BEAR REINTRODUCTIONS:LESSONS AND CHALLENGES

INVITEDPAPER

JOSEPHD. CLARK,U.S. GeologicalSurvey, Southern Appalachian Field Laboratory, 274 EllingtonPlant Sciences Building, Universityof Tennessee, Knoxville,TN 37996, USA, email: [email protected] DJUROHUBER, Department of Biology,Veterinary Faculty University of Zagreb,Heinzelova 55, 10000 Zagreb,Croatia, email: [email protected] CHRISTOPHERSERVHEEN, U.S. Fish and WildlifeService, UniversityHall, Room 309, Universityof Montana,Missoula, MT 59812, USA, email: [email protected]

Abstract: Reintroductionis defined as an attemptto establisha species in an areathat was once partof its historicalrange, but from which it has been extirpatedor become extinct. Historically,one of the most successfulprograms was the reintroductionof 254 Americanblack (Ursus americanus) from Minnesotato the InteriorHighlands of Arkansasin the 1960s; that populationhas grown to >2,500 today. More recent efforts have involved fewer but bettermonitored animals and have sometimes employed techniquesto improve site fidelity and survival. In Pennsylvania,for example, pregnantfemale Americanblack bears were successfully translocatedfrom winterdens, the premisebeing thatthe adultfemales would be less likely to returnbecause of the presenceof young cubs. That winter-releasetechnique was comparedto summertrapping and release in Tennessee;winter releases resultedin greatersurvival and reducedpost-release movements. Homing has not been a problemfor small numbersof brownbears (Ursus arctos) reintroducedto the Cabinet-Yaakecosystem in Montanaand Idaho and to the mountainsof Austria and France. Reintroductionsuccess appears to be correlatedwith translocationdistance and is greaterfor subadultsand females. As with any small population,reintroduced populationsare susceptibleto environmentalvariation and stochasticdemographic and genetic processes. Althoughmanagers have focused on these biological barriers,sociopolitical impedimentsto bear reintroductionare more difficult to overcome. Poor public acceptanceand understandingof bears are the main reasons some reintroductionprograms have been derailed. Consequently,the public should be involved in the reintroduction process from the outset; overcoming negative public perceptionsabout bear reintroductionwill be our greatestchallenge.

Ursus 13:335-345 (2002)

Key words: bear,reestablishment, reintroduction, restoration, translocation, Ursus americanus, Ursus arctos

Termsto describe animaltranslocations are often con- patchesfollowing periodiclocal (Levins 1970, fused. The InternationalUnion for the Conservationof Hanski 1996). Bears, however, may not be well adapted Nature (IUCN) ReintroductionSpecialist Groupdefines to exploit such . Although the considerabledis- reintroductionas an attemptto establisha species (or lower persal capabilities of male black and brown bears have taxonomic unit) in an area that was once part of its his- been well documented(Kemp 1976, Youngand Ruff 1982, torical range, but from which it has been extirpatedor Schwartz and Franzmann1992, Blanchardand Knight become extinct (IUCN 1998). Reintroductionis some- 1995, Stratmanet al. 2001), thatbehavior is not typical of times called repatriation(Reinert 1991). When native all age and sex classes, and rateof populationspread cor- conspecifics already are present, reintroductionsare re- relates only weakly with mobility (Caughley 1977:69). ferred to as supplementationsor augmentations(IUCN Females of at least some bearspecies typicallydo not dis- 1998). Reintroductiondiffers conceptuallyfrom the re- perse and, thus, reside within a portionof their mother's lated methods of nuisance translocations,fostering, and home range(Schwartz and Franzmann 1992). Also, bears rehabilitationbecause the primaryobjectives of thosetech- have relatively low reproductiverates (Bunnell and Tait niques usually are not populationreestablishment. Al- 1981), a characteristicwhich limitspopulation growth and though the latter methods have been well documented naturalcolonization ability (Hanski 1991, Hastings1991). (e.g., Boyer and Brown 1988, Linnell et al. 1997, Stiver Consequently,bears should probablybe consideredpoor et al. 1997, Clark 1999), we discuss those methods only colonizers and reintroductionmay be necessary to expe- to the extentthat they relateto reintroductionas the IUCN dite this otherwise slow recolonizationprocess. it. defines Reintroductionis a costly and time-consumingenter- Reintroductionof bears has been the subject of much prise, with only about 11%of all species reintroductions renewed interest. The distributionand numbersof most resulting in viable populations(Beck et al. [1994] from bear species have been dramaticallyreduced and frag- Earhardt [1999]). In general, reintroductionsuccess is mentedbecause of habitatloss, overexploitation,or some enhancedin instances when there are a large numberof combinationof both (Servheen 1990). Yet, in many cases founders,low environmentalvariation, and access to refu- the causes of historical populationdeclines have been rec- gia, and for species with high genetic variability,a high tified; habitatshave recoveredin many areasand laws are rateof populationincrease with low varianceand low in- in place to preventoverexploitation. traspecific competition (Griffith et al. 1989). Unfortu- In the case of populationfragmentation, long-term per- nately, bears exhibit low population growth with high sistence on depends recolonization of vacant variance,are subject to high environmentalvariation (e.g., 336 Ursus 13:2002 annual fluctuations in food production), and have low HISTORICALPROGRAMS genetic variability relative to their population size Few bear reintroductionefforts have occurred,fewer (Manlove et al. 1980, Wathenet al. 1985). Furthermore, have been successful,and fewer still have been adequately bearshave a stronghoming instinctand often travelgreat documented. Perhapsthe earliest programwas a 1933 distances (Beeman and Pelton 1976, Miller and Ballard Californiaeffort, where about 30 Americanblack bears 1982, Rogers 1987) and experience increased mortality were translocatedfrom Yosemite National Park to the following translocation(Rogers 1986, Fies et al. 1987, Angeles National Forest (D. Updike, CaliforniaDepart- Stiver 1991, Comly 1993, Riley et al. 1994, Blanchard ment of Fish and Game, Sacramento,California, USA, and Knight 1995). Thus, reintroductionin general, and personalcommunication, 2002; Table 1). Managersused bearreintroduction specifically, presents many challenges. a hard release, whereby animals were captured,trans- Our goals for this paper are to provide an overview of ported,and released without an acclimationperiod. Griz- bearreintroduction efforts worldwide, summarize the find- zly bearshad been extirpatedin 1922 from the areawhere ings, and offer guidancefor the future. the black bears were reintroduced.Although the Ameri-

Table1. Characteristicsof bearreintroduction programs and eventualoutcomes. Abbreviations:NP = NationalPark, NF = National Forest, Am. = American, NWR= National WildlifeRefuge.

Distance from capture Number Species and Source Release area site (km) re-leased Date reintroductionmethod Outcome Yosemite NP, Calif., Angeles NF, Calif. USA 350 -30 1930s Am. black bears, hard Success USAa release Cook County, Minn., Interior Highlands, 1,570 254 1958-68 Am. black bears, hard Success USAb Ark., USA release Cook County, Minn., Northern La., USA 2,030 161 Mid- Am. black bears, hard Unknown USAc 1960s release Failure Byelorussiad Bialowieza, Poland 300 >11 1938 Brown bears, soft release of captive bears Vienna, Austria (zoo)' Trentino, Italy 2 1959 Brown bears, soft release Failure of captive bears Zurich, Switzerland (zoo) Trentino, Italy 4 1969, Brown bears,hard release Failure and Este Castle, Italy 1974 of captive bears (zoo)' Northeast Penn., USAf Southeast Penn., USA 440 22 1977-84 Am. black bears, soft Success release with winter-den technique Unknown Shenandoah NP, Virginia, SoutheastVa., USA 298 43 1991 Am. black bears, hard USAg release of nuisance bears Croatia, Sloveniah Austria >400 3 1989-93 Brown bears, hard release Success Northern Montana, USA' CabinetMountains, 20-40 4 1990-94 Brown bears, hard release Pending Mont., USA Am. black soft Great Smoky Mountains Big South Fork,Tenn., 160 14 1996-97 bears, Pending den NP, Tenn., USAj USA release with winter technique and summer acclimation pens Sloveniah Central Pyrenees, 1600 3 1996-97 Brown bears, hard release Pending France Northern and southern Am. black bears, soft with winterden La., USAk Central La., USA 50-180 6 1998-01 release Pending technique hard release Slovenia' Alps, Italy -400 7 1999-01 Brown bears, Pending soft White River NWR, Ark., Felsenthal NWR, Ark., 160 10 2000-01 Am. black bears, Pending USAm USA release with winter den technique a 2002 D. Updike, CaliforniaDepartment of Fish and Game, Sacramento,California, USA, personalcommunication, b Smith and Clark 1994 c Taylor 1971 d Buchalczyk 1980 e Osti 1999 f Alt 1995 g Comly 1993 h D. Huber,University of Zagreb,Zagreb, Croatia,, unpublished data i Servheenet al. 1995 J Eastridgeand Clark2001 k D. Anderson,U.S. Fish and Wildlife Service, Tallulah,Louisiana, personal communication, 1999 1 P. Genovesi, InstitutoNazionale per la FaunaSelvatica, Ozzano Emilia, Italy, personalcommunication, 2001 m B. Wear,University of Tennessee, Knoxville, Tennessee, USA, personalcommunication, 2001 BEAR REINTRODUCTION* Clark et al. 337 can black bearpopulation size today is unknown,the ex- ing them in semi-naturalconditions failed and they were pansion of the populationinto unoccupied grizzly range returnedto captivity.In 1969, another2 Carpathianbrown is mostly due to that reintroductioneffort (D. Updike, bearsfrom a zoo in Zurichwere reintroduced.Both were personalcommunication, 2002). habituatedto people. During an attemptedrecapture, the Another early programreleased 254 American black female was seriously injuredand had to be killed. The bears from Minnesota and Manitoba,Canada, in Arkan- male was sent to a zoo in Verona. The last known attempt sas from 1958 through1968 (Rogers 1973). Animalswere to reintroducecaptive bears in the Trentinoarea was in capturedin culverttraps during summer and translocated 1974 using 2 males froma privateenclosure at Este Castle. to 3 locations in the InteriorHighlands of northwestAr- One bearwas apparentlyillegally shot in 1976. The other kansas. Subsequentmovements were extensivewith some was captured,fitted with a radiocollar,and radiotracked animals traveling up to 435 km from their release sites until it died in an avalanchein 1978. (Rogers 1973, Smithet al. 1990). Reintroductionsat all 3 locations were successful, with the resulting population increasingto >2,500 animalsin 20-years(Smith and Clark RECENTEFFORTS 1994). Numerically,this may be the most successful re- introductionof any largecarnivore anywhere in the world. AmericanBlack Bears In the mid-1960s, Louisianaofficials attemptedto aug- From 1977 to 1984, 22 adult female black bears were ment dwindling Americanblack bear populationsin the translocated440 km from northeasternPennsylvania to Tensas River area and the Atchafalaya River Basin by augmenta sparsepopulation in the southwesternportion capturing 161 bears from Minnesota and hard releasing of the state (Alt 1995; G. Alt and M. Ternent,Pennsylva- them 2,030 km away (Taylor1971). Bearsare now present nia Game Commission,Harrisburg, Pennsylvania, USA, at both locations in Louisiana, but it is unclear to what personalcommunication, 2001). In that effort, some fe- extent they are the result of the releases (Pelton 1991). male bears that had just given birthto cubs or were sus- As in Arkansas,post-release movements were extensive pected to be pregnant at the time of relocation were and mortalitywas high (Taylor 1971). radiotrackedto their winter dens in the source area, re- In an early Europeaneffort, brown bears were reintro- moved, and placed in den sites at the release area. At the duced to Bialowieza in northeasternPoland where the release site, these bearswere placed in known den sites or original population was extirpated in the late 1800s in wooden crates lined with straw. The premise of this (Buchalczyk1980). Seven were hand-raisedcubs brought winter-denmethod was that the combinationof hiberna- from neighboringByelorussia (now Belarus)and reintro- tion, parturition,and cub rearing would keep the adult duced in 1938. All 7 bears were soon killed by poachers females in the reintroductionarea, thus increasingsite fi- or were recapturedafter they came into conflict with the delity. Althoughsome nativebears were present,the aug- local human inhabitants. In anotherrelease, a pregnant mentation effort, along with harvest restrictions, was female from PoznanZoo was placed in a cage in the for- thoughtto have greatlyincreased population growth. Prior est where she gave birth to 2 cubs in January 1938 to augmentation,harvests for the area averaged4 bears/ (Buchalczyk 1980). The spacing of the bars of the cage year, whereas the recent hunter-killincreased to an aver- permittedthe cubs to leave and returnat will. Access to age of 111 bears/year(M. Terent, personalcommunica- the cage was blocked in the fall and the cubs spent the tion, 2001). This winter-dentechnique is considered a winter in the wild, returningonly sporadicallythe next soft-release method because release was precededby an spring. Duringthe Second WorldWar and Germanoccu- acclimation period (Griffith et al. 1989). In a Virginia pationof the area,the caged adultfemale andanother cap- program,43 nuisance American black bears were hard tive bear were released. These bears subsequentlykilled released in the southwest portion of the state (Comly 2 or 3 people and injuredanother and were then shot. At 1993). Mortality was high, with annual survival rates the end of the Second WorldWar there were probably4 averaging0.37 for females and 0.12 for males. Homing bears in Bialowieza but none were known to have sur- behavioralso was evident, with 32 of the 43 bears leav- vived after 1947 (Buchalczyk 1980). ing the release areas. Although 11 bearsremained in the Some 20 years later in the Trentinoarea of the Italian areas of release, females did not reproducethe first year Alps, 3 historical reintroductionefforts in- and the populationwas projectedto decline. volved >6 bears (Osti 1999; P. Genovesi, Instituto To attemptto reduce the homing observedin Virginia, Nazionale per la Fauna Selvatica, Ozzano Emilia, Italy, 2 soft-release techniques were comparedfor reintroduc- personalcommunication, 2001). The first was of 2 young ing black bears from Great Smoky MountainsNational Carpathianbrown bears in 1959-60, born in captivity in Parkin Tennessee to the Big South Fork National River a Viennazoo. Attemptsto rehabilitatethe bearsby hold- and RecreationArea in Tennessee and Kentucky,some 338 Ursus 13:2002

160 km to the northwest(Eastridge and Clark 2001). The In France, bears were extirpated from the Central winter-dentechnique was used to reintroduce8 adultfe- Pyrenees by 1990 (Parde 1997). Two females in 1996 males with cubs. The second method involved reintro- and 1 male in 1997 were reintroducedfrom Slovenia, a ducing6 adultand subadult female bears, not accompanied distanceof 1,600 km (D. Huber,unpublished data). Both by cubs, to the release area during summerand holding females were pregnantat time of captureand had litters them in pens for a 2-week acclimationperiod. After re- of 2 and 3 cubs in 1997. A hunterkilled the second fe- lease, total distancemoved from the release sites, net dis- male in fall 1997. One of her cubs and both cubs of the tance moved, mean daily distance moved, and circuity otherfemale appearedto have survived(P. Quenette, Life for winter-releasedbears were significantlyless than for Project,Saint-Gaudens, France, personal communication, summer-releasedbears. Also, survivalof winter-released 2001). bears (0.88) was greater than summer-released bears In the Italian Alps, 2 male and 3 female bears from (0.20). Reproductionamong reintroduced bears was docu- Slovenia were hard released in 1999 and 2000, respec- mented;sires were thoughtto be transientadults or prog- tively, to augmenta wild populationestimated to contain eny of the reintroducedbears. only 2-4 bears. Two additionalfemales were releasedin Also using the winter-dentechnique, a female with 2 2001 (P. Genovesi, personalcommunication, 2001). cubs were reintroducedfrom MadisonParish, Louisiana, Lastly,4 subadultfemale brownbears were releasedto and placed in a denning box on the southernportion of augment the existing small population in the Cabinet Tensas River National Wildlife Refuge, Louisiana, in Mountainsin northwesternMontana between 1990 and March 1998 (D. Anderson,U.S. Fish and Wildlife Ser- 1994 (Servheenet al. 1995). The bears were released in vice, Tallulah,Louisiana, USA, personalcommunication, springand summer,the time of maximumfood availabil- 1999). In late February1999, anotherfemale andher cub ity in the area. One of the bears emerged with a cub the were taken to the BuckhornWildlife ManagementArea following spring but died of unknown causes approxi- in TensasParish and placed in a den. Both females stayed mately50 weeks afterrelease. Heroffspring disappeared. in the reintroductionarea the next year and at least 1 of The other 3 bears eventually droppedtheir radiocollars the 3 cubs survived. In 2001, 4 females with 9 cubs were and have not been recaptured.As of 2001, therewas evi- similarlyreintroduced. Although 1 female abandonedher dence thatat least 1 of the 3 bearssurvived and may have cubs, all adultsremained in the release areas. reproduced,but without recaptureand monitoring,this Finally, the winter-dentechnique was used in 2000 to cannot be verified. translocate6 females with 12 cubs from easternArkansas to Felsenthal National Wildlife Refuge in southernAr- kansas (B. Wear, University of Tennessee, Knoxville, WHATHAVE WE LEARNED? Tennessee, USA, personal communication,2001). One of the 6 females died following reintroduction,the fate of Homingand Survival 1 was unknown, and the other 4 survived and remained One of the obstacles to bear reintroductionhas been near the release area. Of the 15 cubs reintroducedwith homingbehavior. Translocated American black bears have their mothers, 5 were known to have survived >1 year returned hundreds of kilometers to their capture sites afterrelease. In 2001, 4 more adultfemales with 10 cubs (Beeman and Pelton 1976, McArthur1981, Fies et al. were reintroducedto Felsenthal. 1987, Rogers 1988). Factorsthat may influence homing include age, sex, the presence of cubs, food availability, BrownBears translocationdistance, and geographicbarriers. Hard re- From 1989 to 1993, 2 females and 1 male brownbears leases of subadultAmerican black bearshave been more were translocatedto lower Austria and Styria and hard successful than hard releases of adults (Rogers 1988, released (D. Huber, unpublished data). The area was Eastridge2000), and subadultbrown bears have demon- thought to be inhabitedby a male (Rauer 1997). One stratedfewer propensitiesfor homing than adults(Miller female was 3.5 years of age when transplanted;she estab- and Ballard 1982, Brannon 1987, Servheen et al. 1995). lished a home range of >115 km2. She gave birth to 3 Additionally,females may be bettercandidates for rein- cubs in 1991 and had a second litterof 3 cubs in 1993 but troductionbecause theirhome rangesare relativelysmall died in Septemberof that year in an unexplained acci- (Rogers 1973, Wilsonand Gipson 1975, Clark1985). The dent. The other transplantedfemale was 6 years old and winter-releasetechnique first developed in Pennsylvania establisheda home rangeof >4,730 km2(Gerstl and Rauer for Americanblack bears holds much promise to reduce 1999). She also gave birth in 1993 to 2 cubs before the homing (Eastridgeand Clark2001). radiosignalwas lost. The bearpopulation in Austriawas Conversely,homing has not been a problemin brown estimatedto be 20 to 25 in 2001. bearreintroduction programs, even for males. Brownbear BEAR REINTRODUCTION* Clark et al. 339 reintroductionshave primarilytaken place in spring or 1993,Boyce andWaller 2000). Eastridgeand Clark (2001) early summer,when food is relatively abundant.In Aus- demonstratedhow probabilitiesin Tennessee tria, bears used roe deer (Capreolus capreolus) feeding and Kentuckycould be greatly diminished and the time stations extensively (Rauer 1997); thus, artificial food to populationreestablishment greatly reduced with addi- sourcesmay help minimizehoming. Ironically,man-made tional releases of bears. Releases beyond 3-5 years had obstacles such as roads and developmentsthat are barri- less effect on populationgrowth and, with 7 releases, the ers to naturalimmigration of bears also may have pre- change in growth was much less discernible. Similarly, ventedhoming. Finally,translocation distances have been Griffithet al. (1989) demonstratedthat, after a point, re- relativelylarge (400-1,600 km). A numberof otherstud- leasing a large numberof animals does little to improve ies also identifiedan inverserelationship between the dis- reintroductionsuccess. tance American black bears were translocatedand the To reduce costs, it may be temptingfor wildlife agen- probabilityof return(Beeman and Pelton 1976, Singer cies to release bears without subsequent monitoring. and Bratton 1980, McArthur1981, Rogers 1986, Fies et However, growth projectionscannot be made if released al. 1987). bears are not monitoredto determinedispersal, survival, Vehicle-relatedmortalities are often associated with and reproductiverates. As a result, more bears may be homing and can be expected to be higherfor reintroduced stocked than needed or, conversely, the numberreleased bears the year after release (G.L. Alt, 1995, Black bear may be insufficient. Most principlesof small population populationestablishment in SouthwesternPennsylvania, managementapply to reintroducedbear populations and, Pennsylvania Game Commission, Harrisburg,Pennsyl- as such, they are vulnerableto demographicand environ- vania, USA; Comly 1993, Eastridge 2000). Similarly, mental stochasticity. Therefore,population growth pro- Massopustand Anderson(1984) found that survivalwas jections shouldaccount for such potentialvariation (Saltz lower for translocatedAmerican black bears (0.56) than 1996, Eastridgeand Clark2001). for non-translocatedbears (0.72) in Wisconsin, and sur- vival of translocatedbears in Maine (Hugie 1982) and Genetics Tennessee (Stiver 1991) was lowest duringthe first sev- Griffithset al. (1996) warnedthat species reintroduc- eral months after release. Although hard releases of tion without genetic profiling of native stock risks what American black bears such as occurred in the Interior they termed genetic genocide. For example, it remains Highlandsof Arkansasclearly can be successful,the num- unclearwhether bears that exist in northernLouisiana to- ber of animals released must be largerbecause of higher day, now listed as threatenedunder provisions of the U.S. dispersal and increasedmortality. Bears generate much EndangeredSpecies Act of 1973 (16 U.S.C. 1531-1544), interest today, however, and the public may not tolerate are descendents of native stock (U. a. luteolus) or bears excessive numbersof transientor road-killedbears asso- reintroducedfrom Minnesota(U. a. americanus;Pelton ciated with such hardreleases. 1991, Milleret al. 1998, Warrillowet al. 2001). The same subspecies (IUCN 1998) or suitable substitutes(Seddon PopulationDemographics and Soorae 1999) shouldbe used wheneverpossible. Us- Demographicsof the founders can have a markedef- ing moder techniques of molecular genetics, it is now fect on resultantpopulation growth and chance of suc- possible to comparethe genes of extinctpopulations, even cessful reestablishment(Saltz 1996). One reason for the by use of museumsamples, with potentialsource popula- success of the bear reintroductionprogram in Arkansas tions. was a relativelylarge founder population, which included If a small numberof individuals are used for popula- manyadult females and excluded many older males (Smith tion reestablishment,founder effects or genetic drift can and Clark 1994). occur. For example, bearsthat were moved from Minne- Although a large number of founders increase the sota to the InteriorHighlands of Arkansaswere released chances of successful reestablishmentand rate of popula- in 2 regions,the OuachitaMountains and the OzarkMoun- tion growth,bear reintroduction is expensive, particularly tains. About 20% of black bearsin northernmid-western using labor-intensive soft-release methods. Also, the states where the translocatedbears originatedare brown source populationmay be in short supply,particularly if in color (Rounds1987), as are 22%of Ozarkbears (Smith certainage or sex groupsare targetedor if the sourcecon- and Clark 1994). Presumably, few founders of the sists of endangeredstock. Therefore,it is importantto OuachitaMountains population carried this trait,because know when returnsin population growth on the initial few if any bears with brown coat color are known in the investmentof bearsdecline. For thatpurpose, a few rein- OuachitaMountains (Smith and Clark 1994). Otherless- troductionprograms have attemptedto predictpopulation discerniblegenetic differencesbetween the 2 populations growth and the time to populationestablishment (Comly probablyexist as well and could, in theory,affect popula- 340 Ursus 13:2002

tion fitness. a newborn, did not rehabilitatewell and was eventually killed by local humaninhabitants. Habitat Though results are encouragingin this case, samples Littleis gainedby releasinganimals in areaswhere habi- sizes were small and over a relatively shorttime. Many tatis unsuitable;thus bear reintroductions have often been more attemptsto rehabilitatehabituated bears have failed. precededby habitatevaluations (van Manen 1990, Boyce Generally,rehabilitation is costly and laborintensive and and Waller2000, Hogg et al. 2000). In additionto food there may be more efficient ways to obtain source ani- production,other factors such as roadsand humandevel- mals, even for endangeredspecies. Also, rehabilitation opmentshave been assessed. Centralto an evaluationof may not be possible for potentially aggressive bear spe- habitatat the release site is an understandingof the initial cies becausethe consequencesof personalinjury by reha- cause of the decline. For example, flooding regimes bilitated bears could undermine overall conservation coupledwith timberharvesting may affectAmerican black efforts for the species. In Poland, for example, captive bear denning habitat (White et al. 2001) and may have brownbears released in the 1940s killed at least 2 people; contributedto the decline of some bearpopulations in the resentmentand negative attitudes toward bear reintroduc- lower Mississippi Alluvial Valley (A. Edwards,Univer- tion by the public persists today. It seems clear that fur- sity of Tennessee, Knoxville, Tennessee, USA, unpub- therattempts to rehabilitatebrown bears should not occur. lished data). Additionally,in some partsof Asia, problems Becauseof the difficulties,expense, and potential for nega- resulting from habitatloss and poaching have not been tive encounterswith the public, we view rehabilitationas remedied. Thus, it may not be appropriateto release ad- a technique primarilyfor coping with individual bears ditional bears into areas where such obstacles have not ratherthan reintroduction. been overcome. Behavior WHATARE THE CHALLENGES? It is of the utmost importancethat reintroducedbears Althoughthe biological issues are complex, it appears behave as normal wild individuals. Any alterationsof thatthey can be overcome for most bearspecies. Perhaps theirbehavior that make them more dependenton anthro- more dauntingare the social and political aspects of bear pogenic food sources or more exposed to humans will reintroduction.Reading and Kellert(1993) observedthat likely result in reduced survival. Perhapsmore impor- many species reintroductionshave failed because the so- tantly, the release of habituatedbears can result in non- cioeconomic and political aspects were not adequately acceptanceby the local humanpopulation and the rejection addressed. Although the historic of the entirereintroduction project. reintroductionprogram in Arkansaswas designedto gen- Rehabilitationis a complex effort to preparecaptive- erate as little publicity as possible, it was eventually ter- born or hand-raisedbears for release in the wild. Reha- minated because of public opposition (Smith and Clark bilitation and release of orphanedAmerican black bear 1994). cubs has been attemptedwith some success (Stiver et al. Conversely,the U.S. Fish andWildlife Service proposed 1997, Clark 1999). The release of cubs thathave already to reintroducegrizzly bearsinto the BitterrootEcosystem become habituatedto humansis much more difficult. In of east centralIdaho and western Montana starting in 2002 2000, 3 (Helarctos malayanus) cubs, confiscated to facilitaterecovery of the species in the lower 48 United by local authoritiesin East Kalimantan,Indonesia, were States (Boyce et al. 2001). The proposalrecommended trained for release (G. Fredriksson, University of reintroductionof >25 bears over 5 years, with a goal of Amsterdam/Tropenbos-KalimantanProject, Kalimantan eventuallyreaching >280 bears. That effort was marked Timur,Indonesia, personal communication, 2001). From by a 5-year public process which sought ideas from the the time they arrived(3-7 months old) at the holding fa- public,solicited documents for review andcomment, held cility, the bears' handlerswalked them throughthe forest public meetings and hearings,and providedmultiple op- during the day and kept them in cages at night, where portunitiesfor the public and political intereststo get in- they received additionalfood. After 6-9 months,2 of the volved. Additionally,an EnvironmentalImpact Statement 3 bears would spend nights in the forest but would come would have designatedreintroduced bears as an "experi- back to camp in the evening to obtain food. Within 6 mental, nonessential"population managed by a citizen- months the 2 bears were consuming many of the same scientistcommittee (U.S. Fish andWildlife Service 2000). foods eaten by theirconspecifics in the areaand appeared That designation allows more management flexibility to have similar home range sizes. Encountersbetween under the U.S. EndangeredSpecies Act, especially for the releasedand wild sun bearshave been observed. Un- large carnivoresthat need managementrather than strict fortunately,the thirdbear, which had been confiscatedas protection. More than 24,000 comments were received BEAR REINTRODUCTION* Clark et al. 341 and there were 7 formal public hearings. Even after all tially because the livestock guarding tradition among that,the futureof this projectis doubtful;local public and herdsmenhas been lost in the absence of large predators, political pressureforced the U.S. Fish and Wildlife Ser- and now thatthe predatorshave returned,herdsmen have vice to reverse an earlierdecision to go forwardwith the been slow to readopt those husbandrypractices. Simi- reintroduction (Doddridge 2001). The process cost larly,proposed brown bear reintroductions in the western >$700,000 without a single bearbeing moved. U.S. have been opposed mostly by ranchers. Nevertheless, it is critical that public support,particu- Althoughpublic relations and education programs have larly local support,for reintroductionprograms be gar- been successful in developing supportfor some reintro- nered from the outset. As charismaticmegafauna it is duction programs(Kleiman et al. 1990), traditionaledu- easier to gain supportfor bearsbecause of theirhigh pub- cation programs are often woefully inadequate. Such lic appealcompared with other species (Eckholm [1978] programstend to simply provideinformation about a spe- and Westman[1990] from Reading and Kellert [1993]). cies with the assumptionthat such knowledge will result Most people identify with bears and have a positive view in a shift in attitudinalvalues (Readingand Kellert 1993). of them because they are aestheticallyappealing, are in- This is rarely successful if beliefs are strongly held telligent,are of largesize, have the capacityto standerect, (Rokeach [1979] and Chaiken and Stangor [1987] from andhave an omnivorousdiet (Kellert 1994). In a national Reading and Kellert [1993]) because knowledge is only study of Americans,a significantmajority was willing to one of several factorsinfluencing attitude (Kellert 1994). set aside millions of hectares of national forest land for Changinglong-held beliefs, particularlyfor species that grizzly bearconservation, despite the potentialloss of jobs evoke as much emotion as do bears,will not be easy. Ad- and a reduction in timber harvest (Kellert 1985). The dressing such oppositionwill be one of the greatestchal- North Americanattitude toward bears is highly positive, lenges facing the field of conservationbiology (Reading yet wildlife managersmay be far too conservativein ac- and Kellert 1993). Failure to do so will rarely result in knowledgingthis public viewpointtoward bears and their long-termsuccess of reintroductionprograms. populationenhancement and recovery (Kellert 1994). Despite positive overallattitudes toward bears, attitudes towardbear reintroduction are more negative. These nega- RECOMMENDATIONS tive attitudesare partiallyassociated with the perception Because females typicallydo not dispersewidely, bears that reintroductionwill result in land-use restrictions. areextremely susceptible to habitatfragmentation and are Rural,property-owning, and resource-dependentgroups poorrecolonizers of vacatedhabitats. Yet, given adequate tend to be more utilitarianand dominionisticin their val- habitat and a sufficient number of founders, some past ues towardwildlife (Kellert 1994). For example, fear of reintroductionprograms have shown remarkablesuccess. restrictionsand strong libertarianattitudes have played We expect human-assistedreintroduction to be an increas- major roles in the opposition to black-footed ferret ingly valuabletool for augmentingor reestablishingbear (Mustela nigripes) reintroductions(Reading and Kellert populationsin the future. 1993); this probablyalso is truefor bears,particularly the Homing behaviorand associated low survivalrates of endangeredspecies. Such restrictionscould affect hunt- reintroducedbears have been majorimpediments to suc- ing, mineral extraction, grazing, logging, and access to cessful bear population reestablishment,especially for public lands,although historically, they rarelyhave. With Americanblack bears. Factorsthat may improvesurvival bears, as with other carnivores,there seem to be differ- and decreasehoming include a lengthy translocationdis- ences between urbanand ruralattitudes toward reintro- tance,natural or man-madebarriers to beardispersal from duction. For example, a 1995 Tennessee survey found the release area,abundant food at the release site, and the that61% of local residentswere in favor of an American use of subadults. Translocationof hibernatingfemale black bear reintroductioncompared to 81% for non-lo- Americanblack bears with cubs has been successful and cals (Peine et al. 1995). This is noteworthybecause rural may hold promise for otherbear species. attitudesprobably best reflect those most likely to be af- Centralto the evaluation of habitatat the reintroduc- fected by the reintroductionprogram. tion site is the identificationof the initial cause of popula- Othernegative attitudesare linked to the potentialdan- tion decline; this is often overlooked in reintroduction ger to humansand the destructionof livestock and crops. programs. Total potential range should be estimated so In Arkansas,landowner attitudes toward bears were greatly long-term population goals can be set; population-area influenced by previous experiences with bear damages relationshipsfor bears may be similar to those of other (Clarket al. 1991). In Europe,most oppositionto brown large carnivores (Schoenwald-Cox et al. 1988). Multi- bear reintroductionwas from fear of livestock losses (P. variate assessments of past reintroductionshave identi- Quenette, personal communication,2001). This is par- fied factors contributingto the success or failure of red 342 Ursus 13:2002 wolf (Canis rufus) releases (van Manen et al. 2000) and Act of 1973 (16 U.S. Code 1531-1544), for example,rep- may be worthwhilefor bears. Additionally,a recentanaly- resents a potentiallypowerful use of authority.Next, re- sis suggests that American black bear absence or pres- introduction programs can purchase support through ence in the southernAppalachian mountains in the eastern financial incentives for conservation. Gray wolf (Canis U.S. can be explained by simple landscapemetrics such lupus) reintroductionsin the western U.S., for example, as patchsize anddistribution (Murrow 2001). The analy- are accompaniedby an indemnityfund for livestock dep- sis identifiedcertain habitat fragments that could serve as redationlosses. Finally,public relations programs should lynchpinsfor surroundingpatches if restockedwith bears attemptto persuadepeople to supportthe reintroduction (Murrow2001). Such metapopulationapproaches could program,or at least, not actively oppose it. Enlistingthe greatlyincrease the efficiency of bearreintroduction pro- help of people with similar culturaland socioeconomic grams. backgroundscan be effective. A committee of commu- Although the population consequences of inbreeding nity leaders was established in Montana to work with in bearshave not been documentedas well as with some managersto answerquestions and concernsabout a griz- other carnivores(O'Brien et al. 1985, 1990), genetic is- zly bearaugmentation program in the CabinetMountains sues should not be overlooked. Soule et al. (1986) rec- (Servheen et al. 1995). As demonstratedby the Bitter- ommend a 90% gene retention level from the source root reintroductioneffort, that task will not be trivial. populationto avoid demographicconsequences of a loss Nevertheless,we suggest that conservationistsshould be of genetic diversity,but this may be difficult to achieve more active in developing programsto purchase, pres- for some endangered species (Earnhardt1999). Bear sure, and persuadepublic supportfor bearreintroduction populationson peninsularfringes of theirrange have been efforts. shownto have significantlyreduced genetic variation com- paredto more centralizedpopulations (Waits et al. 1999), an importantconsideration when choosing a source. Tests ACKNOWLEDGMENTS to determinefounder genome equivalents,mean kinship, We thankall the individualsthat we called upon to an- or othermeasures of the relatednessof the foundingindi- swer our questions about bear reintroduction.Those in- viduals to the source populationshould be routinelyper- clude G. Alt, D. Anderson,A. Casellanos, S. Dobey, R. formed (Earhardt 1999). Eastridge,G. Fredriksson,P. Genovesi, W. Kasworm,L. Finally, too little emphasis has been placed on the Morgan-Smith,J. Murrow,M. Pelton,B. Peyton,G. Rauer, sociopolitical aspects of bear reintroduction.The public J. Swenson, M. Terent, F. van Manen, B. Wear,and P. has an overall positive perceptionof bears,but conserva- Yves Quenette. S. Dobey, W. Kasworm, and B. Wear tionists often fail to take advantageof this. However,the helped prepareslides. We thankR. Eastridgeand F. van negative aspects of bears (e.g., personalinjury, property Manenfor reviews of earliermanuscripts. Associate Edi- damage)are often inflatedor given improperperspective. tor D. Shidelerand reviewers J. Keay,R. Reading,and H. Opinionsurveys and conservationeducation through the Reynoldsprovided excellent editorial comments. Finally, mass media and at local levels shouldbe a cornerstoneof we especially thank S. Miller and the IBA Conference any reintroductionprogram (Kleiman 1989). Careshould Committeefor asking us to prepareand presentthis pa- be takenso thatsuch opinion surveys are an accurategauge per. of public sentiment,however, because a few vocal indi- viduals can dominate public meetings and have undue influence on the process. A procedureincorporating de- LITERATURECITED cision analysis,expert opinion, and tradeoffanalysis was ALT, G.L. 1995. Black bear population establishment in southwestern Final for Job 06233, used in the grizzly bear reintroductionprogram in the Pennsylvania. Report Game CabinetMountains in Montanato reconcile the biologi- Pennsylvania Commission,Harrisburg, Pennsylvania, USA. cal needsof the reintroducedbears with the socioeconomic BEEMAN,L.E., ANDM.R. PELTON.1976. Homing of black bears of the local humanresidents andServheen needs (Maguire in theGreat Smoky Mountains National Park. International 1992). Thatsophisticated approach could serveas a model Conferenceon BearResearch and Management 3:87-95. for otherprograms. BLANCHARD,B.M., AND R.R. KNIGHT. 1995. Biological Reading and Kellert (1993) describe 3 basic methods consequencesof relocatinggrizzly bears in theYellowstone for reducingopposition and developing supportfor rein- ecosystem.Journal of WildlifeManagement 59:560-565. troductionprograms: pressure, purchase, and persuasion. BOYCE,M.S., E.M. KIRSCH,AND C. SERVHEEN.2001. Bet-hedging forconservation. Journal of Bioscience27:385- They suggest thatpower and authorityin the form of law applications enforcementto controlaccess anduse of resourcescan be 392. ,AND J. WALLER.2000. The applicationof resource used to The U.S. Species apply pressure. Endangered selectionfunction analysis to estimatethe number of grizzly BEARREINTRODUCTION * Clark et al. 343

bears that could be supportedby habitats in the Bitterroot reintroductions. 10:923-930. ecosystem. Pages 6/231-6/241 in Grizzly bear recovery in HANSKI,I. 1991. Single-species metapopulationdynamics: the Bitterroot ecosystem. Final Environmental Impact concepts, models, and observations. Biological Journalof Statement.U.S. Departmentof the InteriorFish andWildlife the Linnean Society 42:17-38. Service, Missoula, Montana,USA. . 1996. Metapopulationecology. Pages 13-43 in O.E. BOYER,D.A., ANDR.D. BROWN.1988. A surveyof translocations Rhodes, R.K. Chesser,and M.H. Smith,editors. Population of mammals in the United States 1985. Pages 1-11 in L. dynamics in ecological space and time. The University of Nielsen and R.D. Brown, editors. Translocationof large Chicago Press, Chicago, Illinois, USA. mammals. Wisconsin Humane Society, Inc., Milwaukee, HASTINGS,A. 1991. Structured models of metapopulation Wisconsin, USA. dynamics. Pages 57-71 in M. Gilpin and I. Hanski,editors. BRANNON, R.D. 1987. Nuisance grizzly bear, Ursus arctos, Metapopulation dynamics. Academic Press, London, translocationsin the GreaterYellowstone Area. Canadian England. Field-Naturalist101:569-575. HOGG,J., N. WEAVER,J. CRAIGHEAD,M. POKORNY,B. STEELE,R. BUCHALCZYK,T. 1980. The brownbear in Poland. International REDMOND, AND F. FISHER. 2000. Abundance and spatial Conferenceon Bear Researchand Management4:229-232. distributionof grizzlybear food plantsin the Salmon-Selway BUNNELL,F.L., ANDD.E.N. TAIT. 1981. Populationdynamics ecosystem: a preliminaryanalysis andreport. Pages 6/247- of bears-implications. Pages 75-98 in C.W. Fowler and 6/270 in Grizzly bear recovery in the Bitterrootecosystem. T.D. Smith,editors. Dynamicsof largemammal populations. Final EnvironmentalImpact Statement. U.S. Departmentof John Wiley and Sons, New York,New York,USA. the InteriorFish and Wildlife Service, Missoula, Montana, CAUGHLEY,G. 1977. Analysis of vertebratepopulations. John USA. Wiley and Sons, New York,New York,USA. HUGIE, R.D. 1982. Black bear ecology and managementin the CLARK,J.D. 1985. Notes on the death of an introducedbear in northernconifer-deciduous forests of Maine. Dissertation, Arkansas. Proceedingsof the ArkansasAcademy of Science University of Montana,Missoula, Montana,USA. 34:121. INTERNATIONALUNION FORTHE CONSERVATIONOF NATURE. 1998. ,D.L. CLAPP,K.G. SMITH,AND T.B. WIGLEY.1991. Black Guidelinesfor re-introductions.Prepared by the International bear damage and landowner attitudes toward bears in Union for the Conservation of Nature Re-introduction Arkansas. Proceedings of the SoutheasternAssociation of SpecialistGroup. Gland,Switzerland and Cambridge, United Fish and Wildlife Agencies 45:208-217. Kingdom. CLARK,J.E. 1999. Captureand on-site release of nuisanceblack KELLERT,S.R. 1985. Social andperceptual factors in endangered bears and survival of orphanedblack bears released in the species management. Journal of Wildlife Management Great Smoky Mountains. Thesis, University of Tennessee, 49:528-536. Knoxville, Tennessee, USA. . 1994. Public attitudes toward bears and their COMLY,L.M. 1993. Survival,reproduction, and movementsof conservation. InternationalConference on Bear Research translocated nuisance black bears in Virginia. Thesis, and Management9(1):43-50. Virginia Polytechnic Institute and State University, KEMP, G.A. 1976. The dynamics and regulationof black bear, Blacksburg,Virginia, USA. Ursus americanus, populations in northern Alberta. DODDRIDGE,J. 2001. Reevaluationof the recordof decision for InternationalConference on BearResearch and Management the Final EnvironmentalImpact Statement and selection of 3:191-197. alternative for grizzly bear recovery in the Bitterroot KLEIMAN,D.G. 1989. Reintroductionof captive mammalsfor Ecosystem. FederalRegister 66(121):33623-33624. conservation. Bioscience 39:152-161. EARNHARDT,J.M. 1999. Reintroductionprogrammes: genetic , B.B. BECK.A.J. BAKER,J.D. BALLOU,L.A. DIETZ,AND trade-offsfor populations.Animal Conservation 2:279-286. J.M. DIETZ.1990. The conservationprogram for the golden EASTRIDGE,R. 2000. Experimentalrepatriation of black bears lion tamarin,Leontopithecus rosailia. EndangeredSpecies to the Big South Fork area of Kentucky and Tennessee. Update 8(1):82-85. Thesis, Universityof Tennessee,Knoxville, Tennessee, USA. LEVINS,R. 1970. Extinction. Pages 77-107 in M. Gerstenhaber, , ANDJ.D. CLARK. 2001. Evaluation of 2 soft-release editor. Some mathematical questions. American techniques to reintroduce black bears. Wildlife Society MathematicalSociety, Providence,Rhode Island, USA. Bulletin 29:1163-1174. LINNELL,J.C., R. AANES, J.E. SWENSON,J. ODDEN,AND M.E. SMITH. FES, M.L., D.D. MARTIN,AND G.T. BLANK,JR. 1987. Movements 1997. Translocationof carnivoresas a methodof managing and rates of returnof translocatedblack bears in Virginia. problem animals. and Conservation6:1245- InternationalConference on BearResearch and Management 1257. 7:369-372. MAGUIRE, L., AND C. SERVHEEN. 1992. Integrating biological GERSTL,N.J., ANDG. RAUER. 1999. 10 Jahre Braunbar (Ein and social concerns in endangered species management: Projektbilanz: 1989-1999). World Wildlife Fund, augmentation of grizzly bear populations. Conservation Osterreich,Vienna, Austria. (In German.) Biology 6:426-434. GRIFFITH,B., J.M. SCOTT,J.W. CARPENTER,AND C. REED. 1989. MANLOVE,M.N., R. BACCUS,M.R. PELTON,M.H. SMITH, AND D. Translocation as a species conservation tool: status and GRABER. 1980. Biochemical variation in the black bear. strategy. Science 245:477-480. InternationalConference on BearResearch and Management GRIFFITHS,H.I., A. DAVISON,AND J. BIRKS. 1996. Species 4:37-41. 344 Ursus 13:2002

MASSOPUST,J.L., ANDR.K. ANDERSON.1984. Homing tendencies on population genetics. University of Chicago Press, of translocatednuisance black bears in northernWisconsin. Chicago, Illinois, USA. EasternWorkshop on Black BearResearch and Management .1988. Homingtendencies of largemammals: a review. 7:66-73. Pages 76-92 in L. Nielsen and R.D. Brown, editors. MCARTHUR,K.L. 1981. Factors contributingto effectiveness Translocation of large mammals. Wisconsin Humane of black bear transplants.Journal of Wildlife Management Society, Inc., Milwaukee, Wisconsin, USA. 45:102-110. ROGERS,M.J. 1973. Movements and reproductivesuccess of MILLER, D.A., E.M. HALLERMAN,AND M.R. VAUGHAN. 1998. black bears introducedinto Arkansas. Proceedings of the Genetic variationin black bear populationsfrom Louisiana SoutheasternAssociation of Fish and Wildlife Agencies and Arkansas: examining the potential influence of 27:307-308. reintroductionsfrom Minnesota. Ursus 10:335-341. ROUNDS,R.C. 1987. Distributionand analysis of colourmorphs MILLER,S.D., ANDW.B. BALLARD. 1982. Homing of transplanted of the black bear (Ursus americanus). Journal of Alaskan brown bears. Journal of Wildlife Management Biogeography 14:521-538. 46:869-876. SALTZ,D. 1996. Minimizing extinction probability due to MURROW,J.L. 2001. A black bear metapopulationanalysis for demographicstochasticity in a reintroducedherd of Persian the southernAppalachians. Thesis, Universityof Tennessee, fallow deer Dama dama mesopotamica. Biological Knoxville, Tennessee, USA. Conservation75:27-33. O'BRIEN, S.J., M.E. ROELKE,L. MARKER,A. NEWMAN,C.A. SCHWARTZ,C.C., ANDA.W. FRANZMANN.1992. Dispersal and WINKLER,D. MELTZER,L. COLLY, J.F. EVERMANN,M. BUSH, survival of subadultblack bears from the Kenai Peninsula, ANDD.E. WILDT. 1985. Genetic basis for species Alaska. Journalof Wildlife Management56:426-431. vulnerabilityin the cheetah. Science 227:1428-1434. SCHOENWALD-COX,C.M., R.J. BAKER,AND J.W. BAYLESS. 1988. , N. YUHKI,K.W. RICHARDS,W.E. JOHNSON,W.L. Applying the population/areamodel to the planningof large FRANKLIN,A.E. ANDERSON,O.L. BASS,JR., R.C. BELDEN,AND mammaltranslocations. Pages 52-63 in L. Nielsen andR.D. J.S. MARTENSON.1990. Genetic introgressionwithin the Brown,editors. Translocationof largemammals. Wisconsin Floridapanther Felis concolor coryi. National Geographic HumaneSociety, Inc., Milwaukee,Wisconsin, USA. Research6:485-494. SEDDON,P.J., AND P.S. SOORAE.1999. Guidelinesfor subspecific OSTI,F. 1999. I'Orso brunonel Trentino. Arca press, Trento, substitutionsin wildlife restorationprojects. Conservation Italy. (In Italian.) Biology 13:177-184. PARDE,J. 1997. The brown bear in the central Pyrenees: its SERVHEEN,C. 1990. The statusand conservationof the bearsof decline and recent situation. InternationalConference on the world. InternationalConference on Bear Researchand Bear Researchand Management9(2):45-52. ManagementMonograph Series 2. PEINE,J., M. FLY,D. STYNES,AND B. STEPHENS.1995. Visitor , W.F. KASWORM,AND T.J. THEIR. 1995. Transplanting demographics,behavior, opinions and economic impact,Big grizzly bears Ursus arctos horribilis as a managementtool South Fork National River and RecreationArea. Reportto - results from the Cabinet Mountains, Montana, USA. National Park Service, Southern Appalachian Field Biological Conservation71:261-268. Laboratory,U.S. Geological Survey, Knoxville, Tennessee, SINGER, F.J., AND S.P. BRATTON. 1980. Black bear/human USA. conflicts in the Great Smoky Mountains National Park. PELTON,M.R. 1991. Black bearsin the Southeast: to list or not InternationalConference on BearResearch and Management to list? Eastern Workshop on Black Bear Research and 4:137-149. Management10:155-161. SMITH,K.G., ANDJ.D. CLARK.1994. Black bears in Arkansas: RAUER,G. 1997. First experienceswith the release of 2 female characteristics of a successful translocation. Journal of brown bears in the Alps of EasternAustria. International Mammalogy75:309-320. Conferenceon Bear Researchand Management 9(2):91-95. ANDP.S. GIPSON.1990. Historyof black bears READING,R.P., AND S.R. KELLERT. 1993. Attitudes toward a in Arkansas: over-exploitation, near elimination, and proposed reintroductionof black-footed ferrets (Mustela successful reintroduction.Eastern Workshop on Black Bear nigripes). ConservationBiology 7:569-580. Researchand Management10:5-14. REINERT,H.K. 1991. Translocationsas a conservationstrategy SOULS,M.E., M. GILPIN,W. CONWAY,AND T. FOOSE.1986. The for amphibiansand reptiles: some comments,concerns, and millenniumark: how long a voyage, how many staterooms, observations. Herpetologica47:357-363. how many passengers? Zoo Biology 5:101-113. RILEY,S.J., K. AUNE,R.D. MACE,AND M.J. MADEL.1994. STIVER,W.H. 1991. Populationdynamics and movements of Translocation of nuisance grizzly bears in northwestern problem black bears in Great Smoky MountainsNational Montana. InternationalConference on Bear Research and Park. Thesis, Universityof Tennessee,Knoxville, Tennessee, Management9(1):567-573. USA. ROGERS, L.L. 1986. Effects of translocation distance on , M.R. PELTON,AND C.D. SCOTT.1997. Use of pen-reared frequency of returnby adult black bears. Wildlife Society black bears for augmentation or reintroductions. Bulletin 14:76-80. InternationalConference on BearResearch and Management 1987. Factors influencing dispersal in black bear. 9(2):145-150. Pages 75-84 in B.D. Chepko-Sadeand Z.T. Haplin,editors. STRATMAN,M.R., C.D. ALDEN,M.R. PELTON,AND M.E. SUNQUIST. Mammaliandispersal patterns, the effects of social structure 2001. Long distance movement of a Floridablack bear in BEARREINTRODUCTION * Clark et al. 345

the southeasterncoastal plain. Ursus 12:55-58. WARRILLOW,J., M. CULVER, E. HALLERMAN,AND M. VAUGHAN. TAYLOR,D.F. 1971. A radio-telemetrystudy of the black bear 2001. Subspecific affinity of black bearsin the White River (Euarctosamericanus) with notes on its history and present National Wildlife Refuge. The Journal of Heredity statusin Louisiana.Thesis, Louisiana State University, Baton 92(3):226-233. Rouge, Louisiana,USA. WATHEN, W.G., G.F. MCCRACKEN, AND M.R. PELTON. 1985. U.S. FISHAND WILDLIFE SERVICE. 2000. Grizzly bear recovery Genetic variation in black bears from the Great Smoky in the Bitterroot ecosystem. Final EnvironmentalImpact MountainsNational Park. Journalof Mammalogy66:564- Statement.U.S. Departmentof the InteriorFish andWildlife 567. Service, Missoula, Montana,USA. WHITE, T.H., JR., J.L. BOWMAN, H.A. JACOBSON,B.D. LEOPOLD, VANMANEN, FT. 1990. A feasibility study for the potential ANDW.P. SMITH. 2001. Forestmanagement and female black reintroductionof black bears into the Big South Fork Area bear denning. Journalof Wildlife Management65:34-40. of Kentuckyand Tennessee. TennesseeWildlife Resources WILSON,S., ANDP. GIPSON.1975. Status of black bear in the Agency, TechnicalReport 91-3, Nashville, Tennessee,USA. western Ozarks. ArkansasGame and Fish Magazine 6:10- , B.A. CRAWFORD,ANDJ.D. CLARK.2000. Predictingred 11. wolf release success in the southeastern United States. YOUNG,B.F., AND R.L. RUFF.1982. Populationdynamics and Journalof Wildlife Management64:895-902. movements of black bears in east central Alberta. Journal WAITS,L., D. PAETKAU,AND C. STROBECK.1999. Genetics of of Wildlife Management46:845-860. the bears of the world. Pages 25-32 in C. Servheen, S. Herrero,and B. Peyton, editors. Bears: Status, Survey and Received: 13 June 2001. Conservation Action Plan. International Union for the Accepted: 26 February 2002. Conservationof Nature,Gland, Switzerland. Associate Editor: Shideler