Predation by Marine Mammals on of the Eastern North Pacific Ocean and the Bering Sea

CLIFFORD H. FISCUS

Introduction in the northeastern Pacific Ocean eco­ some extent on these . system. Major predators are discussed here. When marine mammals (with the ex­ This paper is an attempt to consolidate Important sources of information on ception of a few cetaceans) forage for and make available information from the predator-prey relation between ma­ food over the continental shelf, usu­ several sources on the relationship be­ rine mammals and squids of the epipe­ ally comprise most of their prey. Along tween squids and their major marine lagic, mesopelagic, and the continental the continental slope fish and cephalo­ mammal predators. Most of the small shelf waters in the eastern North Pacific pods are equally important, and in the cetaceans, several of the large cetaceans, Ocean are the annual reports of agencies deeper waters of the North Pacific Ocean and most prey on and of the United States (National Marine and the Bering Sea squids form a major octopus when available. Fisheries Service, NOAA) and Canada part of their diet. In the eastern North Marine mammals inhabiting the east­ (Department of the Environment) on Pacific Ocean the neritic squid Loligo ern North Pacific Ocean and Bering Sea pelagic fur seal research carried out opalescens is the most important are listed in Table 1. Of the 45 marine during 1958-74. These reports provide preyed on by marine mammals over the mammals listed in Table 1,26 are known information on species identified from continental shelf. However, the data to prey on squid or octopus in varying stomachs of northern fur seals, and gen­ available on consumption of squid by degrees, and it is quite likely that many eral locations where these seals were marine mammals have not been system­ of the remaining 20 may also prey to taken. In the past several years, data atically reviewed. Parts of this paper were originally prepared for a workshop on squids held at the Seattle Laboratory of the NMFS Table 1.- Marine mammals of the eastern North Pacific Ocean, Bering, and Chukchi Seas. Those known to prey on Northwest and Alaska Fisheries Center cephalopods are marked with an asterisk ('). (Names are aller Rice, 1977.) on 19-20 March 1981. The purpose of Common name Scientific name Common name Scientific name the workshop was to gather information Walrus Odobenus rosmarus Spotted Stenella attenuata' on the squid resources of the region, California Zalophus ca/ifornianus' Striped dolphin Stenella coeru/eoa/ba' Northern sea lion Eumetopias jubatus' Saddleback dolphin De/phinus de/phis' principally from central California to the Callorhinus ursianus' Pacific whiteside Guadalupe fur seal Arctocepha/us townsend; dolphin Lagenorhynchus ob/iquidens' Bering Sea, for potential development Sea otter Enhydra lutris' Northern right of squid fisheries. The various species of Harbor seal Phoca vitu/ina richardsi' whale dolphin Lissodelphis borealis' Spotted or larga seal Phoca largha' Whitehead grampus Grampus griseus' squids were listed and described and Ringed seal Phoca hispida' Pseudorca crassidens' information was presented on their dis­ Ribbon seal Phoca fasciata' Shorttin Globicepha/a macrorhynchus' Bearded seal Erignafhus barbatus Killer whale Orcinus orca tribution, seasonal abundance, com­ Northern Mirounga angustirostris' Harbor porpoise Phocoena phocoena mercial utilization, and position they play Gray whale Eschrichtius robustus Dall's porpoise Phocoenoides dalli' Minke whale Balaenoptera acutorostrata Belukha De/phinapterus /eucas' Bryde whale Ba/aenoptera edeni Physeter macrocephalus' Sei whale Ba/aenoptera borealis Pygmy sperm whale Kogia breviceps' Fin whale Ba/aenoptera physa/us Dwarf sperm whale Kogia simus* Blue whale Ba/aenoptera musculus North Pacific giant Humpback whale Megaptera novaeang/iae bottlenose whale bairdii Right whale Balaena g/acialis Goosebeak whale Ziphius cavirostris' Clifford H. Fiscus is with the National Marine Bowhead' whale Ba/aena mysticetus Ginkgo-tooth whale Mesop/odon ginkgodens Mammal Laboratory, Northwest and Alaska Rough-toothed Archbeak whale Mesop/odon car/hubbsi Fisheries Center, National Marine Fisheries dolphin Steno bredanensis' Bering Sea beaked Service, NOAA, 7600 Sand Point Way, N.E., Bottlenose dolphin Tursiops truncatus' whale Mesop/odon stejnegeri Seattle, WA 98115. Spinner dolphin Stenella longirostris' Densebeak whale Mesop/odon densirostris

February 1982, 44(2; from this research have been combined I I and compiled in a series of reports (North \ 0 I, 42 N Pacific FurSeal Commission, 1962, 1969, I 1971,1975,1980; Lander, 1980; Perez'; g and Perez and Big{ 3, I Two abundantsmall cetaceans (Dall's porpoise and Pacilic white-sided dol­ ·f+ +.', Cape Mendocino phin) and the northern fur seal inhabit ( '.., the same subarctic waters and consume , 400 \ essentially the same species of squids \ \ I (Kajirnura et al., 1980). Antonelis and ,I Fiscus (1980) briefly discussed the food I I and feeding habits of pinnipeds of the , California Current. Some demersal spe­ , cies were found in the stomachs of col­ '"'1) lected specimens, evidencing that these 380 pinnipeds, including the fur seal, and the t~:, two small cetaceans wruch feed primarily , +. + in surface and near surface waters on .. t + + '1:':t.-,_ epipelagic and mesopelagic species, also +~( feed over the continental shelf and de­ ..- + ... I scend to the bottom. l ", 360 ...+ +. \ Several other small cetaceans are PACIFIC OCEAN , ...... " abundant in tropical and subtropical + " • + , , waters and two of them, the saddleback ...... ~ dolphin and the shortlin pilot whale, o range northward into the southern fringe 3, ...... _-- ...... , of the region discussed here. Reilly (1977) summarizes the available information on 340 feeding habits of the pilot whale. He describes live population centers for this species; the northernmost (Californian) stretches from about San Francisco Bay, Calif., to northern Baja California, 1260 W 1250 1240 1230 1220 121 0 1200 Mexico. This population may feed most­ lyon the squid Loligo opalescens. The Figure 1. - Locations off California where northern fur seals were col­ saddleback dolprun is most abundant lected whose stomachs contained Loligo opalescens.

'Perez, M. A. 1979. Preliminary analysis of feed­ ing habits of the northern fur seal in the eastern North Pacibc Ocean and Bering Sea, 1958-74. In H. Kajimura, R. H. Lander, M. A. Perez, A. E. York, and M. A. Bigg, Preliminary analysis from the California Bight southward and United States and Canada during of pelagic fur seal data collected by the United States and Canada during 1958-74. Nat!. Mar. also preys heavily on squids. 1958-74. Collections were made in seven Mammal Lab., Northwest and Alaska Fish. Most information on bathypelagic and areas as follows: 1) California, December Cent., Nat!. Mar. Fish. Serv., NOAA, 7600 Sand benthic squids has been obtained by to June 1958-66; 2) Oregon, January to Point Way, N.E., Seattle, WA 98115. (Submitted to 22d Annual Meeting of the North Pacific examining the stomach contents of sev­ May 1958-65; 3) Wasrungton, December FurSeal Commission.) Unpub!. rep.,p. 167- 232. eral species of toothed whales, primarily to June 1958-74; 4) British Columbia, 'Perez, M. A., and M. A. Bigg. 1980. Interim the sperm whale taken by commercial January to July 1958-72; 5) Gulf of Report on the feeding habits of the northern fur seal. In H. Kajimura,R. H.Lander, M. A. Perez, whalers and from specimens stranded Alaska, February to August 1958-68; 6) A. E. York, and M. A. Bigg, Further analysis of on beaches. Western Alaska, May to October pelagic fur seal data collected by the United States and Canada during 1958-74, Part 2. Nat!' 1958-68; and 7) Bering Sea, May to Mar. Mammal Lab., Northwest and Alaska Fish. Methods November 1958-74. Because the ceph­ Cent., Nat!. Mar. Fish. Serv., NOAA, 7600 Sand alopod fauna of Oregon, British Colum­ Point Way, N.E., Seattle, WA 98115. (Submitted Collection of Specimens to 23d Annual Meeting of the North Pacific bia, and Western Alaska is similar to Fur Seal Commission.) Unpub!. rep., p. 4-172. Fur seals were taken at sea by the adjoining areas, only four of the areas

2 Marine Fisheries Review Figure 2.-Locations off Washingtol where northern fur seals were col lected whose stomachs contair.e( ++ + Loligo opalescent>

48'

+ PACIFIC OCEAN +

47' + + + +...... + are discussed. See the annual reports +

mentioned previously for methods of ~ 46' taking. ~ Oregon Dall's porpoise, PacifIc white-sided 0, dolphin, and several other small ceta­ 128' 1270 125' ceans (saddleback dolphin and killer whale) were also taken at sea during fur seal research cruises (see Kajimura et a!., 1980, for description of taking). Sperm whales taken commercially off central California by commercial whalers from 1956 to 1971 (Rice, 1974 and pers. commun.) were examined at the Rich­ ever, if the head and arms, and tentacles from the shore seaward over the conti­ mond, Calif., stations and stomach con­ are relatively intact, identitication usually nental shelf and slope within its range tents retained for identifIcation at the can be made to . Within the past which extends from Mexico northward National Marine Mammal Laboratory 20 years the numbers of species de­ to British Columbia (Recksiek and Frey, (NMML) in Seattle, Wash. scribed in the family from 1978; Bernard, 1980). Young (1972) Stomach contents of stranded speci­ the North PacifIc Ocean and Bering Sea stated that L. opalescens probably is the mens were collected as they became has more than doubled, and more species most abundant species off available and identifIed in the NMML. will likely be identifIed and described in the California coast. Bernard (1980) Squid remains in stomachs ranged from the future. mentions this squid as being present in relatively intact and undigested whole The degree of digestion of prey items large numbers in the shallow waters off specimensto undigestable hard partssuch is important and should be considered British Columbia. It is locally abundant as the gladius, beaks, and statoliths. when analyzing information on prey seasonally and is a major prey of , Unfortunately many identifIcations must species from marine mammal stomachs. , and most pinnipeds and small be based on beaks alone. The head, base Whole or relatively undigested squids cetaceans throughout its range. of arms and tentacles, and the buccal indicate consumption near the point of During all months in which fur seals mass persist in a stomach longer than do capture; however, beaks alone may rep­ were collected (January through June other body parts, and this fragment can resent accumulations of previous feed­ off California and December through be useful in identifying a specimen at ings and the could have traveled June off Washington), L. opalescens was least to genus. a considerable distance from where the found in the stomachs of fur seals col­ Squid beaks from the stomachs of prey was consumed. Prey identifIed from lected off California and Washington. marine mammals, ush, and birds usually the stomachs of dead, stranded speci­ Most fur seals that had consumed it were can be identifIed to family. In some mens usually provide little information taken over the continental shelf and slope instances, however, species identifIca­ on where prey was consumed. Squids, or a relatively short distance seaward of tions can be made where no near rela­ by family, are discussed below. the slope. It was reported in the stom­ tives occur. For example, in the eastern achs of six fur seals collected well off­ North PacifIc OceanL. opalescens is the shore in the eastern part of the Gulf of only representative ofits family presently Squids Taken as Prey Alaska; however, signifIcant numbers known to occur and Onychoteuthis bor­ Loliginidae have not been found in seals taken north ealijaponicus is the only representative of Hecate Strait, British Columbia. L0­ of its genus. There are many species of Loligo opalescens is the only species cations where fur seals were collected gonatids and they cannot be identifIed of this family identifIed from the north­ whose stomachs contained L. opalescens with certainty from beaks alone; how- eastern North PacifIc Ocean. It is found are shown in Figures 1 (California) and 2

February 1982, 44(2) 3 (Washington) (from Kajimura3 and un­ have been the result of an increase in the of the subarctic water mass except for published records\ It was also found in number of cruises seaward of the conti­ Ommastrephes bartramii which ap­ the stomachs of Dall's porpoise and Pa­ nental slope during those months. lo­ parently ventures across the subarctic cific white-sided collected off cations where fur seals were collected boundary on occasion (Clarke, 1966; California and Washington (Kajimura et whose stomachs contained 0. boreali­ Young, 1972). While we have not identi­ aI., 1980) and in saddleback dolphins japonicus are shown in Figures 3 (Cali­ fied 0. bartramii from the stomachs of collected off California (Fiscus and fornia) and 4 (Washington) (footnotes 3 marine mammals collected in the eastern Niggol,1965). and 4). This species is also taken by North Pacific Ocean, the species occurs Dall's porpoise and Pacific white-sided at least seasonally as reported by Bernard Onychoteuthidae dolphins (Kajimura et aI., 1980). (1980), and we have identified several Onychoteuthis boreabjaponicus is the specimens taken in gillnets by Pacific Ommastrephidae only species of this genus identifIed from Investigation (NMFS Northwest transitional and subarctic waters of the Most members of this family are found and Alaska FIsheries Center) vessels North Pacific Oceans. It ranges over the in tropical and subtropical waters south south of the Aleutian Islands. continental slope and seaward, occurring rarely over the continental shelf from Baja California north to British Columbia in the eastern North Pacific Ocean (Young, 1972). The stomachs of three seals taken off Kodiak Island in the Gulf of Alaska in April and May 1960 con­ tained 0. boreabjaponicus; it is taken in surface gillnets in subarctic waters of 42' N the North PacifIc Ocean south of western Alaska and the Aleutian Islands (foot­ note 4). In the western North Pacific Ocean, this species ranges north to the and waters south of the western Aleutian Islands at least sea­ sonally (Naito et aI., 1977). It is a major 40' prey of pinnipeds and cetaceans that forage in offshore epipelagic and meso­ pelagic zones. It was identified from the stomachs of fur seals collected off California during all months in which fur seals were col­ lected (January-June), and off Washing­ 38' ton from January to June. None were reported off Washington in December but only 78 seals were taken during that month. Occurrences of 0. borealtjapon­ icus increased off Washington in April and May; however, this increase could 36'

1

4 Marine Fisheries Review Figure 4.-Lo­ cations off Wash­ ington where northern fur 490 N seals were col­ + lected whose stomachs con­ + tained Onycho­ leulhis boreali­ + japonicus. 48 0 ~ PACIFIC OCEAN +++ +~ +

47 0 + ~++ ++-+-+-* -++

46 0

1320 W 131 0 1300 1290 1280 1270 1260 1250 1240

Gonatidae upper mesopelagic and epipelagic zones. evident in the fur seal's diet than off Three genera, , , Gonatids were identified in the stom­ California and probably were as impor­ and , comprising about 12 achs of fur seals collected off California tant as 0. borealljaponicus seaward of species, represent this family in subarctic and Washington during all months in the continental shelf. Gonatids were well waters of the North Pacific Ocean and which fur seals were collected (January­ represented in the prey of the Pacific Bering Sea (Young, 1972; Okutani, 1973; June off California, November-June off white-sided dolphin and Dall's porpoise Naitoetal., 1977; Anderson, 1978; Bu­ Washington). Off California, gonatids off Washington (Kajimura et al. 1980). blitz, 1981; and Jefferts, 1981). It is comprised a smaller portion of the fur Pike (1950) and Clarke and Macleod extremely difficult or impossible at pres­ seals' diet than did L. opalescens and (1980) reported Gonatidae from the ent to identify these species from beaks Onychoteuthis borealljaponicus; how­ stomachs of sperm whales taken off alone; therefore, this discussion will be ever, they were well represented as prey British Columbia. Locations where fur restricted to the family level. of the Pacific white-sided dolphin and seals were taken whose stomachs con­ Members of this family are found Dall's porpoise which may regularly feed tained gonatids off California and Wash­ principally from the continental slope at greater depths than do fur seals. Gon­ ington are shown in Figures 5 and 6 seaward and in some inshore deepwater atids were well represented among (footnote 4). localities such as Prince William Sound, squids identified from the stomachs of Gonatids are an important prey of fur Alaska, and southeastern Alaska- British sperm whales taken by commercial seals and the smaller cetaceans in the Columbia. Some individuals may be whalers off central California in the Gulfof Alaska from March through June found at all depths from near the bottom 1960's (footnote 4). Sperm whales ap­ over the outer continental shelf, the along the continental slope to surface parently were feeding near the bottom continental slope, and seaward. loca­ waters at night (Roper and Young, 1975). primarily along the continental slope and tions where seals whose stomachs con­ Pearcy et al. (1977) reported vertical into deeper waters. Those taken by com­ tained gonatids were collected in the migrations of three species of gonatids mercial whalers were collected within Gulf of Alaska are shown in Figure 7 off Oregon (Gonatus pyros, G. onyx, 60-100 n.mi. (110-185 km) of San Fran­ (footnote 4). Gonatids have been identi­ and Gonatopus borealis), and most cisco Bay where the whaling stations fied from the stomachs of northern sea cephalopods captured (mostly juvenile were located. lions and harbor seals collected in the and larval forms) were caught in the Off Washington, gonatids were more northern Gulf of Alaska and Prince

February 1982, 44(2) .') Figure 5.-Locations where northern fur seals were collected off California whose stomachs contained Gonatidae.

+

41 0 N

6 William Sound (Pitcher, 1977; Calkins ; Pitcher and Calkins\ Gonatids, inhabitants of the continen­ tal slope and seaward from the slope, are the only cephalopods identified in 390 the stomachs offur seals collected in the eastern Bering Sea from June through September. Locations in the eastern + Bering Sea where fur seals whose stom­ achs contained gonatids were collected +* are shown in Figure 8 (footnote 4). Go­ 370 natids also are prey of the Dall's porpoise in both the Gulf of Alaska and Bering Sea(Kajimuraet~., 1980). The northern PACIFIC OCEAN sea lion and ribbon seal probably prey on gonatids along the continental slope. Lowry et al. (1977) report squids as 350 important food of ribbon seals. Lowry et al. (1979), summarizing available infor­ mation on feeding by the ice seals, re­ ported squids were consumed by ribbon and spotted seals. We have a few records \, 330 of gonatids from the stomachs of ringed seals taken in the Chukchi Sea (footnote 4). Octopus were reported from the stomachs of bearded and spotted seals. 1270 W 1250 1230 121 0 1190 Johnson et al. (1966) identified octopus in the stomachs of a few ringed and bearded seals taken at Point Hope, Alaska, in the Chukchi Sea. Gonatids also formed a significant part of the food of sperm whales taken by Japanese and Soviet whalers in the Gulf of Alaska pinnipeds that range over the offshore pelagic octopus, Ocythoe tuberculata, and Bering Sea (Table 2). and oceanic waters (as opposed to coast­ was identified in the stomachs of a few al species [Repenning, 1976]) are oppor­ fur seals taken off southern California. Results and Discussion tunistic feeders, perhaps attracted to Four of these squids, Abraliopsis spp., Some marine mammals are relatively certain localities or regions by seasonally Octopoteuthis spp., Moroteuthis ro­ restricted in their diet, regularly feeding abundant fishes and squids. Over the busta, and Chiroteuthis spp., and the on only a few types of prey. However, continental shelf, fishes comprise most pelagic octopus Ocythoe were taken in many of the toothed whales and those of their diet, but along the continental relatively small numbers and did not con­ slope (sometimes characterized by nutri­ tribute significantly to the fur seal's food ent-rich upwellings) fish and squids are in this region. ~Calkins, Alaska Department of Fish and equally important prey. In oceanic The northern fur seal, as well as many Game, 333 Raspberry Road, Anchorage, AI< waters squids are usually the most im­ other marine mammals, is migratory. 99502. Pers. Commun. portant prey. Thus, information gathered during pe­ 7Pitcher, D. W., and D. G. Calkins. 1981. Biolo~ of the harbor seal, Phoca vitulina richardsi, In At least nine squids have been identi­ lagic fur seal studies, which was keyed to the Gulf of Alaska. Outer Continental Shelf fied from the stomachs of fur seals col­ periods ofseasonal abundance in certain Environmental Assessment Program Final Re­ port. Juneau Project Office, P.O. Box 1808, lected in the eastern North Pacific Ocean localities, does not provide information Juneau, AI< 99802. and Bering Sea (Table 3) and a small on a year-round basis. For example, no

6 Marine Fisheries Review Figure 6.-Locations where northern fur seals were collected off Washington whose stomachs contained Gonatidae.

300 N

1500 1400

Figure 7. - Locations where northern fur seals were collected in the Gulf of Alaska whose stomachs contained Gonatidae.

February 1982. 44(2} 7 "I '-,

58° N

BERING SEA

57°

56°

+ ++ 55°

,- /---_.... I 54°

53°

173° W 171' 169° 167" 165° 163°

Figure 8. - Locations where northern fur seals were collected in the Bering Sea whose stomachs contained Gonatidae.

Table 2.-Squids identified from the stomachs of sperm whales from the Bering Sea and Gulf of Alaska"'. Family and Aleutian area Gulf of Family and Aleutian area Gulfor species name and Bering Sea Alaska species name and Bering Sea Alaska Family Gonatidae Family Chiroteuthidae 2 Chiroteuthis veranyi 2 Berryteuthis (Gonatus) magister 1,2 2 Family Mastigoteuthidae Gonatus fabricii var separata 2 Mastigoteuthis sp. Gonatopsis makko 1 Family Cranchidae Family Octopoteuthidae Taonius pavo 1,2 Octopoteuthis longipetra 2 Galiiteuthis armata 1,2 Cristalloteuthis behringiana 2 Family Onychoteuthidae Moroteuthis robustus 1,2 Family Ommastrephidae Onychoteuthis (banksii) borealijaponicus 2 Todarodes (sloaneipacificus) pacificus 2 Ommastrephes bartramii 2 Family Histioteuthidae Stigmateuthis 1 Order Octopoda Meleagroteuthis separata 2 Octopus sp, 2

Family Architeuthidae Architeuthis japonica 2 '1; data from Okutani and Nemoto (1964). '2; data from Kodolov (1970).

8 Marine Fisheries Review Table 3. -Cephalopods identified in the stomach contents o.f northern fur seals. Callorhinus ursinus. collected during pelagic fur seal investigations. 1958-74. Loca- Loligo Abrali- Octopoteu- Other Gonatus Berryteu- Gonatopsis Onychoteuthis Moroteuthis Chiroteu- Unident. Year tion' opa/eseens apsis sp. this sp. gonatids spp. this spp. borealis borea/iiaponieus robusta this sp. squid 1958 CA x OR x WA AK(SE. GA.WA. BS)

1959 CA x OR x WA x

1960 AK

1961 CA x OR x WA x BC x

1962 AKIWA. UP. BS) x

1963 AK(BS)

1964 CA x OR x WA x AKIBS) x

1965 CA WA

1966 CA x

1967 WA

1968 WA AKIGA, BS.WA)

1969 WA

1970 WA

1971 WA

1972 WA

1973 AKIBS)

1974 AK 18S) 'SE = southeastern Alaska, GA - Gulf of Alaska. WA - western Alaska, UP - Unimak Pass. BS - Bering Sea. CA - California. OR - Oregon. WA - Washin9ton. AK = Alaska. BC = British Columbia.

collections were made off California ceans, and sperm whales with those to British Columbia; Onychoteuthis bor­ from July to December, off Washington obtained from traditional methods of ealijaponicus may also be sufficiently from July to November, in the Gulf of direct collecting and sampling, we can abundant in this area and seasonally into Alaska from August to February, or in compile a useful base of information for Alaskan waters. At least two (Berryteu­ the Bering Sea from November to April, cephalopod stock assessment and fIshery this magister and Gonatopsis borealis) times when few or no fur seals are pres­ development. of the 12 or more species of gonatid ent. Information is needed on the distri­ Based on frequency of occurrence in squids found in the area are large and bution and abundance of squids for given the stomach contents of marine mam­ probably abundant enough for com­ localities for the entire year to determine mals, the following squids probably are mercial utilization from Washington ifpotentially valuable squids are available sufficiently abundant in some parts of north to the Bering Sea. Populations of in signifIcant numbers to support com­ the North PacifIc Ocean and Bering Sea several genera (Symplectoteuthis, Do­ mercial fIsheries. to support commercial fIsheries: L. opal­ sidicus, and Ommastrephes) within the By combining data available on ceph­ escens, presently fIshed off central and family Ommastrephidae are probably alopod occurrences in the stomachs of southern California, could probably sup­ abundant enough to support commercial fur seals, other pinnipeds, small ceta- port fIsheries from northern California fIsheries from subtropical waters north

February 1982, 44(2) 9 to southern California. One member of biology of cephalopods, p. 89-126. Acad. Fisheries biological production in the subarc­ Press Lond., 615 p. tic Pacitic region, special volume, p. 321-337. the genus Ommastrephes, 0. bartramii, _----:-_ ,and N. Macleod. 1980. Cephalo­ Res. Inst. North Pac. Fish., Fac. Fish. Hok­ occurs north to the subtropical-subarctic pod remains from sperm whales caught off kaido Univ., Hakodate, Japan. boundary region and may be present western Canada. Mar. BioI. (Berl.) 59:241­ North Pacitic Fur Seal CommIssion. 1962. North 246. Pacitic Fur Seal Commission report on inves­ further north seasonally in sufficient Fiscus, C. H. 1979. Interactions of marine mam­ tigations from 1958 to 1961, North Pac. Fur numbers to support a fishery. malsand Pacitic hake. Mar. Fish. Rev.4I(lO): Seal Comm., Wash., D.C., 183 p. Octopus spp. which are found on the 1-9. _-:---:-_.1969. North Pacitic Fur Seal Com­ _--,---__ , and K. Niggol. 1965. Observations mission report on investigations from 1964 to continental shelf throughout the region of cetaceans off California, Oregon, and 1966, North Pac. Fur Seal Comm., Wash., form part of the catch in the bottom Washington. U.S. Fish. Wildl. Serv., Spec. D.C., 161 p. Sci. Rep. Fish. 498, 27 p. _-:---,----_ . 1971. North Pacitic Fur Seal Com­ trawl fishery and are regularly taken by Jefferts, K. 1981. Zoogeography and systematics mission report on investigations in 1962-1963. harbor seals, northern and California sea of cephalopods of the Northeastern Pacitic North Pac. FurSeal Comm., Wash., D.C., 96 p. lions, several of the ice- inhabiting seals, Ocean. M.S. Thesis, Oregon State Univ., _-:---:-_.1975. North Pacitic Fur Seal Com­ Corvallis. n.p. mission report on investigations from 1967 and sperm whales. Johnson, M. L., C. H. Fiscus, B. T. Ostenson, through 1972, North Pac. Fur Seal Comm., Information that can be obtained from and M. L. Barbour. 1966. Marine mammals. Wash.. D.C., 212 p. examinations of marine mammal stom­ In N. 1. Wilimovsky and 1. N. Wolfe (editors), _-:---,----_ . 1980. North Pacitic Fur Seal Com­ Environment of the Cape Thompson region, mission report on investigations from 1973­ ach contents and used for developing Alaska. p. 877-924. U.S. Atomic Energy 1976, North Pac. Fur Seal Comm., Wash.. commercial fisheries include: Identifi­ Comm., Div. Tech. Inf. (Avail. U.S. Dep. D.C., 197 p. Commer., Natl. Tech. Inf. Serv., Springtield, Okutani. T. 1973. Guide and keys to squid in cation of prey, date and general location Va., as PNE-481.) Japan. Bull. Tokai Reg. Fish. Res. Lab. 74: where the marine mammals are cap­ Kajimura, H., C. H. Fiscus, and R. K. Stroud. 83-111. tured, and the size, sex, and reproductive 1980. Food of the PaciJic white-sided dolphin, _:---,-----,-- ,and T. Nemoto. 1964. Squids as the Lagenorhynehus obliquidens. Dall's porpoise, food of sperm whales in the Bering Sea and condition of prey. Clarke (1977) men­ Phoeoenoides dalli. and northern fur seal, Alaskan Gulf. Sci. Rep. Whales Res. Inst. tioned that marine mammals often are Callorhinus ursinus. off California and Wash­ Tokyo 18:111-122. more efficient in capturing large squids ington, with appendices on size and food of Pearcy, W. G., E. E. Krygier, R. Mesecar, and Dall's porpoise from Alaskan waters. U.S. F. Ramsey. 1977. Vertical distribution and than are nets; thus, information on size Dep. Commer., NOAA Tech. Memo. NMFS migration of oceanic micronekton off Ore­ (maturity) of a species in a given location F/NWC-2, 30 p. gon. Deep Sea Res. 24:223-245. Kodolov, L. S. 1970. O. kalmarakh v. Bering­ Pike, G. C. 1950. Stomach contents of whales perhaps can best be obtained through ovom more (Squids of the Bering Sea). Tr. caught off the coast of British Columbia. Fish. the identification of stomach contents. Vses. Nauchno- issled. Inst. Morsk. Rybn. Res. Board Can., Prog. Rep. Pac. Coast Sta­ Khoz. Okeanogr. 72:162-165. [In Russ.1 tions 83:27-28. (Transl. by Israel Program Sci. Transl., 1972, Pitcher, K. W. 1977. Population productivity Acknowledgments p. 157-160.) In P. A. Moiseev (editor), Soviet and food habits of harbor seals in the Prince tisheries investigations in the northeastern William Sound-Copper River delta area, The author thanks Hiroshi Kajimura, Pacific, Part 5. Available from Natl. Tech. Alaska. Final rep., MMC-75/03, Contract NMFS National Marine Mammal labo­ lnf. Serv., Springtield, Va., as TT 71-50127. MM5ACOII, 36 p. Mar. Mammal Comm., ratory, for his assistance in assembling Lander, R. H. (editor). 1980. Summary of north­ Wash., D.C. ern fur seal data and collection procedures, Recksiek, C. w., and H. W. Frey. 1978. 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10 Marine Fisheries Review