The Bowhead Whale, Balaena mysticetus: Its Historic and Current Status

Item Type article

Authors Shelden, Kim E. W.; Rugh, David J.

Download date 10/10/2021 01:24:30

Link to Item http://hdl.handle.net/1834/26463 The Bowhead Whale, Balaena mysticetus: Its Historic and Current Status

KIM E.W. SHELDEN and DAVID J. RUGH

Introduction threatened) or whether specific stocks of vertebrate fish or wildlife that inter­ should be removed from the list of En­ breeds when mature" (16 U.S.c. § Bowhead whales, Balaena mysticetus dangered and Threatened Wildlife (50 1532(16). Subspecies and distinct (Fig. 1), are currently listed as endan­ C.P.R. § 17.11). This paper is an up­ population segments are not defined gered under the Endangered Species Act date of literature that has become avail­ within the statute. In 1992, the U.S. Fish of 1973 (ESA) and depleted under the able since the publication of Braham and Wildlife Service (USFWS) issued Marine Mammal Protection Act of 1972 (1984) and a review of elements that a draft policy providing a definition for (MMPA). The ESA, as amended in may contribute to the evaluation of the "distinct population segments," re­ 1978, requires the status of populations status of stocks of bowhead whales. viewed by Rohlf (1994). This policy listed as "endangered" or "threatened" was revised and published in the Fed­ to be reviewed every 5 years. For bow­ Background eral Register (vol. 59, p. 65885) in 1994, head whales, an evaluation is to be made Under the ESA, "endangered" status and the final policy was published in as to whether the status of these whales is designated to those species facing 1996 (Fed. Regist., vol. 61, p. 4722). should be revised (i.e., changed to extinction "throughout all or a signifi­ Within this policy, three elements will be cant portion of [their] range... " (16 considered in the status designation of Kim E.W. Shelden and David J. Rugh are with § the National Marine Mammal Laboratory, Alaska U.S.c. 1532(6)). "Threatened" status population segments: 1) discreteness, 2) Fisheries Science Center, National Marine Fish­ is provided to "... any species which is significance, and 3) conservation status. eries Service, NOAA, 7600 Sand Point Way N.E., likely to become an endangered species Historically, the distribution of these Bin CI5700, Seattle, WA 98115-0070. within the foreseeable future through­ whales was almost circumpolar in the out all or a significant portion of its northern hemisphere (Fig. 2); however, ABSTRACT-The bowhead whale, Balaena mysticetus, is currently listed as endangered range" (16 U.S.C. § 1532(20)). The heavy exploitation by commercial whal­ under the Endangered Species Act of 1973 definition of species within the ESA is ers seriously depleted them throughout and as depleted under the Marine Mammal not based on the taxonomic categories their range. For management purposes, Protection Act of 1972. Literature on the used in the field of biology. Instead the five stocks (geographically distinct seg­ species is updated since 1984, and elements are reviewed that may contribute to the term refers to "... any subspecies of ments of the population) are currently evaluation of the status of bowhead whale fish or wildlife or plants, and any dis­ recognized (IWC, 1992:27). Three of stocks. tinct population segment of any species these stocks are found in the North At-

Figure I. - The bowhead whale, Balaena mysticetus.

57(3-4) 1 lantic and two in the North Pacific, some east as the Laptev Sea. Between 1940 northern Norway, in the waters of or all of which may be reproductively and September 1990, 37 bowhead Zemlya Frantsa-losifa (Franz Josef isolated. whale sightings have been recorded in Land), near Novaya Zemlya, and near Distribution and Abundance this region, but some of these sightings Severnaya Zemlya (Fig. 3). were not unequivocally bowhead whales The majority of observations oc­ Spitsbergen Stock (Moore and Reeves, 1993:Table 9.2). curred between April and September. Commonly referred to as the Green­ Reviewing records from west to east, Overall, group sizes were small, rang­ land whale, bowhead whales in the east­ sightings were scattered along the coast­ ing from I to 15 individuals, with the ern North Atlantic have been observed line of Greenland, in the waters near exception of two groups observed in in the waters north of Iceland and as far Spitsbergen Island, off North Cape in August in Zemlya Frantsa-losifa, "sev­

140 160 W 180 E 40 N

.*'. *-, f .. .,J'.. '". . BERING SEA •

Figure 2. - Circumpolar distribution of bowhead whales (adapted from Reeves and Leatherwood, 1985).

2 Marine Fisheries Review eral tens of whales" in 1981, and "about 66 individuals" in 1983 (Belikov et aI., 1989:255). In 1989, Clark et al. I iden­ tified at least four bowhead whales from acoustic recordings collected off Spits­ bergen. Additional sightings of lone animals were reported in each year dur­ ing 1990-93 in Zemlya Frantsa-losifa (de Korte and Belikov, 1994). Despite extensive research cruises conducted in the Barents Sea (McQuaid, 1986), Ice­ landic waters (Sigurj6nsson, 1985; Sigurj6nsson et aI., 1988; Sigurj6nsson et al.2), Norwegian and adjacent waters (Christensen et aL3), and continued Danish and Norwegian whaling activ­ ity, very few bowhead whales have been seen. It may be that some of the Spitsbergen stock overwinter within the pack ice (Moore and Reeves, 1993), in o regions inaccessible to unreinforced o vessels. Bowhead whales observed from T'"" aircraft were often found in polynyas or leads within solid ice (Belikov et aI., 1989). o Movement patterns of bowhead to whales within the eastern North Atlan­ tic are not well known. Whalers de­ scribed the migration of animals hunted o between Greenland and Spitsbergen Is­ CD land as following a counterclockwise circuit (Southwell, 1898, as summa­ o rized in Ross, 1993). The whales would '=t winter in the water near Iceland and Jan Mayan, then move northeastward as the c ice front retreated during March and ~ April. From May to June, the animals o ::2: summered along the ice edge between o~------t------­c ----+------il CIl Greenland and Spitsbergen before mov­ ., ing south again with the advancing pack :s: "" o ice. Variations in the routes taken dur­ C\l ing the southbound migration have been attributed to the existence of separate o '=t

1 Clark, C. W., L. M. Brown, K. von der Heydt, A. Baggeroer, and 1. Dyer. 1991. Acoustic de­ o tections of bowhead whales from the east CD Greenland-Spitsbergen stock in spring 1989. Int. Whal. Comm. Unpubl. Doc. SC/43/PSI9, 12 p. 2 Sigurj6nsson, J., T. Gunnlaugsson, P. Ensor, M. o Newcomer, and G. Vikingsson. 1990. North At­ o lantic sightings survey 1989 (NASS-89): ship­ to board surveys in Icelandic and adjacent waters July-August 1989. Int. Whal. Comm. Unpubl. Doc. SC/42/021, 29 p. o (.)Q) o 3 Christensen, 1., T. Haug, and N. 0ien. 1990. o ~~ o Review of the biology, exploitation and present to CD abundance of large baleen whales and sperm <0 whales in Norwegian and adjacent waters. lnt. Figure 3. - Arctic seas of the northeast Atlantic with place names men­ Whal. Comm. Unpubl. Doc. SC/42/105, 28 p. tioned in the text (adapted from Moore and Reeves, 1993).

57(3-4) 3 "tribes" (subspecies or species) of bow­ Little is known of the migration pat­ years to summer along the southern head whales (Scoresby, 1820:211) or terns of bowhead whales found in the coast of Spitsbergen, then return east as segregation of the population into age­ northern Barents Sea. Whalers de­ the ice retreated (Zorgdrager, 1720, or sex-specific groups (Southwell, scribed how a group of whales would summarized in Eschricht and Reinhardt, 1898; de Jong, 1983). arrive out of the east during heavy ice 1866). These whales were said to look and behave differently from the other Spitsbergen whales. However, estab­ lishing the existence of any genetic or morphological variations between these groups would be difficult given the re­ duced size of the current population (Reeves, 1980). There is some confusion as to whether the animals present in these waters to­ day are representative of the historic stock. Jonsgard (1981, 1982) expressed the possibility that the Spitsbergen stock may be extinct. Past sightings may in­ clude migrants from the Davis Strait stock or the East Siberian Sea (Bering Sea stock) (Jonsgard, 1981). Whalers reported incidents in which "unsuccess­ fully harpooned" whales from one stock (Davis Strait or Spitsbergen) were later killed or found dead in the waters in­ habited by the other stock (Eschricht and Reinhardt, 1866; Reeves et aI., 1983). In times when these whales were more plentiful, such overlap in ranges may have occurred; however, it is more likely that the present population in the eastern North Atlantic is a remnant of the severely depleted Spitsbergen stock (Reeves and Leatherwood, 1985; McQuaid, 1986; Moore and Reeves, 1993). Possibly numbering only "in the tens" now (Christensen et ai. 3), this group of animals is thought to originally have been the most populous of the bowhead whale stocks (Braham, 1984; Woodby and Botkin, 1993). The major­ ity of islands within this region were uninhabited, and it is likely that this stock rarely experienced human preda­ tion prior to commercial whaling (Ross, 1993). From 1660 to 1912, about 90,000 bowhead whales were harvested by commercial whalers, a total that does not include mortally wounded whales that escaped, whales captured but cast adrift, or cargos of vessels that were lost or captured at sea (Ross, 1993). Woodby and Botkin (1993) reevaluated the his­ torical harvest data and developed a The early whale fishery in Greenland is depicted in this 1781 engraving from Dow (1967), minimum preexploitation estimate of courtesy of and copyright by Argosy Book Store, Inc., N. Y. about 24,000 whales (an earlier estimate

4 Marine Fisheries Review GREENLAND

Ottawa • Islands'

Hudson Bay CANADA

Figure 4. - Northeast Canada and Greenland with place names mentioned in the text (adapted from Moore and Reeves, 1993). by Mitche1l4 came to 25,000). A maxi­ been fully compiled and analyzed dian Arctic Archipelago (Davis Strait mum estimate was not computed be­ (Ross, 1993; Woodby and Bodkin, stock) and the other found in Hudson cause catch and voyage data (covering 1993). Strait, Hudson Bay, and Foxe Basin over 200 years of whaling) have not (Hudson Bay stock) (as defined in Davis Strait Stock Moore and Reeves, 1993; Fig. 4). This Bowhead whales in the western distinction is uncertain given that it is 4 Mitchell, E. D. 1977. Initial population size of North Atlantic are currently segregated not based on genetic or morphological bowhead whale (Balaena mysticetus) stocks: cumulative catch estimates. In!. Whal. Comm. into two stocks, one occupying Davis evidence; instead the two are separated Unpubl. Doc. SC/29/33, 113 p. Strait, Baffin Bay, and along the Cana­ based on their summer feeding distri-­

57(3-4) 5 butions (Reeves et aI., 1983; Reeves and McLaren and Davis6; Richard et a1. 7). From counts obtained during the fall Mitchell, 1990). Unlike the Spitsbergen stock, whalers migration in 1978, Davis and Koski The Davis Strait stock is believed to did not attribute variations in size and (1980) concluded that the population follow a counterclockwise migration migration patterns to the existence of was "in the low hundreds at most." In pattern similar to the Spitsbergen stock multiple stocks in Davis Strait (Reeves 1979, an estimated 140 ± 33 bowhead (Ross, 1993). Animals move northeast et aI., 1983). The recent reidentification whales were observed migrating through from the Labrador Sea across Davis of a whale photographed in Isabella Bay the 1978 study area, separate from an­ Strait to the west coast of Greenland, in September 1986 and again near Disko other group observed summering in following the pack ice as it recedes. Bay, West Greenland, on 10 April 1990 Isabella Bay (Koski and Davis9). Using Some cross Baffin Bay to Lancaster (Heide-Jl/lrgensen and Finley, 1991) the numbers gathered from aerial photo­ Sound, the summer feeding grounds appears to support the hypothesis that identification surveys, Finley (1990) north of Baffin Island, while others con­ these whales belong to one stock. estimated "as many as" 107 individual tinue north along the Greenland coast Recent research on the Davis Strait bowhead whales were in the vicinity of to Kane Basin or west of Baffin Island stock has focused on summer residents Isabella Bay in late September 1986. To into Barrow Strait (Davis and Koski, and southbound migrants utilizing this number Finley added the Koski and 1980; Reeves et aI., 1983). Some ex­ Isabella Bay (Finley, 1990; Richardson Davis9 estimate from 1979 to obtain a change between the eastern North At­ et aI., 1995). Additional sightings in­ "conservative" estimate of 250 animals lantic stocks and the Bering Sea stock clude the entanglement of a bowhead for the entire stock. Zeh et a1. (1993) has been documented (based on whal­ whale off northwest Greenland in early reanalyzed the Finley (1990) data us­ ing irons used in the Davis Strait fish­ November 1980 (Kapel, 1985) and in­ ing a mark-recapture method sampling ery that were found in whales taken in cidental sightings made during surveys only the best quality photo images from the Chukchi Sea) (Bockstoce and for beluga whales, Delphinapterus 1986 (77 photographs) and 1987 (21 Burns, 1993). Segregation by size and leucas, and narwhals, Monodon mono­ photographs). They obtained an esti­ sex appears to occur at this time: cows ceros, conducted off West Greenland in mate of 214 ± 54 bowhead whales in with calves and subadults are found March 1990, 1993, 1994, and April Isabella Bay, to which they added the around the inlets and islands north and 1991 (Reeves and Heide-Jl/lrgensen8). 1979 Koski and Davis9 estimate, yield­ west of Baffin Island while other adults Monitoring this population has proven ing a total abundance of about 350 bow­ occupy the waters along the northeast to be difficult given its large range and head whales in the Davis Strait stock. coastline (Reeves et aI., 1983; Reeves severely reduced numbers (Finley, Because only well marked animals were and MitcheIl5). The summer resident 1990; Reeves and Heide-Jl/lrgensen8). used in the analysis, this estimate is population ofIsabella Bay (east-central Since 1980, no long-term studies have considered to be "conservative" (Zeh et Baffin Island) has consisted predomi­ been made of this stock outside of aI., 1993). Given the low proportion of nantly of adults and large subadults Isabella Bay (Finley, 1990; Reeves and calves (2.2-3.6%) observed during (Finley, 1990; Richardson et aI., 1995). Heide-Jl/lrgensen8). these surveys, recovery from past whal­ Early whalers believed this segregation Current estimates of Davis Strait ing "has been, at best, exceedingly continued through the southbound mi­ stock abundance are based on sightings slow" (Davis and Koski, 1980), and vi­ gration (Southwell, 1898). Cows, gathered by Davis and Koski (1980), ability of the population may be at risk calves, and young whales followed the Finley (1990), and Koski and Davis9. (Finley, 1990). The Davis Strait stock east coastline of Baffin Island back to has had no appreciable evidence of re­ the Labrador Sea, while large adults covery since the period of commercial ("old males") moved south along the 6 McLaren, P. L., and R A. Davis. 1983. Distri­ whaling, over 80 years ago (Reeves et western coast (Southwell, 1898, sum­ bution of wintering marine mammals off West aI., 1983). Greenland and in southern Baffin Bay and north­ marized in Ross, 1993). This may be ern Davis Strait, March 1982. Rep. for Arctic Pi­ Almost 29,000 bowhead whales were true in part as calves and small juve­ lot Proj., Calgary, Alberta, by LGL Ltd., 98 p. harvested in Davis Strait between 1719 niles «10m in length) are rarely ob­ 7 Richard, P., J. Orr, R Dietz, and L. Dueck. 1993. and the end of commercial whaling in served in Isabella Bay (Finley, 1990). Preliminary report on an aerial survey of belu­ 1915 (Ross, 1993) from an estimated gas in the North Water, 21-26 March 1993. Work. Some whales may migrate south along pap. pres. to Joint Commisso Conserv. Manage. original stock of over 11,700 (Woodby the west coast of Greenland (Kapel, Narwhals and Belugas. Sci. Work. Group, and Botkin, 1993). Between 1922 and 1985) while others may overwinter in Copenhagen, 21-23 June. 1975, only seven whales were taken or 8 Reeves, R. R, and M. P. Heide-Jl'lrgensen. 1994. polynyas and flaw zones within the pack Recent observations of bowhead whales offWest struck and lost by Canadian Eskimos ice (Reeves et aI., 1983; Turl, 1987; Greenland. Int. Wha!. Comm. Unpub!. Doc. SCI (Mitchell and Reeves, 1982; Reeves and 461NAll, 9 p. Heide-Jl/lrgensen8), and after 1979, the 9 Koski, W. R, and R A. Davis. 1980. Studies of the late summer distribution and fall migration Canadian government "explicitly banned of marine mammals in NW Baffin Bay and E the hunting of bowheads without a li­ 5 Reeves, R. R., and E. Mitchell. 1991. Summer Lancaster Sound, 1979. Rep. from LGL Ltd. to cense" under the Federal Cetacean Pro­ segregation in the Davis Strait bowhead whale Petro-Can. Explor. Inc., 214 p. Avail. from stock. Int. Wha!. Comm. Unpub!. Doc. SC/431 Pallister Resour. Manage. Ltd., Suite 105,4116­ tection Regulations (Reeves and Mit­ PS28, I p. 64th Ave. S.E., Calgary, Alberta, Can. TIC 3B3. chell, 1990). Though bowhead whales

6 Marine Fisheries Review are currently designated as endangered and in July 1981, twelve whales were riod of time this stock was hunted were by the Committee on the Status of En­ seen in South Bay, Southampton Island incomplete (Bockstoce and Botkin, dangered Wildlife in Canada (COSE­ (Reeves and Mitchell, 1990). Approxi­ 1983). Whales in this stock were dis­ WIC) (Campbell, 1990), a request to mately 23 different bowheads were covered by commercial whalers in 1848 harvest one whale per year by the Inuit identified by McLaren and Davis 11 dur­ (Bockstoce, 1986), but intensive hunt­ of the eastern Northwest Territories has ing aerial surveys conducted over ing did not begin until 1852 when been made under the Nunavut Final Hudson Strait in winter. No single sur­ whales in the Bering Sea stock were no Land Claim Agreement (Indian and vey has adequately covered the range longer as plentiful in "traditional" whal­ Northern Affairs Canada 1993, summa­ of this stock, and therefore it is not pos­ ing areas (Bockstoce and Burns, 1993). rized in Reeves and Heide-Jprgensen8). sible to extrapolate these counts into a By 1860, the Okhotsk Sea stock was In Autumn 1994, a bowhead whale was population estimate. A conservative es­ severely depleted, and whalers had al­ illegally harvested from the Davis Strait timate by Mitche1l4 placed the current ready resumed whaling in the Bering stock by a Canadian native (Marine Mam­ population at 100 or less. Recent ob­ Sea (Bockstoce, 1986). Mitche1l4 esti­ mal Commission10). servations indicate that "at least a few mated the preexploitation size of the tens of whales" from this stock continue population to be 6,500 based on a total Hudson Bay Stock to occupy a large portion of their former estimated catch of 3,506 whales. Ross As mentioned in the preceding Davis range (Reeves and Mitchell, 1990). (1993) suggested that this estimate may Strait Stock section, there is currently Prior to commercial whaling, this be too high for the reasons stated above no evidence to suggest that the Hudson population consisted of about 580 and offered "a conservative, though Bay stock is discrete from it. However, whales (Mitchell4 as modified by mostly speculative, compromise" of3,ooo separating these two populations based Woodby and Botkin, 1993). From 1860 as a minimum population estimate. on their summer distributions allows for to 1915,565 whales were harvested by Though Scammon (1874) stated that a conservative approach to managing commercial whalers (Ross, 1993). Be­ bowheads were hunted "throughout the each one (Reeves and Mitchell, 1990). tween 1919 and 1976, 28 bowhead whole extent" of the Okhotsk Sea, cer­ From the middle of May until Septem­ whales were killed or struck and lost by tain areas were occupied by concentra­ ber, bowhead whales of the Hudson Bay native hunters in western Atlantic wa­ tions of animals during the summer stock are commonly observed near ters (Reeves et aI., 1983; Reeves and months (Fig. 5). In the northeastern Southampton Island in northwestern Mitchell, 1990). Of these, 23 were from Okhotsk Sea, whales were found in Hudson Bay and along the northern tip the Hudson Bay stock. Since 1980, Penzhinskaya Gulf and Gizhiginskaya of the Melville Peninsula in Foxe Ba­ other incidents of bowheads being fired Gulf. The next area of concentration sin (Moore and Reeves, 1993; Fig. 4). upon have been reported (as cited in was to the southwest in Tauyskaya Bay. Scattered sightings also have occurred Reeves and Mitchell, 1990). There has Farther south, the best whaling grounds along the western coastline of Hudson been no appreciable recovery of this were within the gulfs and bays south of Bay and near the Ottawa Islands and stock, in part attributed to occasional the Shantarskiye Islands and west of Mansel Island of eastern Hudson Bay hunting by natives (Reeves and Heide­ Sakhalin Island; Moore and Reeves (Reeves and Mitchell, 1990). Bowhead Jprgensen8) and predation by killer (1993) provide additional details. Al­ whales observed in Hudson Strait, whales (see section on Natural Mortal­ most all of the areas where summer con­ which connects the Bay to the Labra­ ity) (Reeves et aI., 1983; Reeves and centrations of bowhead whales occurred dor Sea, are believed to be migrants Mitchell, 1990). in the past are still occupied today leaving the Hudson Bay area in the au­ (Table 1). As recent as August 1995, Okhotsk Sea Stock tumn or returning in the spring (Finley during joint Russian-American surveys, et al., 1982; Reeves and Mitchell, 1990). There has been some difficulty in a few dozen bowhead whales were ob­ A portion of the population overwinters assessing the historical distribution and served in a feeding aggregation south in the upper Bay in leads and polynyas abundance of bowhead whales in the of the Shantarskiye Islands (Brownell 12). formed near islands (Low, 1906; Ross, Okhotsk Sea. Right whales, Eubalaena Berzin et ai. (1990) estimated the popu­ 1975; Reeves and Mitchell, 1987). glacialis, and gray whales, Eschrichtius lation in this area to be at least 250­ Sightings usually have been of small robustus, were sometimes misidentified 300 animals. An estimate of abundance groups of bowhead whales, rarely ex­ as bowhead whales, and whaling of 300-400 was made for the entire ceeding two individuals (Reeves et aI., records maintained during the short pe­ Okhotsk Sea based on data collected 1983, Reeves and Mitchell, 1990). Fif­ since 1979 (Vladimirov, 1994:Table 1). teen whales (in groups of 1, 2, 3,4, and However, "no quantitative data are 5) were observed in upper Foxe Basin 11 McLaren, P. L., and R. A. Davis. 1982. Winter available to confirm" these estimates on 20 August 1983 (Orr et aI., 1986), distribution of Arctic marine mammals in ice­ covered waters of eastern North America. Rep. for Petro-Can. Explor. Inc., Calgary, Alberta, by LGL Ltd. Offshore Labrador BioI. Stud. 10 Marine Mammal Commission. 1995. Annual (OLABS) program rep., 151 p. Avail. from 12 Brownell, R. L. NMFS, Southwest Fisheries report to Congress, 1994. Mar. Mamm. Comm., Pallister Resour. Manage. Ltd., Suite 105,4116­ Science Center, La Jolla, CA, 92038. Personal 1825 Conn. Ave. NW., Wash., D.C. 20009, 270 p. 64th Ave. S.E., Calgary, Alberta, Can. T2C 3B3. commun.

57(3-4) 7 Okhotsk Sea

0:o~ShOi Shantar c::JMayy1 Shantar p IS'''''-1/seIiChiy Is. l> Bering Sea

Kamchatka Tauyskaya Bay __.c:::o... Peninsula

. lany Island RUSSIA Okhotsk Sea

J Kuril Islands a f o 100 200 300 400 500km I I I I I I

Figure 5. - Okhotsk Sea and environs with place names mentioned in the text (adapted from Moore and Reeves, 1993).

(Berzin et al. 13). There is some specu­ ing the summer in the northeastern Fedoseev (1984) observed bowhead lation as to whether animals found dur­ Okhotsk Sea form a distinct population whales deep in the ice north of Sakhalin from those in the Shantarskiye region. Island in 1969, 1981, and 1983, in ad­ The winter distribution of both of these dition to one sighting east of Sakhalin 13 Berzin, A. A., S. A. Blokhin, H. Minakuchi, R. L. Brownell, A. M. Burdin, and V. N. groups is unknown. Island in 1981 and another a little over Burkanov. 1995. Bowhead and gray whale popu­ Berzin et al. (1991) noted that by 200 km south ofTauyskaya Bay in 1982 lations in the Okhotsk Sea. In Abstracts of the North mid-November, bowhead whales were (Table 2). Because whalers left the Pacific Marine Science Organization (PICES): workshop on the Okhotsk Sea and adjacent areas, no longer found in the Shantarskiye re­ Okhotsk Sea before the onset of winter Vladivostok, Russia, 19-24 June 1995, p. 44-45. gion, despite the waters being ice-free. storms in early November and did not

8 Marine Fisheries Review Table 1.-Recent sightings of bowhead whales in historic summer concentration areas In the Okhotsk Sea. Commas separate slghtings that were made at different times (days or months) during the survey year. grate north and east, following leads in the sea ice in the eastern Chukchi Sea 2 2 2 5 5 General location of sightings 1982' 1983' 1984' 1986 ,3 1987 ,4 1988 1989 1990 until they pass Point Barrow where they N,E, Okhotsk Sea travel east toward the southeastern Penzhinskaya Gulf U· U U U U 2 U U Gizhiginskaya Gulf U U U 17 U 7 19,36 U Beaufort Sea (Braham et al., 1980; Brahamet al., 1984; Marko and Fraker 18). S, W. Okhotsk Sea Between Bol'shoi Shantar Island Most of the summer (June through Sep­ and Utichiy Island U U U 18 U U 8,14,21,33 U Tugurskiy Bay U 3 U U U U U 1 tember), bowhead whales range through Konstantina Bay 13 2 20 U 47 72 U 44,7+20 the Beaufort Sea (Hazard and Cubbage, Ulbanskiy Bay 15 U 11 U 40 >30 72',1's Akademiya Bay U 39 U U U U U 1982; Richardson et al., 1987; McLaren and Richardson l9; Richardson et al. 20 ; , Data from Berzin et al. (text footnote 39), Surveys occurred during 1-7 September 1982, 30 August-7 September 1983, and 21-23 August, 20 October 1984, Richardson et a],2I). Spatial distribution 2 Data from Berzin et al. (1990), The 1986 sightin9 occurred in early June (N,E, Okhotsk Sea); the 1987 si9hting was in seems to vary between years (Richard­ July; the 1988 sightings occurred in late May (N,E. Okhotsk Sea), on 7 AU9ust (Konstantina Bay, including 2 calves), 22 and on 20 October (Ulbanskiy Bay), son et al., 1987; Davis et a1. ; Thomson 3 Data from IWC (1988:113), The 1986 sighting occurred in September (S.w, Okhotsk Sea), et a1. 23) affected in part by surface tem­ 4 Data from Berzin et al. (1988), The 1987 sighting occurred in October (Ulbanskiy Bay). 5 Data from Berzin et al. (1991), The 1989 sightings occurred on 23 May (19), 24 May (36 including 2 calves), May 28 (8), perature or turbidity fronts and anoma­ the morning of 31 May (14), the afternoon of 31 May (21),1 June (33), late August (72), and 28-30 September ("singly 23 and far apart"). The 1990 si9htings occurred on 24-25 June (1),27 August (44 around the cape separating Konstantina lies (Borstad, 1985; Thomson et al. ). and Ulbanskiy Bay), and 28 August (7 near the cape, 20 spread througn Ulbanskiy and Akademiya Bay). Very few bowhead whales are found 6 U = unknown whether the area was surveyed or not. , This sighting is incorrectly attributed to the 1989 season in Berzin et al. (1991); the sighting occurred during the 1988 in the Chukchi Sea in summer (Miller season only (Brownell, text footnote 12), . et aI., 1986); however, there have been enough sightings to indicate that some Table 2.-Bowhead whales observed outside 01 summer concentration areas in the Okhotsk Sea (from Fedoseev, whales do not migrate to the Beaufort 1984). Sea, at least during some years. Many Date Location Lat. Long, Group size small groups have been observed trav­ 8 Apr. 1969 N. of Sakhalin Island 55Q1O'N 144Q30E 6 eling northwest along the Chukchi Pen­ 11 Apr, 1981 55Q20'N 144Q15E 23 55Q15'N 144QOOE 2 insula in May (Bogoslovskaya et aI., 7 May 1981 56QOO'N 141 Q20E 8+7 9 May 1981 54Q58'N 143Q15E 7 21 Dec,1983 57Q30'N 143QOOE 2 18 Marko, R., and M. A. Fraker. 1981. Spring 1981' E. of Sakhalin Island Not noted in text 3 J. 29 Dec, 1982 S. of Tauyskaya Bay 56Q22'N 150QOOE 4 ice conditions in the Beaufort Sea in relation to bowhead whale migration. Rep. for Alaska Oil , Sightin9 was noted on Figure 1 (Fedoseev, 1984) but not described within the text. Gas Assoc., Anchorage, by LGL Ltd., 98 p, 19 McLaren, P. L., and W. J. Richardson, 1985. Use of the eastern Alaskan Beaufort Sea by bowheads return until June, there are no historic western Bering Sea in winter (Novem­ in late summer and autumn. In LGL and Arctic Science, Importance of the eastern Alaskan Beau­ records on bowhead whale distribution ber to April), generally associated with fort Sea to feeding bowheads: literature review and during this whaling hiatus, leaving the the marginal ice front, and found near analysis, p. 7-35. Rep. for U.S. Minerals Manage. possibility that there originally was a the polynyas of St. Matthew and St. Servo by LGL, Inc., and Arctic Sci., Ltd., 158 p. 20 Richardson, W. J., B. Wtirsig, G. W. Miller, common stock between the Okhotsk Lawrence Islands and the Gulf of and G, Silber. 1986. Bowhead distribution, num­ and Bering Seas (Townsend, 1935). Anadyr (Bogoslovskaya et aI., 1982; bers and activities. In W. J. Richardson (Editor), However, other authors (Bockstoce and Brueggeman, 1982; Braham et al., Importance of the eastern Alaskan Beaufort Sea l4 15 to feeding bowhead whales, 1985, p. 146-219. Botkin, 1983; Lindholm ) have argued 1984; Ljungblad et a1. ; Brueggeman Rep. for U.S, Minerals Manage, Servo by LGL 16 17 that the Okhotsk Sea stock has always et a1. ; Bessonov et a1. ; Fig. 6). From Inc., OCS Study MMS 86-0026, NTIS No. been a population discrete from the April through June, these whales mi­ PB87-124350, 315 p. Bering Sea stock. 21 Richardson, W. J., B. Wtirsig, and G. W. Miller. 1987. Bowhead distribution, numbers and activi­ 15 Ljungblad, D. K., S, E. Moore, 1. T. Clarke, ties.ln W. J. Richardson (Editor), Importance of Bering Sea Stock and 1. C. Bennett. 1986, Aerial surveys of en­ the eastern Alaskan Beaufort Sea to feeding bow­ dangered whales in the northern Bering, eastern head whales, 1985-86, p. 257-368. Rep. for U.S. The Bering Sea stock, also referred Chukchi and Alaskan Beaufort Sea, 1985: with Minerals Manage. Servo by LGL Inc., NTIS No. to as the western Arctic stock or the a seven year review, 1979-85. Rep. for U.S. Min­ PB88-150271, 547 p. Bering-Chukchi-Beaufort stock, has erals Manage. Servo by Nav. Ocean Systems 22 Davis, R. A., W. R. Koski, and G. W. Miller. 1983, Cent., NTIS No. PB87-115929, 443 p. been studied more extensively than any Prelimary assessment of the length-frequency dis­ 16 Brueggeman, 1. 1., B. Webster, R. Grotefendt, tribution and gross annual recruitment rate of the of the aforementioned stocks. This stock and D. Chapman. 1987. Monitoring the winter western arctic bowhead whale as detennined with is widely distributed in the central and presence of bowhead whales in the Navarin Ba­ low-level aerial photogrammetry, with comments sin through association with sea ice. Rep. for U.S. on life history. Rep. to NMFS, Natl. Mar. Mamm. Minerals Manage. Servo by Envirosphere Co., Lab., Seattle, Wash., by LGL Ltd., 91 p. 14 Lindholm, 0.1863. Whales and how tides and NTIS No. PB88-101258, 179 p. 23 Thomson, D. H., D. B. Fissel, J. R. Marko, R. currents in the Okhotsk Sea affect them. [From 17 Bessonov, B" V. V. Mel'nikov and V. A. Bobkov. A. Davis, and G. A. Borstad, 1986. Distribution an unidentified journal with news from the Far 1990. Distribution and migration of cetaceans in of bowhead whales in relation to hydrometeoro­ East of Russia, p. 42-43. In Russ. Trans!. by L. the Soviet Chukchi Sea. In Conference Proceed­ logical events in the Beaufort Sea. Environ. Stud. A. Hutchinson for Univ. Calif. San Diego, La ings, Third Information Transfer Meeting, p. 25­ Revolving Funds Rep. 028, 119 p, Can. Dep. In­ 10lla, 1965, 12 p.] 31. U.S. Minerals Manage. Serv., Alaska OCS Reg. dian N. Affairs, Ottawa.

57(3-4) 9 o-wrangell. Beaufort Sea

Chukchi Sea 70°

Cape 8erdtse-Kamen Cape Netten CANADA

ALASKA

81. Matthew I.

./II 60°

Bering Sea

50°..1..----,..------,.------,.------.------..------' 180° 160° 140° Figure 6. - Bering Sea and environs with place names mentioned in the text.

1982; Bessonov et al. 17; Ainana et al.24; Ze1ensky et al. 25), and about 60 whales traveled northwest past Cape Serdtse­ Kamen 15-28 June 1990 (Mel'nikov 24 Ainana, L., N. Mymrin, L. Bogoslovskaya, and 25 Zelensky, M., V. V. Mel'nikov, and V. V. and Bobkov, 1993). One group of 7 1. Zagrebin. 1995. Role of the Eskimo Society of Bichkov. 1995. Role ofthe Naukan Native Com­ Chukotka in encouraging traditional Native use pany in encouraging traditional Native use of whales was observed off Cape Netten, of wildlife resources by Chukotka Natives and wildlife resources by Chukotka Native people and Chukchi Peninsula, on 26 July 1991 in conducting shore based observations on the in conducting shore based observations on the dis­ distribution of bowhead whales, Balaena tribution ofbowhead whales, Balaena mysticetus, traveling north, and a group of 7 was mysticetus, in coastal waters off the south-east­ in waters of the Bering Sea and Chukchi Sea ad­ seen here on 27 September traveling ern part of the Chukotka Peninsula (Russia) dur­ jacent to the Chukotka Peninsula (Russia) during east (Mel'nikov and Bobkov, 1994). ing 1994. Rep. to Dep. Wild!. Manage., North 1994. Rep. to Dep. Wild\. Manage., North Slope Slope Borough, Barrow, Alaska, by Eskimo Soc. Borough, Barrow, Alaska, by Eskimo Soc. Farther northwest, near Cape Schmidt, Chukotka, Provideniya, Russia, 135 p. Chukotka, Provideniya, Russia, 105 p. single animals were observed on 5 Au-

10 Marine Fisheries Review Aerial view of a bowhead whale mi­ grating past Point Barrow, Alaska, during the 1992 spring migration. A column of vapor, the "blow," exits the open blowholes. White on the chin and tail are characteristic of bowheads, and acquired marks along the dorsal surface allow for reidentification of this individual if photographed again. Photo by Wayne Perryman, NMFS.

A bowhead whale feeds in the Beau­ fort Sea (swimming to the left in this image). The high blowhole, "neck," and broad back are characteristic of this species. Photo by David Rugh, NMFS.

Aerial view of a bowhead whale adult and calf migrating past Point Barrow, Alaska, during the 1992 spring migration. The calf is in the typical swimming position beside the adult and between the pectoral flipper and fluke. Photo by Robin Westlake, NMFS.

57(3-4) gust and 1 September by crew members early winter (late October and Novem­ females harvested in the spring has in­ of the ice-breaker Krasin. Studies con­ ber) they arrive in the Bering Sea creased from 4.7% (1973-77) to 20.4% ducted in 1994 have shown the presence (Kibal'chich et al., 1986; Bessonov et (1986-92) (Braham, 1995). Immature ofbowhead whales throughout the sum­ al.!?), where they remain until the fol­ whales «13 m) made up the largest mer along the southeastern portion of lowing spring migration. percentage of the harvest; 80% prior to the Chukchi Peninsula (127 sightings Segregation by size and sex class 1978 and 60% since 1978 (Braham, in June, 59 in July, 5 in August, and 6 occurs in three overlapping pulses dur­ 1995). Records from the historical com­ in September (Ainana et al.24)) and the ing the spring migration, the first con­ mercial catch were not as well main­ easternmost portion of the Peninsula (21 sisting of subadults, the second of larger tained. Unlike the commercial whalers sightings in June and 39 in August whales, and the third composed of even in the North Atlantic region, whalers in (Zelensky et al. 25)). larger whales and cows with calves the North Pacific sector did not docu­ During late summer (early Septem­ (Rugh, 1990; Angliss et ai., 1995; ment size or sex differences between the ber to mid-October), bowhead whales Nerini et al.3!). The reverse appears to aggregations of animals they hunted migrate west out of the Beaufort Sea, occur along the Chukchi Peninsula; (Bockstoce and Bums, 1993). as evidenced during aerial surveys Russian natives noted the appearance of Whales taken by commercial whal­ (Richardson 26; Ljungblad et ai. 27), ra­ large numbers of mothers with calves ers in the Bering Sea may have been dio-tracking (Wartzok et al. 28), and sat­ in late March and early April followed representatives of a population that did ellite- tracking (Mate and Krutzikow­ by immature and adult animals (Bogo­ not migrate (Bockstoce and Botkin, Sky29). From mid-September to mid­ slovskaya et al., 1982). In the Beaufort 1983; Bockstoce, 1986; Fraker33). Af­ October they are seen in the northeast Sea in summer, aggregations have usu­ ter reducing the number of bowhead Chukchi Sea, some as far north as lat. ally consisted of only juveniles or of whales in the Bering Sea to a point nON (Moore et al., 1986; Moore and large whales that may include calves where they were uneconomical to hunt, Clark30). Whales migrate from Point (Richardson26, Davis et al. 32). In 1983, the whalers turned their attention to the Barrow into the Chukchi Sea heading Cubbage and Calambokidis (1987) Okhotsk Sea. And when the Okhotsk toward Wrangel Island (Mate and found a significant inverse correlation Sea stock was nearly eliminated, the Krutzikowsky29). When they reach the between longitude and size class; en­ whalers explored the Chukchi Sea. Be­ Siberian coast, they follow it southeast counter rates for larger whales increased cause they did not find whales in the to the Bering Strait (Bogoslovskaya et moving west to east in the Beaufort Sea. western Chukchi Sea, they continued al., 1982; Ze1ensky et al. 25). In 1991, Onshore and offshore distributions var­ east to the Beaufort Sea where large fall migrants, presumably returning ied annually, suggesting that "sex- or numbers were found (Bockstoce and from the Beaufort Sea, did not begin age-class segregation patterns are tem­ Bums, 1993). There has been difficulty passing Cape Netten until 10 Novem­ porally and spatially fluid and cannot in assessing the total number of animals ber (Mel'nikov and Bobkov, 1994). By be defined rigidly for any region or pe­ killed in the Bering Sea stock. Only riod" (Moore and Reeves, 1993:351). 19% of commercial whaling records Segregation by size also occurs during collected from 1849 to 1914 have been 26 Richardson, W. J. (Editor). 1987. Importance the fall migration (Braham, 1995). extensively analyzed (Bockstoce and of the eastern Alaskan Beaufort Sea to feeding bowhead whales, 1985-86. Rep. for U.S. Miner­ George et al. (1995) showed a clear Botkin, 1983). Woodby and Botkin als Manage. Servo by LGL Inc., NTIS No. PB88­ trend in progressively smaller whales (1993) reviewed the summaries of esti­ 150271,547 p. harvested between August and Novem­ mated annual kills from pelagic and 27 Ljungblad, D. K., S. E. Moore, J. T. Clarke, ber. Along the Chukchi Peninsula, the shore-based fisheries compiled by and J. C. Bennett. 1987. Distribution, abundance, behavior and bioacoustics ofendangered whales fall migration splits into two pulses Marquette and Bockstoce (1980), in the Alaskan Beaufort and eastern Chukchi (Bogoslovskaya et ai., 1982; Mel'nikov Bogoslovskaya et al. (1982), Bockstoce Seas, 1979-86. Rep. for U.S. Minerals Manage. Servo by Nav. Ocean Systems Cent., NTIS and Bobkov, 1993; 1994), though seg­ and Botkin (1983), Breiwick and No.AD-AI83934/9, 391 p. regation by size or sex class was not Mitchell (1983), Breiwick et al. (1984), 28 Wartzok, D., W. A. Watkins, B. Wiirsig, J. confirmed as the cause. Sonntag and Broadhead (1989), Mit­ Guerrero, and J. Schoenherr. 1990. Movements The sex ratio of whales harvested by chell4, and Braund et al. 34. They created and behaviors of bowhead whales. Rep. from Purdue Univ., Fort Wayne, for AMOCO Prod. Alaska natives from 1973 to 1992 was a time series of harvest data showing Co., Anchorage, 197 p. equal, though the proportion of adult cumulative kill records for each year 29 Mate, B. R., and G. Krutzikowsky. 1995. Ap­ plication of remote methods of large cetacean tracking: bowhead whales. Rep. for U.S. Miner­ als Manage. Servo by Hatfield Mar. Sci. Cent., 3\ Nerini, M. K., D. Withrow, and K. Strickland. 33 Fraker, M. A. 1984. A brief review of recent Oregon State Univ., Newport. OCS Study MMS 1987. Length structure ofthe bowhead whale popu­ information on the responses of bowhead whales 95-0053,174 p. lation derived from aerial photogrammetry, with (Balaena mysticetus) to offshore petroleum op­ 30 Moore, S. E., and J. T. Clarke. 1992. Distribu­ notes on recruitment spring 1985 and 1986. Int. erations. Int. Whal. Comm. Unpubl. Doc. SC/361 tion, abundance and behavior of endangered Whal. Comm. Unpubl. Doc. SC/39/PSI4, 22 p. PSI7, 19 p. whales in the Alaskan Chukchi and western 32 Davis, R. A., W. R. Koski, G. W. Miller, P. L. 34 Braund. S. R.. W. M. Marquette. and J. R. Beaufort Seas, 1991: with a review 1982-91. McLaren, and C. R. Evans. 1986. Reproduction Bockstoce. 1988. Data on shore-based bowhead Rep. for U.S. Minerals Manage. Serv., Anchor­ in the bowhead whale, summer 1985. Int. Whal. whaling at sites in Alaska. Int. Whal. Comm. age, Alaska, by SAIC, Marlt. Servo Div., 237 p. Comm. Unpubl. Doc. SC/38/PS2, 123 p. Unpubl. Doc. SC/40/PSIO, 9 p.

12 Marine Fisheries Review from 1848 to 1991. According to these among ten Alaskan native villages higher levels of epibenthic organisms data, 22,174 bowhead whales have been (IWC,1995;MarineMammalCommis­ were found in the stomachs of small harvested from a stock that may have sion10, Zeh et aI.36). Requests to harvest whales « 10.5 m in length) (Lowry, numbered between 10,000 and 23,000 bowhead whales have also been put 1993), and it appears that copepods be­ before commercial exploitation. From forth by Canadian and Russian natives. come increasingly more important in 1992 to 1994, 113 bowhead whales In 1991, Aklavik hunters in the western the diet oflarger whales (Schell et al. 37). were landed and 35 were struck but lost Canadian Arctic requested a permit to Crustacean zooplankton was the domi­ during native subsistence hunts in kill one or strike two bowhead whales nant prey in both males (89%) and fe­ Alaska (Suydam et aI., 1995). from the Bering Sea stock (Freeman et males (79%) (Lowry, 1993). Several different methods have been aI., 1992). Permission was granted by As evidenced in stomach contents used to compute the current population the Canadian government in August collected between April and June, some size of the Bering Sea stock (reviewed 1991, and one whale was harvested in bowhead whales feed opportunistically by Zeh et aI., 1993). In 1994, the Inter­ Mackenzie Bay in the autumn of 1991 during the spring migration (George and national Whaling Commission's (lWC) (Stoker and Krupnik, 1993; Marine Tarpley, 1986; Carroll et al., 1987; Scientific Committee reviewed data Mammal CommissionlO, Zeh et aI.36). Lowry, 1993). In most cases, half or collected during the 1993 visual and Additional licenses were granted in more of the stomachs were empty (3 of acoustic census (Zeh et aI., 1995; 1993 and 1994, though bowhead whales 6 near St. Lawrence Island, 3 of 4 at Raftery and Zeh35 ) conducted near were not harvested in either year (Ma­ Point Hope, 10 of 12 at Wainwright, and Point Barrow. They settled on a current rine Mammal Commission10, Zeh et 23 of 36 near Point Barrow). Stomachs population estimate of 7,992 bowhead ai.36). Aboriginal harvests along the collected south of Point Barrow were whales (95% c.1.: 6,900-9,200) (IWC, Chukchi Sea coasts usually yielded 8­ from small whales and contained 1995; Marine Mammal CommissionlO). 10 bowhead whales in "good years" in "shrimp," gammarid amphipods or very Zeh et aI.36 continued to refine their es­ the early 20th century, but by the mid­ small «3.1 mm) copepods. Stomachs timate using newly available acoustic 20th century, bowhead whale hunting collected from whales collected near data. The 1988 Bayes empirical Bayes was almost completely replaced by gray Point Barrow generally contained either method yielded a population estimate whale hunting (Krupnik, 1987). Poverty copepods, euphausiids, or both, though of 7,500 (95% c.1.: 6,400-9,200). An in the villages along the Chukchi Sea the dominant prey often varied between alternative method, the NiP4 method, coast has brought about a renewed in­ individuals harvested in the same year. which compared the estimated number terest in hunting one or two bowhead During the fall migration, only 2 of 15 of whales passing within the viewing whales a year (Bogoslovskaya et aI., whales harvested off Kaktovik (central ) 17). range of census observers (N4 and the 1982; Bessonov et aI. Beaufort Sea) and 1 of 6 taken near proportion detected by the hydrophone Barrow had empty stomachs (Lowry, Life History and Ecology array (P4), resulted in an estimate of 1993). Sex- and length-related differ­ 8,000 (95% c.1.: 6,900-9,200) for the Most of what is known of the life his­ ences in diet were not noted in the 11 1993 census. The Bayes empirical tory and ecology ofbowhead whales has samples collected at Kaktovik (4 fe­ Bayes method "provided the best infor­ been derived from studies of the Bering males and 7 males) that underwent de­ mation on population size for assess­ Sea stock. The data presented in this tailed examination. Copepods were pre­ ment purposes" while the NiP4 method section applies only to this stock unless dominant in six of the stomach samples, provided "the best available estimate of otherwise stated. euphausiids in three, mysids in one, and the current size" of the Bering Sea stock one contained almost equal parts of all Feeding (lWC, 1995; Zeh et aI.36). An annual three organisms. Euphausiids were the rate of increase of 3.1 % (95% c.1.: 1.4­ Food habits studies conducted on 35 dominant prey item found in five large 4.7%) was computed for the Bering Sea bowhead whales (21 males and 14 fe­ whales (>14 m) harvested near Point stock (lWC, 1995; Zeh et aP6). males) harvested between 1975 and Barrow in September. A quota of 204 bowhead whales has 1989 by Alaskan natives were reviewed In years when oceanographic condi­ been set for 1995-98 (see section on by Lowry (1993). Most prey species tions are favorable, the lead system near Management), based on a stated need identified from bowhead whale stom­ Point Barrow may serve as an impor­ for 51 whales per year to be divided ach contents were crustacean zooplank­ tant feeding ground in the spring ton, particularly euphausiids and cope­ pods ranging in length from 3 to 30 mm. 35 Raftery, A. E., and J. E. Zeh. 1994. Bowhead Epibenthic organisms, mostly mysids 37 Schell, D. M., S. M. Saupe, and N. Haube­ whale, Balaena mysticetus, population size esti­ and gammarid amphipods, were also nstock. 1987. Bowhead whale feeding: alloca­ mated from acoustic and visual census data col­ common in stomach contents, with only tion of regional habitat importance based on lected near Barrow, Alaska, in 1993. Int. Wha!. stable isotope abundances. In W. J. Richardson Comm. Unpub!. Doc. SC/46/ASI3, 25 p. a small sampling of benthic species (Editor), Importance of the eastern Alaskan Beau­ 36 Zeh, J. E., A. E. Raftery, and A. A. Shaffner. (Lowry, 1993). Age-related differences fort Sea to feeding bowhead whales 1985-86, p. 1995. Revised estimates of bowhead population 369-415. Rep. to U.S. Minerals Manage. Servo size and rate of increase. Int. Wha!. Comm. were difficult to establish given the lim­ by LGL Eco!. Res. Assoc. Inc., NTIS No. PB88­ Unpub!. Doc. SC/47/AS 10,26 p. ited sample size; however, slightly 150271.

57(3-4) 13 (Carroll et aI., 1987). Other important crustaceans he called "squillae and calving interval in bowhead whales has feeding grounds exist in the Beaufort shrimps." Scoresby's drawings were been corroborated by photo-identifica­ Sea (Lowry and Frost, 1984; Ljungblad examined by Ruud (1937) who deter­ tion evidence (Miller et aI., 1992; Rugh et aI., 1986; Schell and Saupe, 1993; mined that the "squillae" were partially et aI., 1992). The highest sighting rate Wtirsigetal., 1985; 1989; Thomson and digested euphausiids. of calves during the spring migration Richardson38), along the northern occurred in 1993. This sighting rate, Reproduction Chukchi Peninsula (Mel'nikov and when graphed with earlier shore-based Bobkov, 1993; 1994; Moore et aI., Although apparent sexual activity census and aerial photogrammetric 1995) and possibly in the Bering Sea occurs among bowhead whales most sightings, provides additional support (Schell and Saupe, 1993). Schell and months of the year, studies of bowhead for a 3-4 year variable recruitment pat­ Saupe (1993) and Schell et a1. 37 believe fetuses indicate conception typically tern (George et aI., 1995). that a significant amount offeeding may occurs during late winter or early Gestation lasts 13-14 months (Nerini occur outside the eastern Beaufort Sea, spring, as summarized by Koski et aI. et aI., 1984) or 12-16 months (Tarpley 41 particularly by older whales in the fall (1993:249). Not all sexual activity leads et a1. ). Calves are 4.0-4.5 m in length and early winter when they are in the to conception; fecundity of some bow­ when born and increase from modal Chukchi and Bering Seas. These con­ heads is in doubt with the discovery of lengths of 4.75-5.0 m in late May to clusions are based on patterns of iso­ pseudohermaphroditism in at least two 6.25-6.5 m in late summer, a growth tope variation found in the visceral fat males with testicular feminization rate of 1.5 cm/day (Koski et aI., 1993), and muscle sampled from three adults (Tarpley et aI. 40), a relatively high inci­ much less than the 2.88 em/day calcu­ and six subadults. This variability sug­ dence rate as only 76 bowheads have lated for southern right whales (Best and gests that older animals are feeding in been closely examined between 1980 Rtither42). Weaning occurs 9-15 months different areas or on different prey types and 1989 (Philo et aI., 1993). Calves are postpartum (Nerini et aI., 1984) with than younger animals. Feeding behav­ usually born between April and early about 95% of the yearlings weaned by ior has been observed during the sum­ June during the spring migration (Koski the next spring migration (Rugh et aI., mer in the Beaufort Sea and the fall in et aI., 1993) probably peaking in May 1992). Yearlings are 6.6-9.4 m long in the Chukchi Sea but not in the winter (Nerini et aI., 1984). Most (76%) photo­ the spring according to data from har­ in the Bering Sea, as summarized by grammetrically measured calves were vested whales (Nerini et aI., 1984) and Lowry (1993). within 1 m of their average length in 7.0-8.7 m in summer based on photo­ The occupation of summer concen­ late spring (4.25-5.25 m) indicating a grammetric lengths (Koski et aI., 1993). tration areas by bowhead whales in the majority are born within a restricted Growth rates appear to slow after wean­ Okhotsk Sea is believed to be influ­ period (Koski et aI., 1993). Pregnancy ing. Small bowheads reidentified be­ enced by distributions of prey (Berzin rates have been documented as 0.21 tween years had growth rates of much et al. 39), though feeding aggregations pregnancies/mature female/year (Tar­ less than 1 m/yr (Koski et aI., 1992; 41 32 were only reported in 1979 and 1995 pley et aI. , which includes results from Davis et aI. ). Carbon isotope analy­ (Brownell I2, Berzin et a1. 39). In the Nerini et aI., 1984) or 0.20-0.35 sis, using apparent oscillations in iso­ Davis Strait stock, feeding behavior was (George et aI., 1995) with a possible topic ratios in tissue samples from ba­ observed in bowhead whales summer­ increase in pregnancy rates of whales leen to indicate annual cycles, also sug­ ing in Isabella Bay, Baffin Island harvested since 1985 (George et aI., gests bowhead whales grow slowly, tak­ (Finley, 1990). Most of this activity took 1992). These pregnancy rates suggest ing on the order of two decades to reach place in deep (>200 m), glacial troughs that mature female bowheads have calv­ sexual maturity (Schell et aI., 1989; where large concentrations of Calanus ing intervals of3.5-7.1 years (Nerini et Schell and Saupe, 1993). This growth copepods occurred (Finley, 1990; aI., 1984) or 4 years (Tarpley et aI. 41 ), rate is much slower than that of other Nicklin, 1995; Richardson et aI., 1995). comparable to calving intervals of right baleen whales (Lockyer, 1981; 1990; Information on food habits of the whales (Kraus et aI., 1986; Bannister, Payne et aI., 1990). Conventional tech­ Spitsbergen stock is sparce. Scoresby 1990; Best, 1990; Hamilton and Mayo, niques to age bowhead whales have been (1820) found Greenland whale stom­ 1990; Payne et aI., 1990). The 3-4 year considered unsuccessful (Nerini43), in achs contained predominantly small part because of the poor correlation be­ tween whale size and indicated age, but 40 Tarpley, R. J., G. G. Stott, R. F. Sis, M. J. some evidence, such as ivory or stone 38 Thomson, D. H., and W. J. Richardson. 1987. Shively, and G. H. Jarrell. 1985. Further obser­ Integration. In W. J. Richardson (Editor), Impor­ vations on the morphology of the reproductive harpoon heads found in five recently tance of the eastern Alaskan Beaufort Sea to feed­ tract of the bowhead whale, Balaena mysticetus. ing bowhead whales 1985-86, p. 449--479. Rep. In T. F. Albert (Editor), Third conference on the to U.S. Minerals Manage. Servo by LGL Ecol. biology ofthe bowhead whale, Balaena mysticetus: 42 Best, P. B., and H. RUther. 1989. Aerial photo­ Res. Assoc. Inc., NTIS No. PB88-150271. extended abstracts and panel discussion, Anchor­ grammetry of southern right whales-a prelimi­ 39 Berzin, A. A., V. L. Vladimirov, and N. Y. age, p. 115-119. nary report. Int. Whal. Comm. Unpubl. Doc. SCI Doroshenko. 1985. The distribution and numbers 41 Tarpley, R., R. Weeks, and G. Stott. 1988. 411PS4, 18 p. ofcetacea in the Okhotsk Sea: results from aerial Observations on reproductive morphology in the 43 Nerini, M. K. 1983. Age determination tech­ surveys. Int. Whal. Comm. Unpubl. Doc. SC/371 female bowhead whale (Balaena mysticetus). Int. niques applied to mysticete whales. M.S. thesis. 05,7 p. Whal. Comm. Unpubl. Doc. SC/401PS8, 50 p. Univ. Wash., 51 p.

14 Marine Fisheries Review harvested whales, suggest that bowhead proportion of immature whales in the net in Japan (Nishiwaki and Kasuya, whales may live >50 years (Philo et al., population. For modeling purposes, 1970) and another in the waters of 1993) or>75 years (George et al., 1995). Chapman (1984) used an estimated northwest Greenland in a net used to Results from female bowhead whales crude birth rate of 0.072 (S.E. 0.014). capture beluga whales (Kapel, 1985). examined for corpora albicantia or cor­ Koski et al. (1993) reviewed available Between 1976 and 1992, only three pora lutea indicate sexual maturity be­ aerial photograrnmetric data along with ship-strike injuries were documented gins when whale lengths exceed 14.2 rates ofsightings ofcalves from ice-based out of a total of 236 bowhead whales m, and perhaps as small as 12.3 m observers. These data suggested variable examined from the Alaskan subsistence 41 (Nerini et aI., 1984; Tarpley et aI. ). recruitment in bowheads with a 3--4 year harvest (George et al., 1994). These low Aerial photograrnmetry indicates whales cyclicity (also described in Rugh et al., numbers suggest that either bowhead as small as 12.2 m were accompanied 1992; Withrow and Angliss, 1992). The whales do not often encounter vessels, 22 by calves (Davis et al. ). Smaller females mean crude birth rate for all available in­ avoid interactions with vessels, or that tend to calve later in the spring migration formation from 1982-89 (summarized in interactions usually result in the death than larger females: 1.5% (1168) of the Koski et al., 1993:264) was 0.052. Even of the animal. Exposure to man-made adults with calves photographed in the with this low birth rate, the Bering Sea noise and contaminants may produce spring were <13.5 m long compared to bowhead population is increasing, sug­ short- and long-term effects (Richard­ 12% (7/59) in the summer (Koski et al., gesting a low rate of mortality (Koski et son and Malme, 1993; Bratton et aI., 44 1993). There is virtually no information al., 1993; Whitcher et al. ). 1993) that may compromise the health on size at maturation for males, except and reproductive performance of some Morbidity and Mortality that, in general they tend to be smaller whales. The effects ofrepeated encoun­ than females (Nerini et al., 1984). Although bowhead whale mortality ters with seismic vessels, drill rigs, hy­ Using aerial photogrammetric tech­ caused by the subsistence harvest is drocarbons, and heavy metals are un­ niques, Angliss et al. (1995) summa­ fairly well documented (see section on known. Anthropogenic impact is a func­ rized and revised results from surveys Distribution and Abundance), particu­ tion of the extent that industrial activi­ conducted from 1985-92 (originally larly since 1977 in the Bering Sea stock, ties coincide with the bowhead whales' 31 reported in Nerini et al. ; Withrow and little is known about naturally occurring seasonal occupation of certain regions Angliss, 1992; 1994). These show that diseases and death in bowhead whales and the whales' tolerance level of the 5.2% ofthe sampled population of bow­ (e.g., Heidel and Albert, 1994). Since impacts (Richardson and Malme, 1993; head whales in the Barrow area in the 1964, at least 36 cases exist in which Bratton et aI., 1993). spring were calves «6 m), 53.7% were the cause of death could not be estab­ It is not known whether bowhead juveniles (6-13 m), and 41.1 % were lished (Philo et aI., 1993). Some ofthese whales suffer from stress-induced bac­ adults (>13 m). These authors also deaths may be the result of hunter-in­ terial infections similar to those ob­ showed that age segregation occurred flicted wounds. Types of human-in­ served in captive cetaceans (Buck et al., during the spring migration: 80% of the duced injuries include embedded shrap­ 1987). Studies of harvested bowhead measured animals early in the migra­ nel and harpoon heads from hunting whales have provided information on tion were juveniles, dropping to 20% attempts, rope and net entanglements bacterial, mycotic, and viral infections by the end of the migration. Adults with from harpoon lines and crab pots, and but not the level to which they contrib­ calves did not appear until the last few collisions with vessels, summarized in ute to mortality and morbidity (Philo et weeks of the migration. This seasonal Philo et al. (1993). Bowhead whales fre­ aI., 1993). Skin lesions, found on all pattern was also evident in aerial pho­ quently react to being struck with a har­ harvested bowhead whales, were not tographic data showing increased de­ poon by rotating rapidly (Philo et aI., malignant or contagious. However, po­ grees of graying on the peduncles (a 1993). This type of reaction near ropes tentially pathogenic microorganisms sign of aging) as the migration season or nets would only increase the likeli­ inhabit these lesions and may contrib­ progressed (Rugh, 1990) and in sizes hood of entanglement. Whales with ute to epidermal necrosis and the spread of whales harvested at Barrow (George ropes caught in their baleen and with of disease (Shotts et aI., 1990). Expo­ et aI., 1995). Cubbage and Calam­ scarring caused by rope entanglement sure of these roughened areas of skin bokidis (1987) described segregation of have been observed during the harvest to environmental contaminants, such as bowheads by size in the Beaufort Sea, (Philo et aI., 1992) and in aerial photo­ petroleum products, could have signifi­ further confounding efforts to represen­ graphs of live animals (NMML, unpubl. cant effects (Shotts et al., 1990; Aibert45); tatively sample the population for calv­ data). Although incidental take of bow­ although, Bratton et al. (1993) con­ ing rates. On the other hand, Clarke et head whales is apparently rare, there has al. (1987) found little analytical evidence been one reported entrapment and death 45 Albert, T. F. 1981. Some thoughts regarding of geographic or temporal segregation of of a young bowhead whale in a fishing the possible effect of oil contamination on the bowhead whale, Balaena mysticetus. In T. F. calves during the fall migration. Albert (Editor), Tissue structural studies and other Population modeling by Breiwick et 44 Whitcher, B., J. E. Zeh, D. J. Rugh, W. R. investigations on the biology ofendangered whales Koski, and G. W. Miller. 1996. Estimation of in the Beaufort Sea, p. 945-953. Rep. to Bur. Land al. (1984) underlined the need for im­ adult bowhead whale survival rates. Int. Whal. Manage. from Dep. Vet. Sci., Univ. Md., Coil. Park, proved estimates of calf production and Comm. Unpubl. Doc. SC/48/ASI2, 22 p. NTIS Accession No. PB86-l53566.

57(3-4) 15 cluded that such encounters were not also provide some protection from killer whaling which ended over 100 years likely to be hazardous. whale attacks. The frequency of attacks ago, there are still only 3Q0-400 whales. Of 130 bowhead whales examined is unknown and killer whale distribu­ The Bering Sea may have had a stock between 1980 and 1989, only one whale tion in northern waters has not been well that was eliminated, except for the com­ had a tumor, though this may be the re­ documented (George et al., 1994). In the ponent that migrated to the Beaufort sult of incomplete necropsies performed Davis Strait stock, about one-third (5 Sea. This stock was reduced from at on the other whales (Migaki and Albert, of 16) of the summer residents photo­ least 10,300 animals, and has been re­ 1982; Philo et aI., 1993). The tumor, graphed in Isabella Bay showed evi­ covering slowly over the past 80 years located on the liver, was benign. Ac­ dence of killer whale scarring along to a current population of about 8,000. cording to Philo et ai. (1993:295), "it is their flukes and sometimes their flip­ There is evidence that bowhead whales unlikely that tumors are major contribu­ pers (Finley, 1990). Two attacks were are long-lived animals. It is therefore tors to bowhead whale morbidity or witnessed during Finley's study and possible that in the greatly reduced mortality." Marine caliciviruses first appear to occur frequently; Inuit hunt­ stocks, some of the animals have sur­ discovered in California sea lions, ers even have a phrase for the behav­ vived nearly since the termination of Zalophus californianus californianus, iors observed during an attack, "Ardlin­ commercial whaling, but fecundity rates are known to be widespread and trans­ gayuq." Of 195 whales examined dur­ are so low that very few new whales are ferable between terrestrial and marine ing the Alaskan subsistence harvest being added to the respective stocks. hosts (Smith et al., 1986; Barlough et (1976-92), 8 had been wounded by Calving intervals of 3-4 years and the aI., 1986; Smith et aI., 1987; Smith et killer whales (George et aI., 1994). possibility that bowheads whales do not al. 46). Though not isolated from bow­ Seven of the eight bowhead whales were become sexually mature until they are head whale tissues, there is evidence greater than 13 m in length, suggesting 20 years old may, in part, explain these that these whales may be hosts for these either that scars are accumulated over slow recovery rates in stocks with only viruses (Smith et aI., 1986; Smith et aI., time, or young animals do not survive a a few hundred whales. 1987; Smith et aI. 47). Symptoms of killer whale attack. Hunters on St. The IWC is the primary body respon­ calicivirus infection include formation Lawrence Island reported two small «9 sible for conservation and management of cysts, open sores, inflammation of the m) bowhead whales found dead as a of bowhead whale populations world­ lungs, brain, heart, and stomach and result of killer whale attacks (George wide. All stocks of bowhead whales are intestine lining, and abortion (Barlough et aI., 1994). Overall, the frequency of classified as "protected" by the IWC. et aI., 1986; Smith et aI., 1986; Smith attacks on bowhead whales in the As mentioned in the Introduction, the et aI., 1987; Smith et aI. 46). How much Bering Sea stock appears to be low United States further classified all bow­ these viruses contribute to natural mor­ (George et aI., 1994). Attacks witnessed head whales as endangered under the tality and possibly to reduced reproduc­ in the Okhotsk Sea at the tum of the ESA and depleted under the MMPA. tion in the bowhead whale population century are reported in Mitchell and Currently, the bowhead whales in the is unknown (Philo et aI., 1993). Reeves (1982). Recent accounts include Bering-Chukchi-Beaufort Seas repre­ Evidence of ice entrapment and pre­ information on the distribution and sent the largest surviving stock. This is dation by killer whales, Orcinus orca, abundance of killer whales in the the only stock, mandated by the IWC has been documented in almost every Okhotsk Sea, but additional attacks have and through exemptions under the ESA bowhead whale stock. The percentage not been reported (Vladimirov, 1994). and MMPA, still harvested by aborigi­ of whales entrapped in ice is considered From the available data, it is not pos­ nal hunters in Alaska. Thus far, a quota to be small, given that this species is so sible to assess the level of depredation to harvest bowhead whales from the strongly ice-associated (Tomilin, 1957; on bowhead whales by killer whales, Bering Sea stock and the other stocks Mitchell and Reeves, 1982; Nerini et al., particularly in terms of size-class selec­ has not been provided to Russian or 1984; Philo et aI., 1993). The ice may tion and encounter rates. Canadian natives by the IWC (Stoker and Krupnik, 1993; IWC, 1995). As Management determined by the IWC's Scientific 46 Smith, A. w., D. E. Skilling, and K. Benir­ Clearly, bowhead whale stocks are Committee, "the total of whales landed schke. 1981. Investigations of the serum antibod­ ies and viruses of the bowhead whale, Balaena slow to recover, and some might not in the four years 1995-98 should not mysticetus. In T. F. Albert (Editor), Tissue struc­ recover at all. The Spitsbergen stock exceed 204, with a maximum number tural studies and other investigations on the bi­ was reduced from 24,000 to a few of 68 strikes in 1995,67 in 1996,66 in ology ofendangered whales in the Beaufort Sea, p. 233-254. Rep. to Bur. Land Manage. from "tens" of whales and has not recovered 1997, and 65 in 1998." (lWC, 1995:22). Dep. Vet. Sci., Univ. Md., College Park, NTIS in the past 80 years. The Davis Strait Any unused portion of the strike quota No. PB86-153566. and Hudson Bay stocks declined from will be carried forward from that year 47 Smith, A. W. 1979. Serum antibodies and vi­ ruses. In J. J. Kelley and G. A. Laursen (Editors), about 12,300 whales to less than 450 and added to the strike quota of any Investigation of the occurrence and behavioral currently, although significant whaling subsequent years, provided that no more patterns of whales in the vicinity of the Beaufort has not occurred in 80 years. The than 10 strikes are added to the strike Sea Lease Area, p. 445-457. Rep. to Bur. Land Manage. from Nav. Arctic Res. Lab., Barrow, Okhotsk Sea stock was originally quota for anyone year. The average Alaska, NTIS No. PB86-153582. around 3,000 whales, but after severe number of whales harvested per year

16 Marine Fisheries Review from 1989 to 1993, including those and DeMaster49). Because the Alaska 1985; Ahmaogak53), there is insufficient struck and lost, was 42 (Suydam et ai., Eskimo subsistence harvest "is man­ evidence about cumulative and long­ 1995). aged under an international regime that term effects of anthropogenic noises Since 1981, the harvest has been differs from the MMPA amended leg­ (Richardson and Malme, 1993). The monitored by the Alaska Eskimo Whal­ islation," the IWC quota supercedes any spring season appears to be a particu­ ing Commission (AEWC) through a quotas set by the MMPA(Small and larly critical period in the bowheads' Cooperative Agreement with the Na­ DeMaster49). Reviews of observer data annual cycles as this is the time all, or tional Oceanic and Atmospheric Ad­ and vessel logbooks indicate that no most, of the population migrates, often ministration (NOAA). This Cooperative incidental takes of bowhead whales by into areas covered by dense ice, where Agreement will remain in effect until commercial fisheries have occurred in migration routes are constrained and the year 2000. Also since 1981, the U.S. waters (Small and DeMaster49). most likely to be blocked by noise AEWC has channeled funds to the North Incidental takes of bowhead whales in sources (Richardson et al. 52). Concep­ Slope Borough (NSB) for censusing the fisheries have rarely been reported and tion and parturition happen during the bowhead whale population as it migrates are thought to not be an issue of con­ spring, as well as some feeding. This is past Point Barrow in the spring (Monta­ cern; in particular because the habitat also the season of the most intense har­ gue, 1993). The AEWC has also been selected by bowheads (ice-covered vest by subsistence hunters. responsible for allocating IWC quotas seas) limits commercial or sport fisher­ Currently, bowhead whales in the among its member communities and has ies activities (Small and DeMaster49). Bering Sea are classified as a "strategic worked to improve hunting methods and Impacts from industrial development stock" under the amended MMPA. technology to reduce the number of (particularly offshore oil extraction) are Stocks are classified as strategic when whales struck but lost. Emphasis has ofconcern as most habitat of the Bering one of the following three criteria are been placed on promoting understand­ stock of bowheads is within active or met: 1) "level of direct human-caused ing of the needs of native Alaskan whal­ potential lease zones. But studies indi­ mortality exceeds the potential biologi­ ers and obtaining quotas that will meet cate that bowhead behavior is often tem­ cal removal level"; 2) "based on the best these needs while still ensuring recov­ porarily affected when exposed to close available scientific information, [the ery of the bowhead whale population. approaches by ships, seismic vessels, stock] is declining and is likely to be Roles have included habitat manage­ and aircraft. Reactions are less obvious listed as a threatened species under the ment and protection in light ofincreased when the noise source is fairly constant, Endangered Species Act of 1973 within commercial activity in the Arctic such as with distant seismic or drilling the foreseeable future"; or 3) [the stock] (Stoker and Krupnik, 1993). work, but migrating bowheads some­ "is listed as a threatened or endangered Under the 1994 reauthorization of the times adjust their course to divert species under the Endangered Species MMPA, the National Marine Fisheries around stationary sources of man-made Act of 1973 (16 U.S.C. 1531 et seq.), Service (NMFS) and U.S. Fish and noise (LGL and Greeneridge50, Hall et or is designated as depleted under this Wildlife Service (USFWS) developed ai. 51 , Richardson et ai.52). Although act" (Marine Mammal Commission48). a method for establishing minimum al­ there has been concern expressed by The level of human-caused mortality lowable incidental takes of marine residents of the Arctic (Ahmaogak, and serious injury during the subsis­ mammals by commercial fisheries. This tence harvest of Bering Sea bowhead potential biological removal level whales, 42 animals per year, does not (PBR) is defined as "the maximum 49 Small, R. J., and D. P. DeMaster. 1995. Alaska exceed the PBR nor the IWC quota for number of animals, not including natu­ marine mammal stock assessments 1995. U.S. 1995. The lower end of the optimum Dep. Commer., NOAA Tech. Memo. NMFS­ ral mortalities, that may be removed AFSC-57, 93 p. sustainable population (OSP) range was from a marine mammal stock while al­ 50 LGL and Greeneridge. 1987. Responses of calculated to vary between 6,500 and lowing that stock to reach or maintain bowhead whales to an offshore drilling opera­ 10,500, using an initial stock size of tion in the Alaskan Beaufort Sea, autumn 1986. its optimum sustainable population" Rep. for Shell W. E. P. Inc., Anchorage, Alaska, 12,599 (95% C.1. 10,945-l7,43l)(IWC, (Marine Mammal Commission48). Us­ from LGL, Ltd., King City, ant., Can., and 1995) and the assumption that the maxi­ ing the minimum population estimate Greeneridge Sci. Inc., Santa Barbara, Calif, 371 p. mum net productivity level (MNPL) is 51 Hall, J. D., M. L. Gallagher, K. D. Brewer, P. (Nmin) of7,524 bowhead whales for the R. Regos, and P. E. Isert. 1994. ARCO Alaska 60% of carrying capacity (K). Objec­ Bering Sea stock, one-half the maxi­ Inc. 1993 Kuvlum exploration area site specific tive criteria for defining when this popu­ mum theoretical net productivity rate monitoring program: final report. Rep. for ARCO lation should be downlisted to threat­ Alaska Inc., Anchorage, from Coastal Offshore (0.5Rma..) of 4%, and a recovery factor Pac. Corp., Walnut Creek, Calif., 219 p. ened or delisted under the ESA have not (F,) of 0.5, resulted in a PBR of 75 ani­ 52 Richardson, W. J., C. R. Greene Jr., J. S. Hanna, been developed to date. mals from the current population (Small W. R. Koski, G. W. Miller, N. J. Paternaude, and M. A. Smultea. 1995. Acoustic affects of oil pro­ duction activities on bowhead and white whales visible during the spring migration near Pro Bar­ 53 Ahmaogak, G. 1989. Protecting the habitat of 48 Marine Mammal Commission. 1995. The row, Alaska-1991 and 1994 phases. Rep. for the bowhead whale. In L. Rey and V. Alexander Marine Mammal Protection Act of 1972, as U.S. Minerals Manage. Servo from LGL, Ltd., (Editors), Proceedings of the sixth conference of amended. Mar. Mamm. Comm., 1825 Conn. Ave. King City, ant., Can. OCS Study MMS 95-0051, the Comite Arctique International, New York, 13­ N.W., Wash. D.C. 20009, 299 p. 539 p. 15 May 1985, p. 593-597.

57(3-4) J7 The agency plans to develop objec­ ____ , , and . 1991. COSEWIC fish and marine mammal subcom­ Results of aerial surveys to study the distri­ mittee status reports: VI. Can. Field-Nat. tive criteria to determine whether this bution and abundance of whales in the Sea of 104:1-6. stock's current classification is accurate Okhotsk in 1988-1990. In L. A. Popov (Edi­ Carroll, G. M., J. C. George, L. F. Lowry, and K. tor), Research papers on marine mammals of O. Coyle. 1987. Bowhead whale (Balaena or, if not, whether it should be listed as the North Pacific, 1989-1990, p. 4-14. mysticetus) feeding near Point Barrow, threatened or removed from the list of VNIRO, Moscow, U.S.S.R [Environ. Can. Alaska, during the 1985 spring migration. endangered and threatened wildlife. Transl. 4083780, 200 p.] Arctic 40: 105-110. Best, P. B. 1990. Natural markings and their use Chapman, D. G. 1984. Estimates of net recruit­ One possible approach which will be in determining calving intervals in right ment of Alaska bowhead whales and of risk investigated is that used by IUCN­ whales off South Africa. S. Afr. J. Zool. associated with various levels of kill. Rep. lnt. World Conservation Union. This system 25:114-123. Whal. Comm. 34:469-471. Bockstoce, J. R 1986. Whales, ice and men: the Clarke, J. T., S. E. Moore, and D. K. Ljungblad. of classification consists of three levels history of whaling in the western Arctic. Univ. 1987. Observations of bowhead whales (i.e., critical, endangered, and vulner­ Wash. Press, Seattle, 400 p. (Balaena mysticetus) calves in the Alaskan _--,__ and D. B. Botkin. 1983. The histori­ Beaufort Sea during the autumn migration, able), each level representing a differ­ cal status and reduction of the western arctic 1982-85. Rep. lnt. Whal. Comm. 37:287-293. ent probability of extinction within a bowhead whale (Balaena mysticetus) popu­ Cubbage, J. c., andJ. Calambokidis. 1987. Size­ specific period of time (Mace and lation by the pelagic whaling industry, 1848­ class segregation of bowhead whales dis­ 1914. Rep. lnt. Whal. Comm., Spec. Iss. cerned through aerial stereophotogrammetry. Lande, 1991). The mCN criteria also 5:107-141. Mar. Mamm. Sci. 3:179-185. take into account extent of occurrence, ____ and J. J. Burns. 1993. Commercial Davis, R, and W. Koski. 1980. Recent observa­ area of occupancy, number of locations whaling in the North Pacific sector. In J. J. tions of the bowhead whale in the eastern Burns, J. J. Montague, and C. J. Cowles (Edi­ Canadian high Arctic. Rep. lnt. Wha!. Comm. or subpopulations, and number of ma­ tors), The bowhead whale, p. 563- 577. Soc. 30:439-444. ture animals. Additional objective cri­ Mar. Mamm. Spec. Pub!. 2. de Jong, C. 1983. The hunt of the Greenland teria based on recently revised Recov­ Bogoslovskaya, L. S., L. M. Votrogov, and I. I. whale: a short history and statistical sources. Krupnik. 1982. The bowhead whale off Rep. Int. Whal. Comm., Spec. Iss. 5:83-106. ery Plans, as appropriate, will also be Chukotka: migrations and aboriginal whaling. de Korte, J., and S. E. Belikov. 1994. Observa­ considered. Because population esti­ Rep. Int. Whal. Comm. 32:391-399. tions of Greenland whales (Balaena mysti­ mates are extremely low for the Spits­ Borstad, G. A. 1985. Water colour and tempera­ cetus), Zemlya Frantsa-Iosifa. Polar Rec. ture in the southern Beaufort Sea: remote sens­ 30(173): 135-136. bergen, Davis Strait, Hudson Bay, and ing in support of ecological studies of the Dow, G. F. 1967. Whale ships and whaling; a Okhotsk Sea stocks, these should con­ bowhead whale. Can. Tech. Rep. Fish. Aquat. pictorial history of whaling during three cen­ turies. Argosy Antiquarian Ltd., N.Y., 446 p. tinue to retain endangered status. Sci. 1350, 68 p. Braham, H. W. 1984. The bowhead whale, Balaena Eschricht, D. F., and J. Reinhardt. 1866. On the mysticetus. Mar. Fish. Rev. 46(4):45-53. Greenland right-whale (Balaena mysticetus, Literature Cited _..,---,_ . 1995. Sex and size composition of Linn.), with special reference to its geographic Ahmaogak, G. 1985. Comments regarding the bowhead whales landed by Alaskan Eskimo distribution and migrations in times past and development of a policy pertaining to research whalers. In A. P. McCartney (Editor), Hunt­ present, and to its external and internal char­ in the U.S. Arctic. Inuit Stud. 9(2):27-32 ing the largest animals: native whaling in the acteristics. In W. H. Flowers (Editor), Recent Angliss, R. P., D. J. Rugh, D. E. Withrow, and R. western Arctic and subarctic, p. 281-313. Can. memoirs on the cetacea, p. I-ISO. Ray Soc., C. Hobbs. 1995. Evaluations of aerial photo­ Circumpolar lnst. Stud. Whal. 3, Occas. Publ. Lond. gram metric length measurements of the 36,345 p. Fedoseev, G. A. 1984. [Encountering whales in Bering-Chukchi-Beaufort Seas stock of bow­ ____ , M. A. Fraker, and B. D. Krogman. the ice fields of the Sea of Okhotsk.] head whales (Balaena mysticetus). Rep. Int. 1980. Spring migration of the western Arctic Ekologiya 3:81-83. [In Russ., partial transl. Whal. Comm. 45:313-324. population of bowhead whales. Mar. Fish. by S. Smrstik (NMML) for this review.] Bannister, J. L. 1990. Southern right whales off Rev. 42(9-10):36-46. Finley, K. J. 1990. Isabella Bay, Baffin Island: western Australia. Rep. lnt. Whal. Comm., ____ , B. D. Krogman, and G. M. Carroll. an important historical and present-day con­ Spec. Iss. 12:279-288. 1984. Bowhead and white whale migration, centration area for the endangered bowhead Barlough, J. E., E. S. Berry, D. E. Skilling, and distribution, and abundance in the Bering, whale (Balaena mysticetus) of the eastern A. W. Smith. 1986. The marine calicivirus Chukchi, and Beaufort Seas, 1975-78. U.S. Canadian Arctic. Arctic 43:137-152. story-part I. Compend. Continuing Educ. Dep. Commer., NOAA Tech. Rep. NMFS ___ , G. W. Miller, M. Allard, R A. Davis, Practicing Vet. 8:F5-F13. SSRF-778, 39 p. and C. R Evans. 1982. The belugas (Delphi­ Belikov, S. E., Yu. A. Gorbunov, and V. I. Bratton, G. R, C. B. Spainhour, W. Flory, M. napterus leucas) of northern Quebec: distri­ Shil'nikov. 1989. Distribution of pinnipedia Reed, and K. Jayko. 1993. Presence and po­ bution, abundance, stock identity, catch his­ and cetacea in Soviet arctic seas and the tential effects of contaminants. In J. J. Bums, tory and management. Can. Tech. Rep. Fish. Bering Sea in winter. Sov. J. Mar. BioI. J. J. Montague, and C. J. Cowles (Editors), Aquat. Sci. 1123,57 p. 15(4):251-257. The bowhead whale, p. 701-744. Soc. Mar. Freeman, M. M. R, E. W. Wein, and D. F. Keith. Berzin, A. A., V. L. Vladimirov, and N. V. Mamm. Spec. Publ. 2. 1992. Recovering rights: bowhead whales and Doroshenko. 1988. Results of aerial surveys Breiwick,1. M., L. L. Eberhardt, and H. W. Braham. Inuvialuit subsistence in the western Canadian to study the distribution and abundance of 1984. Population dynamics of western Arctic Arctic. Can. Circumpolar Joint Fish. Manage. cetaceans in the coastal waters of the Sea of bowhead whales (Balaena mysticetus). Can. J. Committ., Stud. Whal. 2, ISS p. Okhotsk in 1986--1987. In N. S. Chernysheva Fish. Aquat. Sci. 41:484-496. George, J. C., L. M. Philo, R. Suydam, R. Tarpley, (Editor), Scientific research on sea mammals ____and E. D. Mitchell. 1983. Estimated and T. F. Albert. 1992. Summary of the 1989 of the northern part of the Pacific Ocean in initial population size of the Bering Sea stock and 1990 subsistence harvest of bowhead 1986-1987, p. 16-22. VNIRO, Moscow, of bowhead whales (Balaena mysticetus) from whales (Balaena mysticetus) by Alaskan Es­ U.S.S.R. [Can. Transl. Fish. Aquat. Sci. 5506, logbook and other catch data. Rep. lnt. Whal. kimos. Rep. Int. Whal. Comm. 42:479-483. 195 p.] Comm., Spec. Iss. 5:147-151. _~__ , , K. Hazard, D. Withrow, -.,..-7""7" ' , and . 1990. Brueggeman, J. J. 1982. Early spring distribu­ G. M. Carroll, and R Suydam. 1994. Fre­ Aerial surveys to determine the distribution tion of bowhead whales in the Bering Sea. J. quency of killer whale (Orcinus orca) attacks and number of polar whales and beluga whales Wildl. Manage. 46:1036-1044. and ship collisions based on scarring on bow­ in the Sea of Okhotsk in 1985-1989. In A. A. Buck, J. D., L. L. Shepard, and S. Spotte. 1987. head whales (Balaena mysticetus) of the Berzin (Editor), Questions relating to the ra­ Clostridium perjringens as the cause of death Bering-Chukchi-Beaufort seas stock. Arctic tional exploitation of marine mammals in the of a captive Atlantic bottlenosed dolphin 47(3):247-255. far eastern seas, p. 22-34. TINRO, Vladi­ (Tursiops truncatus). J. Wildl. 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