preservation or improvement. Results provided new insights into black bear diet, bioenergetics, foraging strategies, activity patterns, sociobiology, communication, and bear-human interactions. Drawbacks were few study animals and limited study area. Comparative studies are needed in new locations, involving additional age and sex classes, additional physiological data, and additional behavior regimes such as might be exhibited by translocated bears, nuisance bears, or bears whose ranges have been altered by fire, development, insect defoliation, or extensive timber management. Int. Conf. Bear Res. and Manage. 8:321-327 Ecological studiesof elusive, wide-rangingmammals are often hamperedby the inability to observe study animals closely for significant periods. Consequently, researchershave studied wide-rangingspecies in large enclosures (McMahan 1964, Dzieciolowski 1967, LeResche and Davis 1973, Stringham1974, Ozoga et al. 1982,andothers),on leashes(Wallmo 1964,Watts1964, Healy 1967, Neff 1974, Bauer 1977, Crawford 1982, Lautenschlagerand Crawford 1983, Thill and Martin 1989, and others), or by temporarilyreleasing tame animals (Buechner 1950). Otherresearchershave observedwild, free-ranginganimalsthatwerehabituatedto vehicles (Schaller 1972, Bertram1973, Douglas-Hamilton 1973) or people (Graf 1955; Brown 1961; Goodall 1965;Geist 1971; Sade et al. 1976;Fossey and Harcourt 1977;Chepko-SadeandSade 1979;Rogers 1981, 1987a; Rogerset al. 1987; Mech 1988). Eachstudymethodhas its advantages,dependingupon the researchquestions. During1985-1989,we developedmethodsfor habituatingfree-rangingblackbearsto observersandfor using computersto recordandanalyzedetailedbehavioraland ecological data rapidly. Areas of study includedblack bear habitat use, diet, bioenergetics, activity patterns, sociobiology, behavior, communicationmethods, and bear-humaninteractions. The objective was a better understandingof blackbearenvironmentalrequirements and behaviorto aid decision-makingby forest managers andcampgroundmanagersandto increasepublicunderstandingof black bears. This paperdiscusses methods, benefits, and shortcomingsof the study. We thankC. Cowden,P. Miles, V. O'Connor,S. Paul, and S. Scott for help in developing techniquesand collecting data; Kawishiwi Ranger District personnel for relaying 2-way radio transmissions;SuperiorNational Forestchief biologistE. Lindquistandpilots C. Palumbi, D. Bowman, and S. Durst for help in radio-tracking particularlywide-rangingbears;C. Robbins,G. Pritchard, andD. Hewittfor help withdigestivestudies,residentsof the study area for their interest and support;and R. Archibald,L. Darling, K. Elowe, B. Peyton, and two anonymous reviewers for thoughtful reviews of the manuscript. STUDY AREA The study was conducted in the SuperiorNational Forestin northeasternMinnesota(47049'N, 91 45' W). The primarystudyareawas the contiguousterritoriesof 3 principal study animals and was 6 km in diameter. Longermovementsduringlate summer,fall, and estrus increasedthe overall studyareato approximately24 km by 34 km. The primarystudyareacontaineda networkof logging roads,a paved 2-lane highway, a 32-unitcampground, 24 summerhomes, and the researchfacilities. The homes and researchfacilities were all located on KawishiwiRiverandBirchLake,a waterway150-650m wide thatformedthe westernedge of the study area. The study area was almost entirely forested and had gently rolling terrain with occasional rock outcrops. Mixed coniferous-deciduousforest predominated(Rogers 1987b). Standsrangedin age fromnewly clearcutto 322 BEARS-THEIR BIOLOGYAND MANAGEMENT virginstands>100-years-old.Wetlandsincludedstreams and beaverimpoundments,scatteredbogs and marshes, anda 2.5-halake,each typicallyborderedby alder(Alnus rugosa) swamps. Bear foods available in season between May and Septemberincluded wild fruit, beaked hazelnuts (Coryluscornuta),digestiblevegetation,andHymenopteran insects (Rogers 1987b). Wild fruit was available from, July to early September,if crops were successful, and included wild strawberry(Fragaria spp.), dwarf red raspberry(Rubuspubescens),serviceberry(Amelanchier spp.), lowbush blueberry (Vaccinium angustifolium), sourtop bilberry (V. myrtilloides),pin cherry (Prunus pensylvanica),choke cherry(P. virginiana),wild sarsaparilla(Aralia nudicaulis),red raspberry(R. strigosus), red-osierdogwood (Cornusstolonifera),roundleafdogwood (C. rugosa), highbush cranberry (Viburnum trilobum), and mountain-ash(Sorbus spp.). The only hardmastwas beakedhazelnuts,which were availablein AugustandearlySeptemberin occasionalyears. Acorns (Quercus spp.) and other fall mast species were essentially nonexistent. Bears entered dens in October and emergedin April. Snow was presentfromlateOctoberor Novemberto April. METHODS Habituating Black Bears To Observers To habituatebearsto humanpresence,researchersestablisheda beef fatfeeding site, whichwas visitedrepeatedly by 18 males and8 females.Within4 bearvisits, eachbear becameaccustomedto the presenceof a recliningperson within30 m. Within8 visits, mostreadilytookfood from, or within5 m of, an immobileor slowly moving person. Personality,wariness,and time requiredfor habituation variedwithindividuals.Veryfew becameaccustomedto being touched.All became accustomedto talking. Two adults, a male and a female, remaineddefensive after2 and 10 weeks of occasional encounters and were not studied closely. Three other females (401, Patch, and Terri), all 2-years-old, were more receptive and were capturedand radio-collaredfor furtherstudy away from the feeding site. These 3 females were initially timid when they were approachedin new areas.Theyretreated,usuallyunseen, for up to 2 hoursbeforecirclingdownwindandstopping 40-80 m away to smell the person and the beef fat he carriedandto listen to his voice. After<30 minutes,the bears approached,often demonstratingtheir uneasiness by lunging, blowing, and slapping vegetation before calming down and resumingtheir familiarfeeding rou- tines. Over the next 50-100 hoursof contact,each bear became accustomedto being approached,fed, and accompaniedin the forest. Supplementalfeeding was then stopped.The bearscontinuedto approachobserversfor food butreadilylearnedthatopen,emptyhandsmeantno food. After a food-check, the bears typically resumed foraging and ignored observers. However, if food was hiddenin pocketsorbackpacks,thebearsoftencontinued theirattentionorreturnedrepeatedlyto observers.Therefore,no food was broughtduringdatacollectingsessions. Observersfollowed <10 m behindforagingbears. At greaterdistances,the bears sometimes lost trackof observers and became wary of their rustlings. Bears also sometimesbecame wary when observersapproachedto within a few meters, although this wariness waned as bears became more habituated. An intent look from a bearindicatedits concernoveranobserver'sproximityor actions. Bears sometimesgave "threat"displays (lunging, slappingvegetation,blowing) whenobserversgot in theirway in unusuallyfood-richareas. Most startledor defensive reactions (jumping away, whirling around, startingup trees) were due to sudden,seemingly aggressive acts by observers(sneezing, tripping,falling, accidentallyhittingthe bear)or by the suddenappearanceof anextraobserverin densebrushor on an overheadledge. Therefore,data collecting sessions were conductedby lone observers. "Threat"displayswere actuallyexpressionsof fearor uneasiness ratherthan threats. No bear attacked,and bearsretreatedif an observeracted aggressively. Bears were calmed by giving them more room and by talking. Although difficult to describe, it became easy to read bears'expressionsandanticipatetheirmovements.Thus, observersavoided causing "threat"or defensive behaviors. Withtime (100-150 hours),mutualtrustdeveloped, "threats"all but disappeared,and the fully habituated bearsforaged,slept,andappearedto be comfortablewith a single observer1-5 m behindor occasionallycloser to see certainfoods. Sensitive, experiencedobserverssaw <1 "threat"per 100hoursof observation,withmost of the threatsoccurringin dusk or darknesswhen it was more difficult to anticipatebears' movements. Observersuseddimflashlightsto aidobservationsand computerentries at night. The bears typically slept or nursedfrom 1-2 hoursaftersundownto 0.5 hoursbefore sunrise. Observersspent nights in sleeping bags 1-8 m from the sleeping bears. To begin a new year of observations, researchers reinforcedthepreviousyear'shabituationby restingnear dens and day beds before extensive movement began. This presence was especially importantwith newborn BLACKBEARS* Rogers and Wilker STUDYINGRESEARCHER-HABITUATED litters because cubs developed wariness of strangersin late MarchandearlyAprilaroundthe time of emergence (Rogers 1989). Prolongedhandlingof cubs duringthat time (the thirdmonthof life) furtherhabituatedthem to people (Rogers 1989 and unpublisheddata), making them less likely to distracttheir mothers with distress calls when observersaccompaniedthem later. The artificialfeedingsite was essentiallydiscontinued in 1988 and 1989. Bearuse of the feeding site hadwaned by then, and observerswere concentratingon obtaining datafromhabituatedbearsratherthanon habituatingnew ones. Only 1 habituatedfemale (Terri)visited the site in 1988 and 1989 because Bear 401 had been thrownfrom a treeby anotherfemale andkilled on 10 June 1987, and Patchhad shifted her territory3 km away by the end of 1987 and did not returnin 1988. She was killed by a hunter16 km away on 5 September1988. Terri'suse of the site in 1988 and 1989 was limitedto a few visits each year in late May (after springforbs became unpalatable but before ant pupaebecame abundant)and in early fall (afterberriesand hazelnutsdisappeared).On each visit, she was given a few grams of food to entice her onto a platformscale, and she was given more food duringthe fall 1988 huntingseason to diverther from any hunters' baits and to increase the chance of her producingcubs (which she did) for study of mother-cubinteractions. Data Collection Procedures Field equipment(Table 1) includeda Husky Hunter field computer (Husky Computers,Inc., Sarasota,FL 34236), which is a small, immersible, shock resistant computerworn in a harnessthatleaves both handsfree. The computerwas programmedto acceptentrycodes for describing activity, food consumption, habitat, and weather(Rogers et al. 1989a). A clock in the computer automaticallyrecordeddatesandtimesof entries.Activities recordedincludedwalking, sitting, standing,reclining, running,sleeping, climbing, defecating, urinating, digging, grooming, scent-marking,nursing,vocalizing, playing, drinking, bathing, wading, and eating. For eating, numberof bites of each food was recorded. Data were recordedcontinuouslyduringobservation periodsof 24 or48 hours. Twenty-four-hourobservation periods providednatural,around-the-clocktime frames for collecting dataon activity,food consumption,habitat use,andweatherandwererepeatedapproximatelyweekly to keep pace with changes in phenology. Forty-eighthour observationperiods were used when digestibility data were collected in additionto the above data. The longer observationperiod was needed to collect fecal droppingscontaining foods observed eaten during the 323 Table 1. Equipmentneeded for extended bear observations. 1. Shock resistant,immersible,field computerwith harnessthat leaves handsfree. AA batteriesfor change at 24 hours. 2. I4. Backpackwith no framestructureto catch on vegetation. Lightweightreceiver(in ziploc bag to preventraindamage) and a folding mini-antenna. Two-way radio. 5. Small transmitterfor locating the observerin case of injury. i 6. Scat marker(if doing digestion studies-see text). 7. Pocket-sizednotebook. 8. Habitatforms for describinghabitat. 9. Two dozen gallon-sized, prelabelled,datedziploc bags for collecting unknownfoods and (if doing digestion studies) scats. 3. 10. Two ballpointpens with waterproofink for takingnotes, filling out habitatforms, and writingcollection times on scat bags. 11. Compass,laminatedtopographicmap, and laminatedaerial photo. 12. Two quartsof drinkingwaterin collapsible goatskincontainers. (As the wateris used, more room becomes availablefor scats). 13. Insect repellent. 14. Light sleeping bag in a compressionsack; waterproofshell for the sleeping bag (if observingovernight). 15. Dry socks and t-shirtin a ziploc bag (if observingovernight). 16. Flashlightwith fresh batteriesand an extrabulb. 17. Plastic trashbag for keeping the above items dry in backpack when swimming across streams. 18. Rain suit (optional). first 24 hours. Passage time typically was 4-12 hours. Foodseatenduringthe first24 hoursof observationwere identifiablein scats because a colored scat markerwas fed at the beginning of an observation period and a different colored marker was fed at 24 hours. Scats collected between passage of the markerscontainedthe first 24 hours' consumption. Scat markers included colored beads, corn, peanuts,pieces of colored balloon, or combinationsof these items. After it was determined that passage rate was essentially the same for all these materials,pieces of coloredballoons were used because they were most easily obtainedand most easily seen in scats. Approximately 10-20 balloon pieces, each 12 cm2,were fed in a 50-grampiece of fat. The 48-hourobservationperiodswere conductedin 4 consecutive 12-hour shifts. The 24-hour observation periodswere conductedin 2-3 consecutiveshifts of 8-12 hourseach or in 1 24-hourshift. The latterwas possible becauseobserversrestedwhile the bearsslept at nightor napped in daytime. Longer shifts were precluded by 324 BEARS-THEIR BIOLOGY AND MANAGEMENT hungerand the weight of collected scats. Observationperiods were begun, and shift-changes made, only when bears were awake, so sleep-activity cycles were not disturbed.Activity was detectablefrom telemetry signals. The incoming observer located the bearby telemetryandthen waited200-300 m away until telemetry signals indicatedactivity or the observer alreadywith the bearinformedhim by 2-way radiothatthe bear was active. The incoming observertalked the last 100-200 m to identify himself. Habitat Description A goal in collecting habitatuse datain this studywas to provide national forest managers with information useful in habitatmanagementdecisions. Forthatreason, habitatswereclassifiedin U.S. ForestServiceterms-by forest type, tree size, and tree density. In addition,Ecological LandType (ELT),silviculturalhistory,andnaturaldisturbances(suchas insectdefoliation,winddamage, fire, old-growthforests breakingup) were recordedto determinefactors that influence habitatquality. However, not all of this informationwas immediatelyevident uponenteringa new foresttype. Consequently,observers recordedonly the time when entering a stand and recordeddescriptiveparameterswhile moving through it. To understandfully why bearsused particularforest types,it was necessaryto describenotonly the foresttype in generalbut the microhabitats(or forest components) thatwere used within thattype. To recordmicrohabitat use, computercodes were created to denote common microhabitats(e.g., the differenttypesof forestopenings, rock crevices, thickets, certain tree species, old dens, game trails)andwere used to denotemicrohabitatsassociatedwith recordedactivities. Thus,each activityentry hadanaccompanyingmicrohabitatentryif themicrohabitat differedfrom the generalforesttype. Forexample,if an importantcomponentof a foresttype was blueberries growingon patchesof shallow,sunlitsoil atedges of rock outcrops,computerentrieswould indicate,in code, that the bear,e.g., ate 234 bites of blueberriesfrom edges of rockoutcrops.An objectivewas to understandwhy bears used particularhabitatsat particulartimes (time of day, time of year, time of life), considering food, water, security, weather, insects, reproductive status, social constraints,and recentevents. Post-observation Handling of Data After an observationperiod was completed,the data were uploaded from the field computerto a personal computer for viewing, correction of entry errors, and printing. The computer was programmedto sort and tabulatethe data and create the following time-activity budgets(Rogerset al. 1989b): 1. Amountof time spentin each activity, 2. Amount of time spent in each vegetative cover type, 3. Numberof bites takenof each food in each cover type, and 4. Amount of time spent in each activity in each weathercondition. (Weatherwas classified by temperaturerange,humidityrange,and sky condition.) The computercalculatedthese time-activitybudgets for single 24-hourobservationperiods,for 48-hourperiods, or for any specified combination of observation periodsso changesin activitythataccompanychangesin physiology, food distribution,food abundance,weather, or other factors could be discovered. Observerswrote narrativesto describeplay, social interactions,vocalizations, predation,and otherunusualobservationsthatdid not easily fit the computerformat. Routes of bearswere mapped, including resting locations, intensive feeding locations,andhourlyprogress.Uncertaintiesaboutroutes and locationswere resolved by checkingan aerialphoto of the study area. Supplemental Studies In the days following an observationperiod,followwereconductedto determineamountsingested studies up of each food and the nutritionalvalues of those foods. After seeing hundredsof bites during an observation period, observers used their knowledge of bite size to collect 100 bite-sized samplesof each currentbearfood andweigh themfresh andoven-dried(48 hoursat 48 C). Foreach food, the weightperbite (freshandoven-dried) was multipliedby the numberof bites the bearhadeaten of thatfood in 24 hoursto estimatedaily consumptionof eachfood. This informationwas also calculatedby cover typeto determinefeedingefficiency in thedifferentcover types. The oven-driedsampleswere then sent to Washington State University for determinationsof gross energy (by bomb calorimetry),percent protein (Kjeldahl method),digestibleprotein(PritchardandRobbins1990), and digestible energy (Pritchardand Robbins 1990). Bodyweight changes were determinedby enticing the bears onto a platformscale whenever they visited the artificialfeeding site. Boundariesof territoriesandannualrangeswere estimated from the maps of 24-hour bear movements and frominterimradiolocations. Territorieswereconsidered to be areas of exclusive or nearly exclusive use by a female and her offspring. Annual ranges included all STUDYINGRESEARCHER-HABITUATED BLACKBEARS? Rogers and Wilker areasknownto be traversedin a year,includingthe sallies made during estrus and the extraterritorialforaging movementsmadeduringlatesummer.Theavailabilityof differentcover types withinterritoriesandannualranges were determinedfrom aerialphotos,U.S. ForestService and MinnesotaDepartmentof NaturalResources compartmentexam records,privatetimbercompanyinventories, anda U.S. Fish andWildlife ServiceGeographicInformationSystem (GIS) database. Use and availability werecomparedto determinehabitatpreferencesin different seasons and years. EVALUATIONOF METHODS Data Collected The habituatedbears provided over 2,000 hours of observationdata,including6 24-hourperiodsin 1987,23 24-hourperiodsin 1988, and20 24-hourperiodsin 1989. Human Influences on Bear Behavior Grossbehaviorof the 3 habituatedfemaleswas similar to thatof 103 bearspreviouslyradio-trackedin northeastern Minnesota (Rogers 1987b) with no differences in territorysize, nuisancebehavior(see below), food habits, socialrelationships,dailyactivitypatterns,seasonaltravel patterns,or denning habits. Habituatedbears further appearedto behave normally in that they nursed their cubsperiodicallythroughthedayandnightanddefecated at fairlyregularintervalsthatvariedseasonallywithfood type and abundance.Bearspursuedandcapturedyoung animals, markedsmall trees with urine, markedlarger trees by rubbing, and investigated the marks of other bears. Sleeping bears showed apparentlynormalsleep thatincludedbothREMandnon-REMsleep. REMsleep is seldom seen in poorlyhabituatedanimals(Allison and VanTwyver 1970). The bears also prepareddens and became lethargic in preparationfor hibernationwhile being accompanied. A prehiberating female cub that fell asleep with an observer'shandon her femoralartery outside her den on 12 October1989 showed a pulse rate of only 22 beats/minute. Mothers ran to distress cries of cubs 1-2 m from observersbut seemed to discountobserversas possible sourcesof danger. Habituatedbearschasedor fled from wolves (Canis lupus) or otherbears, and they remained awareand responsiveto distant,unidentifiedsoundsbut apparentlyignored the rustlings of nearby observers because the focus of bears' ears and eyes was usually in directionsotherthantowardthe observer. The direction of their muzzles while doing the special breathingthat accompaniedintense scenting also was usually directed 325 away from observers. Observationsof the ways bears sensed their environmentgave insight into the kinds of stimuli thatmost influencedtheir actions. However, some aspectsof growthandbehaviorwere influenced by the study. Habituatedbears reproduced earlier(at 3, 4, and4 yearsof age) thanmost otherbears of the region (Rogers 1987b) due to supplementalfood given to them duringhabituationefforts and September huntingseasons. Bears' foragingwas sometimes interruptedbriefly when observers got in the way. Mating activity was slightly disruptedby observers in that 4 strangemales that approachedhabituatedfemales were initially apprehensiveuntil observers stood back 1520 m. The malesthencourted,copulatedwith,andrested with the females and within 1-2 hours toleratedcloser (10-15 m) observation. A habituatedadult (Patch)and severalhabituatedcubs occasionallyinvolved observers in play, especiallywhenthey awokefromnaps. This was remediedby steppingback a few meters. Potential Problems Problemsof excessive hunting losses, habitualnuisance problems,and appreciableinjuryto observersdid not occur. No one was bitten. Habituatedmotherswere as tolerantof observersas were other bears. One bear occasionally slappedif she was teased with food or was eye-to-eye with a personon herlevel within0.5 m. Slaps that were apparentlyfull power created only welts or scratcheswithno appreciableinjurydueto thedullnessof adultblackbearclaws. No threatbehaviorswereof types used beforefights (deep-throated,pulsingvocalizations; bellowing, open-mouthedthreats;stiff approacheswith backsarchedandnoses nearlytouchingthe ground). We recognize that some danger exists in working closely with largeanimalsandthatsensitivityandgood manners are requiredto avoid provoking defensive behaviors. However,the bearsin this studyshowedrestraintandno propensityto seriously injure. No habituatedbearbecamea habitualnuisanceat any of the 24 homes within the study area or at the 32-unit campground,whichwas less than500 m fromthefeeding site. In fact, the feeding site diverted bears from the campgroundand reduced nuisance problems (Rogers, unpublisheddata). Two bearsbrieflyate dandelionsand 1 bearlicked grease from a disposal areain yardsof the homes. Nuisancebehaviorappearedto be more a functionof homerangelocationandnaturalfood scarcitythan familiaritywith people. When fruit, nuts, or ant pupae wereabundant,habituatedbearssometimesforagedwithin 200 m of homes, the campground,or the feeding site without checking for food or being seen. Most of the 326 BEARS-THEIR BIOLOGY AND MANAGEMENT homeownersdevelopeda protectiveinterestafterseeing the bearsand being informed(wheneverpossible) about new findings. Mortalityof habituatedbearsduringhuntingseason(1 Sep-mid-Oct) did not exceed mortality in the overall population, possibly due to partial protection by researchers.Habituatedbearswere studiedthrough7 bearyears of hunting seasons with only 1 hunting death, showing a slightly lower mortality rate than the 16% hunting mortalityreportedfor other radio-collaredfemalesin Minnesotaduringthe sameperiod(D. Garshelis, Biologist, MinnesotaDep. Nat. Resour.,pers. commun. 1989). Of the 3 radio-collaredbears, 1 is still alive, 1 was killed by a largerfemale and 1 was killed by a hunter 16 km outside the bear's territory. In 1989, observers accompaniedthe last survivinghabituatedadult during most daylighthoursof the huntingseason until she and her cubs enteredprehiberation lethargyin late September. Benefits of Habitat Studies Diet and habitatuse data from the habituatedbears were directlyuseful to local forestmanagersin identifying opportunityareasfor blackbearhabitatpreservation orimprovement.Ona broaderscale, thenew information aided developmentof a habitatsuitabilityindex model forblackbearsof theregion(RogersandAllen 1987)and, hopefully, will aid in refining the model. Some of the informationprovided by habituatedbears has proven difficult to obtain in other ways. For example, a commonly eatensucculent,jewelweed (Impatienscapensis), has not been identifiedin scats in this or previousstudies due to its high digestibility. Grassesin springdiets have not previously been identified as to species or source covertypesbecausegrasseswithoutseed headscannotbe identified in scats. In a previous study in this region (Rogers 1987b),intensivespringuse of blackash (Fraxinus nigra) swamps and alderswamps by bearsfeeding onbluejointgrass(Calamagrostiscanadensis)wasmissed because feeding in those cover types is so efficient and brief that bears were seldom radio-located in them. Habituatedbears also revealed the importanceof large refugetrees (e.g., Pinus strobus>50 cm dbh)to mothers and cubs in spring, confirming an earlier finding by Elowe (1987). Habituatedbearsprovidedthe firstinformationon amountseaten in the wild and how consumptiondiffersbetweenyearsandwithphenologicalprogression within years. Research Needs Majordrawbacksof thisstudywerefew studyanimals and the limited study area. Comparativestudies using habituatedbearsin ecologically differentareasacrossthe continentare needed. Such studies should providenew insights into bear behavior and habitatneeds and may explain regional differences in black bear growth and reproductiondueto possibleregionaldifferencesin feeding efficiency or diet quality. Anothershortcomingof thepresentstudywas the lack of intensive study of males. Few males remainedat the artificialfeeding site long enough for thoroughhabituation. Males born in the study area dispersed despite abundantfood at the feeding site, and other subadults moved on aftera few days or weeks. Maturemales that visited the feeding site duringmating season appeared more interestedin estrousfemales thanfood and did not returnlater. Thereis a need to determinedifferencesin foragingstrategiesof large-bodied,wide-rangingmales comparedto smallerbodied,territorialfemales. Studies of bearsthatmightexhibitadditionalor alteredbehavior regimes such as translocatedbears, nuisancebears, and bears whose ranges have been extensively changed by fire, development,insectdefoliation,or extensivetimber managementarealso needed. Studiesof habituatedbears with implantedphysiologicaltransmitters(to determine heartrate,body temperature,etc.) areneededfor a better understandingof bioenergeticsandthe factorsthatinfluence habitatselection. Studiesof habituatedbearsdo not eliminatethe need for extensive studies of large samples of radio-collared bears to learn age-specific and sex-specific movement patternsand survival rates. Habituatedbear studies are best used for habitatuse studiesandto adddetailsto data patternsobtainedby moreextensivemethods. Nevertheless, in some situations and locations (e.g., the cloud forests of South America)habituatedbear studies (e.g., studiesof spectacledbears[Tremarctosornatus])maybe more immediately feasible than more extensive and expensive studies (B. Peyton, Bear Specialist, InternationalUnion for the Conservationof NatureandNatural Resources [IUCN], pers. commun. 1989). Habituatedbearstudiesarelaborintensivebutcan be accomplished at relatively low cost with few radiocollars and minimal flying time. Computerprograms developedin this studyareavailableto researchersinterested in similarstudies of any species. Researchersare welcome to visit the study area and accompany the habituatedbears to become acquaintedwith field techniques and computerprograms. STUDYINGRESEARCHER-HABITUATED BLACKBEARS* Rogers and Wilker LITERATURE CITED ALLISON, T., ANDH. VANTWYVER.1970. OZOGA, J.J., The evolution of sleep. Nat. Hist. 79:59-65. W.B. 1977. Forage preferencesof tame deer in the aspen type of northernMichigan. M.S. Thesis. Mich. Technol. Univ., Houghton. 85pp. BERTRAM,B.C.R. 1973. Lion populationregulation.EastAfr. Wildl. J. 11:215-225. BROWN,E.R. 1961. The black-taileddeerof westernWashington. Wash. State Game Dep. Biol. Bull. 13. 124pp. BUECHNER,H.K. 1950. Life history,ecology, andrangeuse of thepronghornantelopein Trans-Pecos,Texas. Am. Midl. Nat. 43:257-354. BAUER, CHEPKO-SADE, B.D., ANDD.S. SADE. 1979. Patterns of group splitting within matrilinealkinship groups: a study of social group structurein Macaca mulatta (Cercopithecidae: Primates). Behav. Ecol. and Sociobiol. 5:67-86. H.S. 1982. Seasonalfood selectionanddigestibilCRAWFORD, ity by tame white-taileddeer in centralMaine. J. Wildl. Manage. 46:974-982. R. 1967. Food of the red deer in an annual DZIECIOLOWSKI, cycle. Acta Theriologica 12(36):503-520. DOUGLAS-HAMILTON,I. 1973. On the ecology and behaviour of theLakeManyaraelephants.EastAfr.Wildl.J. 11:401403. K.D. 1987. Factorsaffectingblack bearreproductive ELOWE, success andcub survivalin Massachusetts.Ph.D. Thesis. Univ. of Mass., Amherst. 7 pp. FOSSEY,D., AND A. HARCOURT. 1977. Feeding ecology of free- rangingmountaingorilla(Gorillagorilla beringei). Pages 415-447 in T. Clutton-Brock,ed. Primateecology. Academic Press, New York. GEIST, V. 1971. Mountain sheep: a study in behavior and evolution. Univ. of ChicagoPress, Chicagoand London. 383pp. GOODALL, J. 1965. Chimpanzees of the Gombe Stream Reserve. Pages 425-481 in I. DeVore, ed. Primate behavior:field studiesof monkeysandapes. Holt,Rinehart, and Winston, New York. GRAF, W. 1955. The Rooseveltelk. PortAngeles, Wash.:Port Angeles Evening News. 105pp. HEALY, W.M. 1967. Foragepreferencesof captive deer while free ranging in the Allegheny National Forest. M.S. Thesis. Pa. State Univ., UniversityPark. 93pp. LAUTENSCHLAGER, R.A., AND H.S. CRAWFORD. 1983. Haltertrainingmoose. Wildl. Soc. Bull. 11:187-189. LERESCHE,R.E., AND J.L. DAVIS. 1973. Importance of nonbrowsefoods to moose on theKenaiPeninsula,Alaska. J. Wildl. Manage. 37:279-287. MCMAHAN,C.A. 1964. Comparativefood habitsof deer and threeclasses of livestock. J. Wildl. Manage. 28:798-808. MECH, L.D. 1988. Artic wolf. Voyageur Press, Stillwater. 128pp. D.J. 1974. Foragepreferencesof trainedmuledeeron the NEFF, Beaver Creek Watersheds. Ariz. Game and Fish Dep. Spec. Rep. 4. 61pp. L.J. VERME, AND C.S. BENZ. 327 1982. Parturitionbehaviorandterritorialityin white-taileddeer: impacton neonatalmortality.J. Wildl. Manage. 46:1-11. PRITCHARD,G.T., AND C.T. ROBBINS. 1990. Digestive and metabolicefficiencies of grizzly and black bears. Can.J. Zool. (in press). ROGERS,L.L. 1981. Walking with deer. Pages 29-34 in Records of North Americanbig game. The Boone and CrockettClub. Alexandria,Va. 409pp. 1987a. Seasonal changes in defecationrates of freerangingwhite-taileddeer. J. Wildl. Manage. 51:328-331. . 1987b. The effects of food supplyandkinshipon social behavior, movements, and population growth of black bearsin northeasternMinnesota. Wildl. Monogr.No. 97. 72pp. . 1989. Fosterfamilies found for desertedcubs. Minn. Volunteer52(305):26-31. , AND A.W. ALLEN. 1987. Habitat suitability index models:blackbear,UpperGreatLakesRegion. U.S. Fish and Wildl. Serv. Biol. Rep. 82(10.144). 54pp. , A.N. MOEN,ANDM.L. SHEDD. 1987. Rectal tempera- turesof 2 free-rangingwhite-taileddeer fawns. J. Wildl. Manage. 51:59-62. , G.A. WILKER,ANDV.D. O'CONNOR. 1989a. BEAR: animal recordingsystem. U.S. Dep. Agric., For. Serv., North CentralFor. Exp. Stn., SR 1, Box 7200, Ely, MN 55731. 1 floppy disk + 15pp. _, _ , AND . 1989b. BEARWATCH: animal dataanalysis system. U.S.Dep. Agric., For. Serv., North CentralFor. Exp. Stn., SR 1, Box 7200, Ely, MN 55731. 2 floppy disks + 37pp. SADE, D.S., J. SCHNEIDER,A. FIGUEROA,J. KAPLAN, K. CUSHING, P. CUSHING, J. DUNAIF, T. MORSE, D. RHODES, AND M. STEWART. 1976. Populationdynamics in relationto social structureon Cayo Santiago. Yearb. of Phys. Anthropol. 20:253-262. G. 1972. The Serengetilion: a study of predatorSCHALLER, prey relations. Univ. of Chicago Press, Chicago. 480pp. S.F. 1974. Mother-infantrelationsin moose. Nat. STRINGHAM, Can. 101:325-369. THILL,R., ANDA. MARTIN,JR. 1989. Deer and cattle diets on heavily grazed pine-bluestemrange. J. Wildl. Manage. 53:540-548. O.C. 1964. Arizona's educateddeer. Ariz. Game WALLMO, and Fish Dep. Wildl. Views 11(6):4-9. , ANDD.J. NEFF.1970. Directobservationof tameddeer to measure their consumptionof naturalforage. Pages 105-110 in Range and wildlife habitatevaluation-aresearch symposium. U.S. Dep.Agric., For. Serv. Misc. Publ. 1147. 220pp. C.R. 1964. Foragepreferencesof captive deer while WATTS, free-rangingin a mixed oak forest. M.S. Thesis. Pa. State Univ., UniversityPark. 65pp.