POLARBEARS OF THE ARCHIPELAGO OF THE RUSSIAN

STANISLAVE. BELIKOV,Ministry of EnvironmentalProtection and NaturalResources, All-RussianResearch Institutefor Nature Conservation,Moscow, , email:[email protected] GERALDW. GARNER,U.S. NationalBiological Service, Alaska Science Center,1101 East TudorRoad, Anchorage,AK 99503, USA, email:[email protected] 0YSTEINWIIG,1 Norwegian Polar Institute,Oslo, Norway ANDREIN. BOLTUNOV,Ministry of EnvironmentalProtection and NaturalResources, All-RussianResearch Institutefor Nature Conservation,Moscow, Russia, email:[email protected] YURIA. GORBUNOV,Arctic and AntarcticResearch Institute,St. Petersburg,Russia

Abstract: The SevernayaZemlya Archipelagowas visited by Russianexplorers during 1930-32, and observationsof polarbears (Ursus maritimus) were first recordedincidental to expedition membershunting polar bears for food and fur. A joint Russian, Norwegian, and U.S. researcheffort on ecology in the SevernayaZemlya Archipelagowas initiated in 1991. A total of 11 bears were captured,including 5 adult females which were fitted with satellite collars. Two satellite collars failed soon after deployment(<25 locations), but 3 units functionedfor an extended period of time (>50 locations). A majorityof the movementswere confined to the easternKara Sea, but 1 bear moved into the westernLaptev Sea before returningto the easternedge of the KaraSea near Cape Cheluskin. One bearhad a very restrictedmovement pattern closely associatedwith the 2 northwesternislands in the SevernayaZemlya group (Komsomolets and Pioner ). The combined locations for the 5 collared bears depict an overall range that is centeredaround the SevernayaZemlya Archipelago. Minimumestimates of distances traveledfor 3 bears with >50 locations averaged4,183 km. The daily rates of movement for the study period varied between bears and rangedbetween 6.4 and 14.7 km/day. The rate of movement duringminimum sea ice cover was the highest (12.7 km/day) for 4 defined periods, while rates of movement for the other 3 periods were very similar (8.5-9.3 km/day). Polar bears in this region of Russia do not move long distances to maintaincontact with the sea ice, in contrastto polar bears in the Chukchi Sea in eastern Russia which move over large areas to maintaincontact with the sea ice throughoutthe year.

Ursus 10:33-40

Key words: , , movements, polar bear, sea ice, Severaya Zemlya, Ursus maritimus.

The SevernayaZemlya Archipelagowas discoveredin were found only in the southern part of by a Russian hydrographicexpedition, SevernayaZemlya. Total 105 bearswere hunted but only the approximateshape of the eastern shore of duringthe expedition, 66 were adult males, 24 the archipelagowas mapped. The first expeditionto map single females, 6 females with cubs, and 9 cubs. the Severaya Zemlya Islands occurredduring 1930-32. No bearswere huntedin January,1 in February, The archipelagowas uninhabitedprior to this effort. The 15 in March, 11 in April, 14 in May, 11 in June, 4-man expeditionwas based on DomashniIsland, a small 9 in July, 15 in August, 14 in September,9 in in the western part of the archipelago. Mapping October,4 in November,and 2 in December. A was accomplishedduring 2 trips by researchersthat cir- total of 78 of all bears killed were in a vicinity cumnavigatedthe islandsusing dog teams and sleds. The of the expedition base. High numberof bears diet of expedition team members and dogs included the killed in August and Septemberwas not condi- meat of ringed seals (Phoca hispida), white whales tionedby especiallyhigh densityof bearsin these (Delphinapterus leucas), and polar bears (Ursus months. The cause was that this was time for maritimus). Urvantsev(1935:122) summarizedinforma- gettingfood supplybefore wintering. Bears were tion collected on polar bears duringthe expedition: met moreoften in May when they migratednorth from the TaimyrPeninsula shore. In this month In winter, approximatelyfrom December to paths of bear's tracks of north direction were March, when sea surface is frozen, polar bears observed near western of Komsomolets move south of However capes Severnaya Zemlya. and Pioner islands. some bearsstay in the region. In the second year of the expedition,ice conditionswere not so hard Additionalinformation on polarbears of the region has as a year before, there was a lot of open water been obtainedby polar stationpersonnel on the mainland and bears were met more often. Maternitydens shore and within the islands. Almost all informationon

1Presentaddress: Zoological Museum, University of Oslo, Oslo, Norway, email:[email protected] 34 Ursus 10:1998 polarbears was obtainedincidental to bearhunting. Dur- A joint researchproject between Russian,Norwegian, ing the initialdevelopment of this region,polar bears were and American scientists on polar bear ecology in the consideredharmful animals that damaged human prop- Severaya Zemlya Islands of the east-central Russian erty (Koshkin 1937). Normally, bears were killed when Arctic was initiatedduring May 1991. The objective of they approachedpolar stationsand settlements,and there the researchwas to determinemovement patterns of po- was little effort to hunt bears on the sea ice. Polar bears lar bears near the SevernayaZemlya Archipelagoand to visited settlementsin the region during all seasons, but define the areas occupied by this population. observationswere duringwinter. The dearthof - Fundingfor this study was providedby the U.S. Fish servationsmay be attributableto both females and males and Wildlife Service, the All-Union Research Institute using over-winterdens, as reportedby Popov (1939) and for Nature Conservationand Reserves, and the Norwe- Rutilevski(1939) in the TaimyrPeninsula area. Accord- gian Polar Institute. Appreciationis extended to P.E. ing to Rutilevski(1939) all bearsused dens, but the dura- Fjeld, Norwegian Polar Institute, Oslo, Norway; V. tion of denningappeared to dependon age, sex, and status Baranov, Arctic and Antarctic Research Institute, St. of the animals. Young bears weaned the previous spring Petersberg;W. Bartmann,Dortmann Zoo, Germany;L. and adultmales stay in dens about50 days. Females with Bogdan, All-Russia Research for Nature Conservation, yearling cubs stay in dens about 100 days, while barren Moscow; and A.L. Malev, Kazan Zoo, for assistance in females occupy dens for 115-125 days. Parturientfe- the field, and to the Aeroflot ME-8 helicopter crew for males remainin maternitydens for 160-170 days. professional and efficient helicopter service. S. Arthur The distributionand migration of polar bears in the and T. Olson assisted in computeranalysis and prepara- Severaya Zemlya region appeardependent on ice con- tion of graphics. ditions and abundanceof prey species (primarilyringed seals and beardedseals, Erignathusbarbatus). It is be- lieved that the majorityof polar bears move north fol- STUDYAREA lowing the receding ice during the summermonths and The study areaconsisted of the SevernayaZemlya Ar- returnsouth duringearly winter with the advancingice. chipelago and the seasonal and permanentpolar pack ice In addition, bears searching for seals move locally in the adjacentLaptev and KaraSeas (Fig. 1). The archi- (Kirpichnikov1938, Popov 1939, Rutilevski1939). Some pelago lies approximately50 km north of the Taimyr bears remain in restricted areas, often associated with Peninsulaon the Russianmainland. The climate is harsh shorelineor islands. Rutilevski (1939) reportedthat po- with wintertemperatures frequently below -30 C, accom- lar bears were observednear Cape Cheluskinthroughout paniedby prevailingnortherly winds often exceeding 30 the year. Ice conditionsof the Vilkitski Straitmay deter- m/sec. mine the presenceof bearsnear Cape Cheluskinby influ- Ice conditionsin the Laptev and KaraSeas are severe, encing the abundanceof prey. In May and June, bears with ice formingduring early September(Fig. 1A) in the moved west along the shorelinewhere seals are abundant presenceof the remainingremnant ice. In VilkitskiStrait, in the fast ice. During 1933 approximately350-400 bears the average ice accumulationin October-Januaryis 7- passed near Cape Cheluskin, and 53 bears were killed 11 cm/10 days; in February-April,5-7 cm/10 days; and there between October 1932 to 10 September1933. Po- in May, 1-3 cm/10 days. Ice thickness near Cape lar bear hunting in the archipelagocontinued incidental Cheluskin by 1 Januaryis approximately88 cm, by 1 to other activities through 1956, when hunting of polar March, 140 cm, and by 1 June, 172 cm. bears was prohibitedthroughout the entire Russian Arc- Shorefast ice is usually completely formed by Sep.- tic due to concerns of populationdepletion. Oct. and persists throughJune. In the northeasternKara The first organizedRussian researchon polar bears in Sea, fast ice is extensive because of numerousislands. the Severnaya Zemlya Archipelago was conducted in Along the easterncoasts of the archipelago,the shorefast 1982 when the 4 largest islands were surveyed during a ice zone is comparativelynarrow (Fig. 1B). Multi-year 4-day period. Seven dens were found, and females with ice as well as icebergs are often includedin fast ice along cubs occupied 3 (Belikov and Randla 1987). The survey the shores of the archipelago. In some years multiyear was not replicatedand occurredduring a portion of the fast ice occurs in bays and straitsof the archipelago. Ice den emergence period; therefore, extrapolationto total cover consists mainly of ice formedduring fall. The pro- denning population during the 1981-82 winter is not portion of 2-year and multiyearice in fields is about 3- possible. 5%. Hummockedice is more common near shore. POLARBEARS OF SEVERNAYAZEMLYA * Belikov et al. 35

I,' 4 ,.sland ~ 0 , vSland j ea 80

' / /7i /

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90 100 90 100

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Shmidtl / \ / Shmidtia n /

Arkticheskiy / Ca ichekiy

/ \ S ' Komsonolets / / /, Komso olets /

I/ *[- /; Krasnoi Strait / rasnoi Strait ~~~/ //~ | ^^^~ 1/ Laptev 5Sn / ,

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KaraSet K / /

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Maximumice extent / / Vilkitski Strait / / / ilkitski Strait /

/ heluskin heluskin

/ e, I^ Tamiyr Penins/a // Tamiyr Penin;ula

Fig. 1. Average seasonal sea ice conditions in the Severnaya Zemlya Archipelago, Russia. A. Periods of stable ice formation. B. Limits of fast ice and polynyas at the end of May. C. Position of southern ice edge and southern edge of consolidated ice during late August. D. Position of southern edge of remnant and solid remaining ice in late September. 36 Ursus 10:1998

Winds perpendicularto the shorelinecause the appear- ice 1 June-15 July; minimum sea ice 16 July-15 Sep- ance of polynyas, which may be ice free or covered by tember;and advancing sea ice 16 September-15 Octo- young ice. In February-June,polynyas arepresent about ber. Cumulative distance moved by each bear was 50% of the time to the west of the archipelago. East of calculatedby summingthe greatcircle distancesbetween the SevernayaZemlya Archipelago,polynya occurrence consecutivelocations (1 location/5-or 6-day dutycycle). decreases from 90% in Februaryto 20% in June. Daily rates of movement (km/24 hours) were calcu- In Vilkitski Strait, the sea ice usually begins melting lated from the movement data by dividing the distance by mid-June,and near Cape Arcticheski, during late June. between consecutive locations for a bear by the time in- By the end of June about 10-20 cm of the sea ice has terval in hours. Rates of movement were tested for dif- melted. By mid-Augustalmost all of the shorefastice is ferences between individualsand seasons using analysis broken. However, the Severaya Zemlya region usually of varianceprocedures (ANOVA; Steel andTorrie 1980). does not become completelyice free. Remnantice fields, When differences were detected, the Student-Newman- which include solid ice, remainnear the easternand west- Kuel (S-N-K) multiple comparison test procedurewas ern sides of the archipelagothrough the end of summer used to define which differences were significant (Steel (Figs. 1C and ID). The active process of ice breakup and Torrie 1980). takes place during summer. In August, 10-30% of the Concurrentice informationwas synthesized from ice areamay remainice covered in the region. This ice cover data collected during aerial ice reconnaissancesurveys usually consists of fragmentsof ice fields andbroken ice. conducted by the Russian HydrometerologicalService, and remotely sensed ice data that was collected by Rus- sian satellites. Informationwas summarizedusing the METHODS standardizedRussian procedures for interpretationof sea ice data (Volkov 1981) and plottedon maps for compari- Captureand Marking son to polar bear movements. Bears were captured10-15 May 1991 using helicopter immobilizationprocedures (Lentfer 1968, Larsen 1971, Schweinsburget al. 1982) and remote projectile injec- RESULTS tion of Telazol? (Haigh et al. 1985, Stirling et al. 1989). Tranquilizedbears were measured,weighed using a spring Captureand Marking scale, blood-sampleswere drawn,and a vestigial premo- Four adult female polar bears and 6 accompanying lar tooth was extractedfor aging by counts of cementum COYs (mean litter size = 1.5, SE = 0.29) were captured annuli (Hensel and Sorensen 1980). Cubs-of-the-year (Table 1). In addition, 1 non-lactatingfemale (N7988) (COYs) were measured,weighed, and markedwith ear accompaniedby 2 adult males was also captured. Two tags and lip tattoos. bears were collared near the southwesternregion of the Adultfemales were markedwith lip tattoosand ear tags SevernayaZemlya Islands, while the remaining3 bears and fitted with satellite telemetry collars (Telonics Inc., were collared farthersouth near the Russian mainland Mesa, Ariz.) that transmittedfor 7 hours every 5 days (2 (Fig. 2). Bear densities were low with a total of 23 bears bears) or 6 hours every 6 days (3 bears). The satellite (including COYs) sighted during 59 hours and 20 min- telemetrysystem has been describedin detailby Fancy et utes of aerial searching. al. (1988) and Harriset al. (1990). One location/bearper 5- or 6-day duty cycle was selected based upon proce- Movementsand Sea Ice Information dures describedby Arthuret al. (1998). Expected bat- Cumulative distances moved for 3 bears that were tery life was 24 months, although realized battery life trackedfor >1 year were similar (Table 2), but the daily usually was shorter(Garer et al. 1989). rates of movement differed between bears (ANOVA, P < 0.0005). Multiplecomparison tests (S-N-K,P < 0.05) Movementsand Sea Ice Information indicated that bear N7982 moved at a higher daily rate Data were pooled into 4 seasonal categoriesfor analy- than the other 4 bears (Table 2), but the reason for this sis based on the temporal dynamics of sea ice in the differenceis unknown. Daily movementrates of the non- Severnaya Zemlya region (maximum ice, receding ice, lactatingfemale (N7898) did not differ from movement minimumice, and advancingice). The 4 periods repre- ratesof 3 females capturedwith cubs-of-the-year(N7980, sent broadcategories of sea ice conditions during 1991- N7984, and N7986; Table 2). Rates of movement be- 93: maximum sea ice 16 October-31 May; receding sea tween seasons (Table 3) also differed (ANOVA, POLARBEARS OF SEVERNAYAZEMLYA * Belikov et al. 37

Table 1. Characteristics of bears captured and marked near the Severnaya Zemlya Archipelago, Russia, May 1991.

Animal Weight Lengtha Chest No. Age/sex (kg) (cm) girth (cm) Date

N7980 10/F 148 218 119 10 May N7981L COYb/M 10 79 50 10 May N7981R COY/M 9 74 44 10 May N7982 8/F 160 207 116 11 May N7983 COY/F 14 70 48 11 May N7984 7/F 185 194 122 14 May N7985 COY/M 20 105 65 14 May N7986 13/F 192 193 128 15 May N7987R COY/F 28 112 66 15 May N7987L COY/M 32 112 66 15 May N7988C 4/F 160 174 116 15 May a Length for adults is straightline; length for COYs is total length. b Cubs of the year. c Non-lactating female.

P = 0.085), with rates of movement duringthe period of ern or eastern side of the archipelago,depending on the minimumice cover higherthan the other 3 periods (S-N- wind direction. Duringearly September,ice began form- K, P < 0.05). ing among solid remainingice and by the end of Septem- The majorityof the movements were confined to the ber was formingamong remnant ice. In winter 1991-92, easternKara Sea, but 3 bears(N7982, N7984, andN7988) ice growth was more intensive than usual. For instance, did move into the westernLaptev Sea, with 2 returningto fast ice nearCape Cheluskinduring the whole winterwas the Kara Sea prior to collar failure (Fig. 2A-E). One 20-25 cm thickerthan average. bear (N7988) moved east into the Laptev Sea before re- Fast ice was formed to usual limits by early January turning to the eastern edge of the Kara Sea near Cape 1992 in the archipelagoincluding Vilkitski Strait (Fig. Cheluskin, where the collar apparentlyfailed (Fig. 2E). 1B), with little change until ice-breakup. No polynyas One bear (N7980) had restrictedmovements (Fig. 2A) were presentbeyond the fast ice duringJanuary and Feb- thatwere closely associatedwith Komsomoletsand Pioner ruary 1992. During March-April 1992, only limited Islands. The combinedlocations for the 5 collaredbears polynyas were observed. By May-July 1992, an exten- depict a overall range that is centered around the sive zone of remnantice and open water appearedbe- SevernayaZemlya Archipelago(Fig. 2F). yond the fast ice zone both west and east of the archipelago.The fast ice zone in Vilkitski Straitwas bro- Ice Conditions ken by 1 August 1992, and by mid-Augustit was broken During spring 1991 the limits of shorefast ice in the in the westernentrance of the strait. In summer1991 and Severnaya Zemlya Islands archipelago were similar to 1992, fast ice in Shokalskiand Krasnoi Armii Straits,near the averagecondition (Fig. 1B). Fast ice along the west- Pioner and Komsomolets Islands and aroundSedov Is- ern archipelagoincluded considerable portions of second land, was not broken. Ice formationbegan early in au- year ice. Drift ice was also composed of up to 80% sec- tumn 1992 (Fig. 1A): among solid ice by 1 September ond year ice. In April-June 1991 polynyas were not and among remnantice by mid-September. present beyond the shorefast ice zone west and east of the archipelago. A zone of open water and remnantice east of SevernayaZemlya Archipelago and in Velkitski DISCUSSION Straitappeared during early July. Although sample size is small, movements of polar Shorefastice in the westernentrance to Vilkitski Strait bearsin this region of the RussianArctic appearsto be of was partlybroken by mid-July and was completely bro- 2 types: bears that remain closely associated with land ken by the end of July. The straitwas completely broken (islandsor shoreline),and individualswho remainon sea by mid-August. During summer 1991, fast ice in ice duringa majorityof the year. Bears N7980, N7984, Shokalski and Krasnoi Armii Straitsnear Pioner Island and N7986 displayedthe landform-basedmovement pat- and around Sedov Island was not broken. Throughout tern (Fig. 2A, C), while bears N7982 and N7988 had a the entire summer,solid ice approachedeither the west- more ice-related movement pattern(Fig. 2B, E). Bear 38 Ursus 10:1998

Bear #7980 Bear#7982 A. 10 May 1991 - 22 Feb 1993 Novayae Zemlya 11 May 1991 -2 Jul 1992 N 122 655 / = Locations, Days n = 67 Locations, 418 Days 7~ --

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\ LaptevLapte Sea :~~~~~~~~~~~~~~~~~~l~,

I <, i~~~~~~~~~~~~~~f ;? ^--^ a-^-----i-^~ \ '/ (\F Taimyr Peninusla ?' Taimyr Peninusia

Bear #7986 ' - Novaya Zemlya D 15 May1991 23 Sep 1991 ~....Novaya~~~~~~~~~\e m l Z y a n = 13 Locations, 131 Days

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Bear#7988 - . Novaya Zemlya E. 15 May 1991 29 Oct 1992 n = 73 Locations, 532 Days

Kara Sea

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I

\I ,(;Taimyr Ta Peninusla

Fig. 2. Movements of female polar bears in the Severnaya Zemlya Archipelago, May 1991-Feb 1993, as determined by satellite telemetry. A-E. Individual bear movementss (star denotes capture location). F. Cumulative area encompassed by movements of 5 collared female polar bears in the Karaand Laptev Seas, May 1991-Feb 1993. POLARBEARS OF SEVERNAYA ZEMLYA * Belikov et al. 39

Table 2. Movementcharacteristics of female polar bears Garer et al. 1994a) were recordedduring the portionof nearthe SevernayaZemlya Archipelago, Russia, 1991-92. the yearwith minimumice cover, suggestingsimilar habi- tat use strategiesduring this time period for the 2 areas. Number Total Number Meanrate of In female bears in the Viscount Melville Animal of distance of days movement contrast, polar no. locations (km) tracked km/day(SE) Soundregion of the CanadianArctic had movementrates of 6.9 km/day (SE = 5.9) to 8.8 km/day (SE = 6.3) from N7980 122 655 6.1 4,028 (0.62)aa October N7982 67 4,098 418 14.1 (0.84)b April through (data derived from Messier et al. N7984 21 1,746 125 8.9 (1.53)a 1992, Table IV). Messier et al. (1992) speculated that N7986 13 876 131 9.7 (1.97)a this increasedmovement during the summermonths was N7988b 57 10.4 3,493 532C (0.91)a relatedto the periodof hyperphagiawhen seals were most a Mean rates of movement with differentletters are statistically vulnerableto predation. different (Student-Newman-Kuel,P < 0.05). Rates of movementof SevernayaZemlya Archipelago b Non-lactatingfemale at capture,but was in a maternityden from 1 bears duringthe receding, minimum,and maximumice November 1991 through8 April 1992 (161 days). c were one thirdless than those re- Total movement is based on a 371-day period, with the denning periods approximately period deleted. ported for bears in eastern Russia (Garer et al. 1990), while the rateof movementby SevernayaZemlya Archi- bears the ice was less Table3. Dailyrates of movementby ice season for female pelago during advancing period polarbears near the SevernayaZemlya Archipelago, Russia, than half the rate of bears in the Chukchi Sea (7.5 com- 1991-92. pared to 18.9 km/day, respectively). These differences are likely relatedto differences in the sea ice regime for the regions, with the ice extent in the ChukchiSea Rateof movement(km/day) region Ice Number having a difference of approximately1,400 km between season locations mean SE maximum ice cover and minimum ice cover (Garer et Recede 53 9.3aa 0.94 al. 1990). Minimum 70 12.5b 0.81 The area defined by movements of these 5 collared Advance 31 7.5a 1.21 bearsindicate a populationcentered around the Severnaya Maximum 121 9.9a 0.62 Zemlya Achipelago (Fig. 2F). Earlier speculation that a Mean rates of movement with dissimiliarletters are different the Laptev and Kara Seas were comprised of separate (Student-Newman-Kuel,P < 0.005). populationsof polarbears are not supportedby these data. However,these dataare biased because the captureeffort was limited to bears associatedwith the and N7980 had very restrictedmovements during the 655- archipelago northeastern of the Kara Sea the coast of day trackingperiod and was associatedwith fast ice near portions along the Peninsula. The western and easternbound- Komsomoletsand Pioner Islands and the adjacentpolynya Taimyr aries of the cannotbe defined until furtherre- that is normallypresent northwestof the 2 islands (Fig. population search occurs in the Sea and western of 1B). Bear N7988 appearedto have denned along the Laptev portions shoreline of OctoberRevolution Island in the Shokalski the KaraSea. Strait during winter 1991-92, according to sensor data. This bear made the most extensive movement of the 5 LITERATURECITED collared bears into the Laptev Sea and was moving to- wardthe KaraSea when the collar failed October ARTHUR,S.M., G.W. GARNER,AND T.L. OLSON.1998. Identifying during and 1992 mitigating errors in satellite telemetry of polar bears. (Fig. 2E). Ursus 10:413-419. Polar bear movements in other are influenced regions BELIKOV,S.E., ANDT.E. RANDLA.1987. Fauna of birds and sea ice distributionand et al. In by type (Gamer 1994b). mammalsof the Severaya Zemlya. Pages 18-28 in Fauna contrastto the dynamicnature of the sea ice in the Chukchi and ecology of birds and mammalsof Middle .M.: Sea, the sea ice of the Karaand the Laptev seas is more Nauka. (In Russian.) stable, and polar bears in this region of Russia do not FANCY, S.G., L.F. PANK, D.C. DOUGLAS, C.H. CURBY, G.W. need to move long distancesto maintaincontact with the GARNER, S.C. AMSTRUP,AND W.L. REGELIN. 1988. Satellite sea ice as they do in the Chukchi Sea in eastern Russia telemetry:a new tool for wildlife researchand management. U.S. Fish and Wildl. Serv. Resour. Publ. 172. (Garer et al. 1990, 1994a). However, higher average 54pp. GARNER,G.W., S.C. AMSTRUP,D.C. DOUGLAS,AND C.L. GARDNER. daily movement rates for both areas (12.5 km/day for 1989. Performanceand utility of satellite telemetry Severaya Zemlya; 18.5 km/day for the Chukchi Sea, during 40 Ursus 10:1998

field studies of free-rangingpolar bears. Proc. Int. Symp. in Svalbard. J. Wildl. Manage. 35:27-36. Biotelemetry 10:66-76. LENTFER,J.W. 1968. A techniquefor immobilizingand marking , S.E. BELIKOV,M.S. STISHOV,V.G. BARNES,JR., AND S.M. polar bears. J. Wildl. Manage. 32:317-321. ARTHUR.1994a. Dispersal patternsof maternalpolar bears MESSIER,F., M.K. TAYLOR,AND M.A. RAMASAY.1992. Seasonal fromthe denningconcentration on WrangelIsland. Int.Conf. activity patternsof female polar bears (Ursus maritimus)in Bear Res. and Manage. 9:401-410. the Canadianarctic as revealedby satellitetelemetry. J. Zool., , S.T. KNICK, AND D.C. DOUGLAS. 1990. Seasonal London 226:219-229. movements of adult female polar bears in the Bering and PoPov,L.N. 1939. Huntingmammals of easterncoast of Taimyr Chukchi seas. Int. Conf. Bear Res. and Manage. 8:219- peninsula. Pages 87-121 in Proceedingsof researchinstitute 226. of polar agriculture,live-stock farming,and hunting. Issue , I. STIRLING, AND S.E. BELIKOV. 1994b. Habitat 8. (In Russian.) considerationsfor polarbears in the northPacific rim. Trans. RUTILEVSKI,G.L. 1939. Hunting mammals of Cheluskin North Am. Wildl. and Nat. Resour. Conf. 59:111-120. peninsulaand the VilkitskiStrait. Pages 7-60 in Proceedings HAIGH, J.C., I. STIRLING, AND E. BROUGHTON. 1985. of researchinstitute of polaragriculture, live-stock farming, Immobilizationof polarbears (Ursus maritimus Phipps) with and hunting. Issue 8. (In Russian.) a mixture of tiletamine hydrochloride and zolazepam SCHWEINSBURG,R.E., L.J. LEE,AND J.C. HAIGH.1982. Capturing hydrochloride. J. Wildl. Dis. 21:43-47. and handlingpolar bears in the Canadianarctic. Pages 379- HARRIS,R.B., S.G. FANCY,D.C. DOUGLAS,G.W. GARNER,S.C. 386 in L. Neilsen, J.C. Haigh, and M.E. Fowler, eds. AMSTRUP,T.R. MCCABE,AND L.F. PANK. 1990. Tracking Chemicalimmobilization of NorthAmerican wildlife. Wisc. wildlife by satellite:current systems and performance.U.S. HumaneSoc., Inc., Milwaukee. 447pp. Fish and Wildl. Serv. Fish Wildl. Tech. Rep. 30. 52pp. STEEL,R.G.D, ANDJ.H. TORRIE.1980. Principles and procedures HENSEL,R.J., ANDF.E. SORENSEN.1980. Age determinationin of statistics. Second ed. McGraw-HillBook Co., New York, live polar bears. Int. Conf. Bear Res. and Manage. 3:93- N.Y. 633pp. 100. STIRLING, I., C. SPENCER, AND D. ANDRIASHEK. 1989. KIRPICHNIKOV,A.A. 1938. Marineice and migrationsof some Immobilizationof polarbears (Ursus maritimus) with Telazol mammals. Zoologicheski Zhurnal 17(2):273-279. (In in the Canadianarctic. J. Wildl. Dis. 25:159-168. Russian.) URVANTSEV,N.N. 1935. Two years on the SevernayaZemlya. KOSHKIN,V.N. 1937. Some data on hunting fauna of the L.: Glavsevmorputi. 363pp. (In Russian.) Begichev Island. Pages 89-101 in Arctic problems, No 2. VOLKOV,N.A. 1981. Aerial ice reconnaissance text book. (In Russian.) Gidrometeoizdat,Leningrad, Russia. 240pp. (In Russian.) LARSEN,T. 1971. Capturing,handling and markingpolar bears