The Gray , robustus

DALE W. RICE, ALLEN A. WOLMAN, and HOWARD W. BRAHAM

Introduction eastern stock which breeds along the 1970; Mitchell and Mead I), but has west coast of North America, and the been extinct there for several cen­ The gray whale, Eschrichtius "Korean" or western stock which ap­ turies. robustus (LilIjeborg, 1861), is readily parently breeds off the coast of recognized by its mottled gray color eastern Asia (Rice and Wolman, Distribution and Migration and lack of a dorsal . Instead of 1971). Both stocks were severely Eastern North Pacific this fin, it has a low hump, followed depleted by the early 1900's. Under by a series of 10 or 12 knobs along the legal protection, the eastern stock has Most of the stock spends dorsal ridge of the tail stock; these are recovered substantially- one of the the summer feeding, mostly in the easily seen when the arches to few stocks of great to do so. northern Bering and southern Chukchi dive. The adult gray whale is 36-50 The western stock has not recovered. Seas (Pike, 1962; Rice and Wolman, feet long and weighs between 16 and The gray whale formerly occurred in 45 tons. the North Atlantic (van Deinse and The gray whale is currently con­ Junge, 1937; Cederlund, 1939; Fraser, 'Mitchell, E. D., and 1. G. Mead. fined to the North Pacific Ocean (Fig. 1977. History of the gray whale in the Atlantic Ocean. (Abstr.) In Proceedings of the 2nd 1). Because it uses coastal habitats ex­ Conference on the Biology of Marine Mam­ tensively, the gray whale was especial­ The authors are with the National Marine mals, , California, 12-15 December ly vulnerable to shore-based Laboratory, Northwest and 1977, p. II. (Available from first author, Arctic Fisheries Center, National Marine Fisheries Biological Station, Department of Fisheries and operations. Two stocks occur in the Service, NOAA, 7600 Sand Point Way N.E., Oceans, 555 SI. Pierre Blvd., Ste. Anne de North Pacific: The "California" or Bin C15700, Seattle, WA 98115. Bellevue, Quebec, H9X 3R4, Canada).

120' E 150' E 180' 150'W 120'W 90'W 60'W 30'W O'

Figure I. - Geographic distribution of the gray whale. Simple hatching indicates the summer feeding grounds. Small dots indicate the migration routes. Stippling indicates the winter grounds. In the Atlantic, the gray whale has been extinct for at least several hundred years; early historical records are indicated by large dots, subfossil finds by triangles. Perhaps extinct is the population that formerly spent the winter in southern Japan (large dot).

46(4),1984 7 1971; Bogoslovskaya et al., 1981). An '¢.~~-- - unknown number of individuals sum­ ~.~ __.---- '1 mer along the west coast of North -~..:::: America in apparently isolated loca­ tions south of Alaska from , Canada, as far south as , (Patten and Samaras, 1977; Sprague et al., 1978). In the , sightings have been made of small groups as far east as long. 130 0 W during August (Rugh and Fraker, 1981); in the , gray whales were found along the Siberian coast as far west as 174°08'E in late September (Marquette et. al., 1982). A new~orn gray whale calf in Laguna Ojo de Liebre, Baja California, Mexico. In October and November, the Each dimple on the snout and lower lip marks the site of a hair. Photo by D. W. RIce. stock begins leaving the , exiting the through Unimak Pass, Alaska, mainly in November and December (Rugh and Braham, 1979; Braham, 1984; Rugh, 1984). The The major calving areas are Laguna Unimak Pass (Braham et aI., 1977; whales migrate near shore along the (with 9 percent of the Braham, 1984). This migration is coast of North America from Alaska calves), Laguna Ojo de Liebre (53 per­ completely coastal, at least to the east all the way to central California (92 cent), Laguna San Ignacio (II central Bering Sea (Nunivak Island). percent pass within 1.4 km of Cape percent), and Estero Soledad (12 per­ Most in Alaska travel within Sarichef, Unimak Pass, and 94 percent cent). Minor calving areas (each with I km of the coast, especially in the pass within 1.6 km of the Monterey­ < 6 percent) are San Juanico Bight, southeastern Bering Sea, and at least Point Sur area of central California). Bahia Magdalena, Bahia Almejas, and some apparently feed during migra­ After passing Point Conception, Bahia Santa Marina (Rice et al., 1981). tion (Braham, 1984). During the Calif., the majority take a more direct Calving rarely occurs during the south­ northward migration, the sequence is offshore route across the southern bound migration north of Baja as follows: Newly pregnant females, California Bight to northern Baja California (Rice and Wolman, 1971; followed by anestrous females, adult California. Southbound migrating Sund, 1975). A few calves are also males, and immature males and gray whales swim at about 7.7 born on the eastern side of the Gulf of females; cows with calves are the last kmlhour, and thus travel about 185 California at Yavaros, , and animals to leave the , and km per day (pike, 1962). Bahia Reforma, , Mexico most migrate after the other whales. Migrating gray whales are temporal­ (Gilmore, 1960). Contrary to many The peak of the migration passes ly segregated according to sex, age, published statements, there is no Point Piedras Blancas, Calif., about I and reproductive status (Rice and evidence that San Diego Bay, Calif., May (Poole3). was ever a calving area (Henderson, Wolman, 1971). During the southward Western North Pacific migration, the sequence of passage off 1972). Recent studies have revealed California is as follows: Females in late that the vast majority of gray whales in The Korean stock formerly oc­ pregnancy, followed by females that Baja California (other than cows with cupied the northern in have recently ovulated, adult males, calves) spend the winter outside the the summer, as far north as Penzhin­ immature females, and then immature lagoons in Bahia Sebastian Viscaino skaya Bay, and south to Akademii 2 males. The earliest southbound and Bahia de Ballenas (Rice et al. ). and Sakhalinskiy Gulfs on the west migrants (mostly late-pregnant The northbound migration begins and the Kikhchik River on the east. females) usually travel singly, whereas in mid-February, and by April whales Southbound whales migrated along later migrants usually are in pods of begin showing up in the southern Ber­ the coast of eastern Asia to winter two or more. The mean pod size ing Sea, which they enter through through Unimak Pass is about two. 'Poole, M. M. 1981. The northward migra­ This stock winters mainly along the tion of the California gray whale, Eschrichtius 'Rice, D. W., A. A. Wolman, and D. E. rabustus, off the central California coast. west coast of Baja California. The Withrow. 1984. Distribution and numbers (Abstr.j In Proceedings of the 4th Biennial Con­ pregnant females assemble in certain of gray whales on their Baja California winter ference on the Biology of Marine , shallow, nearly landlocked lagoons grounds. Unpub!. manuscr. Nat!. Mar. December 14-18, 1981, San Franc., Calif., p. Mammal Lab., Nat!' Mar. Fish. Serv., NOAA, 96. (Available from author, Biology Depart­ and bays where the calves are born 7600 Sand Point Way N.E., Bin CI5700, Seat­ ment, Sonoma State University, Rohnert Park, from early January to mid-February. tle, WA 98115. CA 94928. 8 Marine Fisheries Review Circular mud plume pattern produced by a feeding gray whale in the northern Bering Sea. This behavior is believed to be associated with the whale returning to the location it was just at to resume feeding. Photo by H. W. Braham.

calving grounds off the south coast of waters of the Chukchi and Bering , passing from late Seas, they feed primarily on benthic November to late January. Until the gammaridean amphipods. Forty­ turn of this century, another migra­ three species have been identified tion route led down the eastern side of from stomachs, but, depending on Japan to winter grounds in the Seto area, one of seven species is usually Inland Sea, Japan (Omura, 1974). dominant (Pontoporeia femorata, P. Nishiwaki and Kasuya (1970) and affinis, Anonyx nugax, Ampelisca Bowen (1974) hypothesized that three macrocephala, A. eschrichti, recent records of gray whales in Japan Nototropis brueggeni, or N. ekmam). involved vagrants of the California In some areas polychaete worms are stock rather than being Korean stock their main food. Incidentally ingested survivors. It is likely that any rem­ benthos include gastropods, asci­ nants of the Korean stock are in such dians, bivalves, priapulids, decapod low numbers (Brownell, 1977) as to be , isopods, sipunculids, below a critical population size suffi­ hydrozoans, anthozoans, cumaceans, cient for recovery. This stock holothurians, sponges, and fish (Am­ therefore may be almost extinct. modytes sp.) (Zimushko and Len­ skaya, 1970; Bogoslovskaya et aI., Life History and Ecology 1981). Gray whales may play an im­ portant role in the rate of turnover of Aerial view of a feeding gray whale Feeding surfacing in the northern Bering the epibenthos on their summer Sea, near St. Lawrence Island, Gray whales are predominantly feeding grounds (Nerini and Oliver, Alaska. Note the heart-shaped blow bottom feeders that apparently ingest 1983; Nerini, 1984). and the trailing mud plume caused their food by suction (Ray and Little if any food is consumed dur­ when the whale expells water and debris out the side of its mouth Schevill, 1974); only rarely do they ing the southbound migration off the when surfacing after feeding on feed in midwater or at the surface. On U.S. continental coast, although rare­ organisms along the bottom of the their summer grounds in the shallow ly small quantities of decapod nauplii sea. Photo by H. W. Braham.

46(4),1984 9 (Pachyche/es rudis and ?Fabia sp.) are season or 0.46 per year. The period of Anisakis simp/ex (0.3); and two eaten (Rice and Wolman, 1971). gestation is about 13.5 months; fetal acanthocephalans, Corynosoma sp. There are reports that they do feed to growth accelerates during the last half (5.7) and Bo/bosoma sp. (0.3). Ob­ some extent just before (Sund, 1975) of pregnancy and decelerates just vious pathogenic effects are produced and while on their winter grounds off before birth (Rice, 1983). During only by the liver fluke Lecithodesmus Baja California (Swartz and Jones, southward migration, late pregnant goliath, but it is not known whether 1982) although the frequency of this females (exclusive of their conceptus) this ever causes mortality. behavior is unknown. In the interval average 25-30 percent heavier than The , Orcinus orca, ap­ between their southward and north­ other adult females. Most births occur pears to be the only predator on gray ward migration past San Francisco, within a period of 5-6 weeks, with a whales. Evidence from necropsy of 39 the whales without calves lose from peak occurring about 27 January; ex­ gray whales that stranded on St. 0.21 percent to 0.37 percent of their treme recorded dates are 26 December Lawrence Island indicated that 16 had body weight per day. This weight to 1 March (Swartz and Jones, 1983). been killed by killer whales (Fay et aI., reduction is sufficient to account for Lactation lasts an average of about 1978). The mortality rate from killer the estimated energy expenditure dur­ 7 months, ending in August. Females whale attacks is unknown. However, ing the winter. Blubber thickness and are usually in anestrus from August to the frequency of tooth scars on gray oil yield also decrease during winter. November or December. However, whale carcasses indicates that killer Apparent feeding has been observed females that fail to ovulate or con­ whale attacks are often unsuccessful. during the northbound migration ceive during the winter are probably Moderate numbers of gray whale beginning in southeastern Alaska in anestrus for the following 12 calves strand in and near the nursery (Braham, 1984), but again the fre­ months. lagoons (Swartz and Jones, 1983). A quency and quantitative evidence Males attain puberty at an few adults strand every year associated with energy expenditure estimated mean age of 8 years (range, throughout the range, but the number for this is unknown. 5-11 years) and a mean body length of seems low compared with the size of about ILl m. The average weight of the population. Rates of mortality Reproduction the testes of adult males during due to stranding cannot be calculated. Females attain puberty at an southward migration in December Total annual mortality estimates estimated mean age of 8 years (range, and January is 38 kg, and the mean for animals older than 8 years, 5-11 years) and a mean body length of diameter of the seminiferous tubules calculated from the age composition about 11.7 m (see Rice and Wolman is I77/-tm. During northward migra­ as determined by ear-plug readings, (1971) for additional details on tion in February and March, mean were 0.095 for females and 0.081 for reproduction). testes weight and tubule diameter are males; a similar estimate for sexually Female gray whales normally come 22 kg and 148 fllll, respectively. mature females, based on ovarian into estrus biennially in late From July through October, the testes corpora counts, was 0.082 (Rice and November and early December. Most average 23 kg. These differences sug­ Wolman, 1971). These estimates are individuals ovulate only once each gest a marked seasonal sexual cycle in probably biased upwards because the season, although whales failing to the male, with a peak of sperma­ population was increasing during the conceive after their first ovulation togenetic activity in late autumn or 1950's and 1960's when the data were may experience a second estrous cycle early winter. collected. Reilly (1981) estimated the the same season. Multiple ovulations adult natural mortality rate at 0.056 are extremely rare. Mean ovulation Natural Mortality and the juvenile mortality rate at rates are 1.20 per breeding season for No infectious diseases have been 0.132 during that period. The sex nulliparous females and 0.96 per reported in gray whales. Epizoites of ratio is essentially equal throughout breeding season (0.52 per year) for gray whaies include the following life. parous females. There is little (percentage of occurrence in paren­ evidence of postpartum ovulation or theses) (Rice and Wolman, 1971): The Exploitation and Population Size of ovulation at any other time of the barnacle Crypto/epas rhachianecti History of Exploitation year. However, increase in follicle size (100) and the cyamids Cyamus scam­ following stillbirth or early loss of the moni (99.7), C. ceti (99.4), and C. Eskimos living on the shores of the calf suggests that females might kess/eri (98.1). Endoparasites include northern Bering Sea and the Chukchi ovulate following such an event. the trematodes Lecithodesmus goliath Sea have hunted whales for perhaps Most conceptions occur within a (0.6), Ogmogaster penta/ineatus several thousand years. In Alaska, the 3-week period during southward (>22), and O. antarcticus (33); two catch is mostly of bowhead whales, migration, with a peak about 5 apparently undescribed species of the Ba/aena mysticetus, with very few December; a few occur as late as cestode Priapocepha/us, one in the gray whales taken, usually less than January on the winter grounds. The small intestine (30) and the other in one per year (Marquette and Braham, pregnancy rate is 0.86 per breeding the large intestine (0.3); the nematode 1982). However, on the Chukotka

10 Marine Fisheries Review coast of the U.S.S.R. the catch has America also may have taken a few Between 1959 and 1969, 316 gray been almost entirely gray whales. gray whales (Mitchell, 1979). whales were killed under Special Since 1969 gray whales have been From 1846 until about 1900, Scientific Permits off California. taken by the Soviet Government for American whalers exploited gray From 1966 to 1969 the combined the Chukchi Eskimos using one whales mostly on their wintering scientific and U.S.S.R. catches modern-style catcher boat (Ivashin grounds, but also took a few in north­ averaged 221 per year. and Mineev, 1981). The total ern waters during the summer. On the aboriginal catch since 1967 has aver­ basis of available historical records, Current and aged about 165 gray whales per year Henderson (1972) estimated that the Initial Stock Sizes (Table 1). The current catch limit set total catch from 1846 to 1874 was Scammon (1874) estimated that the by the International Whaling Com­ about 8,100. During the peak of this California gray whale population was mission (IWC) is 179 per year. fishery from 1855 to 1865, the annual probably not over 30,000 in Several Indian tribes on Vancouver catch averaged 474 whales. Catches 1853-1856, and that by 1874 the Island and in the State of during the three winter seasons from number did not exceed 8,000 or traditionally hunted gray whales, but 1883-84 to 1885-86 were 58, 68, and 10,000. After a careful analysis of the have not done so since 1928. Indians 41, respectively (Townsend, 1887). historical data, however, Henderson farther north along the coast of North Modern-style whaling began on the (1972) concluded that the population west coast of North America in 1905. did not exceed 15,000-20,000 prior to A few gray whales were taken in the the initiation of commercial exploita­ winter off Baja California and tion in 1846. California, mostly between 1925 and In 1885-86, Townsend (1887) Table 1.-Catches of California gray whales by aboriginal whaling, 1948·82. 1929. Factory ships took an average estimated that only 160 gray whales

Year U.S.S.R.' Alaska' Total of 48 gray whales per year in the Ber­ migrated south past San Simeon, ing Sea from 1933 to 1946 (Table 2), Calif. Andrews (1914) wrote that "For 1948 19 19 1949 26 26 after which commercial whaling for over 20 years [preceding 1910] the 1950 10 11 gray whales was banned by the Inter­ species had been lost to science and 1951 12 13 1952 42 44 national Convention for the Regula­ naturalists believe it to be extinct." tion of Whaling. Howell and Huey (1930) said it was 1953 37 1 38 1954 38 3 39 1955 59 59 1956 121 122 1957 95 96

1958 145 3 148 Table 2.-Catches of California gray whales by modern·style whaling, 1913·47' 1959 187 6 193 1960 156 156 Baja Bering and 1961 207 208 Year California California Washington Alaska' Chukchi Seas' Total 1962 147 147

1963 178 1 179 1913 1 1964 188 2 190 1914 19 19 1965 175 1 176 1966 194 194 1920 1967 125 125 1921 1922 1968 135 135 1969 139 140 1924 1 1970 146 151 1925 100 33 133 1971 150 153 1926 41 1 42 1972 181 182 1927 29 3 32 1928 9 1 12 1973 173 173 1929 2 2 1974 181 184 1975 171 171 1933 2 4 1976 163 163 1934 54 54 1977 186 187 1935 34 34 1936 102 102 1978 182 2 184 1937 14 14 1979 178 4 182 1938 54 54 1980 179 3 182 1939 29 29 1940 1981 135 0 135 105 105 1982 160 4 184 1941 57 57 1942 101 101 1943 99 99 'Data from Ivashin and Mineev (1978) and with addition of figures for 1978·82 from un· 1945 30 30 published data of the AII·Union Research In· stitute of Marine Fisheries and Ocean· 1947 ography (VNIRO), Moscow. 'Data from Marquette and Braham (1982): and unpublished data of the National Marine 'Data summarized from Rice and Wolman (1971), except that the figures for 1943, 1946, Mammal Laboratory. Actual values may be and 1947 have been changed to agree with those in Kleinenberg and Makarova (1955). low because the taking of gray whales is ' (shore stations at Port Armstrong and Port Hobron). often not reported. 'Peiagic whaling.

46(4),1984 11 18 present is shown in Figure 2. Results 16 of these assessments suggest that the population has not increased to the "0 14 ~ level that it would reach if there were 12 0" -5 no current exploitation. The popula­ c: 10 tion has recovered, however, to the ~ level (which was presumably stable) ~ c: that it was at before commercial whal­ '2 '" ing began in the mid-19th century. 0- 0" Reilly (1981) estimated the max­ "- imum sustainable yield (MSY) as 480 gray whales per year; however, in a 1~00 7 1810 1820 1830 1840 1850 1860 1870 1880 1890 1900 1910 1920 1930 19'10 1950 1960 19 0 1980 recent revision, he gave a new Year estimate of 320 per year, which would Figure 2. - Gray whale population t~ajectory based on an. e~timated occur at a population size of 11,380 maximum population size of 24,000 pnor to 1800 and an abongmal take whales 5 • His revised model also through that time of 600 whales per year. Peaks and vall~ys relate to predicted that, under an annual take periods of heavy exploitation and recovery froll? commercIal whalIng. of 180 whales per year, the population This modeling best fits the current populatIon census of about 16,000±3,000 (from Reilly, 1981). would reach 19,000 by the year 2150 and continue rising slowly thereafter, achieving stability at a level of 19,620.

Management One potential threat to the Califor­ " .. doubtful whether more than a tion size in 1979-80 off California was nia gray whale population may be in­ few dozen individuals survive." estimated at 15,647 (95 percent con­ creasing industrial development and However, K. W. Kenyon4 says that he fidence interval of 13,450-19,201); the vessel traffic in the calving lagoons commonly observed gray whales rate of annual net increase (less the and in other vital habitats along the migrating past La Jolla, Calif., during 1.2 percent harvest mortality) during migration route and on the feeding the 1930's. the preceding 13 years was 2.5 per­ grounds. In the recent past, con­ Systematic shore counts of the cent, with a standard error of 0.96 siderable harassment has been caused southward migration were initiated at (Reilly et aI., 1980; Reilly, 1981; Reilly by commercial cruise boats which San Diego, Calif., in 1952-53, and et aI., 1983). This indicates that the take people into the calving lagoons to continued intermittently until 1976-77 eastern North Pacific population has see the whales and by small pleasure (Gilmore, 1960; Rice, 1961). These recovered to, or now exceeds, its size craft brought overland down the new counts indicated a steadily increasing prior to commercial whaling. Baja California highway. Harassment population until 1959-60. Computer simulation models ofthe now may be under better control than From 1967-68 to 1973-74, a shore gray whale population size from 1800 in the past. Under existing U.S. laws, count was made every winter at through 1980 were run by Reilly regulation and enforcement are being Yankee Point near Monterey, Calif. (1981). Various forms of dynamic defined and steps taken to control where 90 percent of the whales pass response of vital parameters to vessels that interfere with gray whales within 2 miles of shore and boat traf­ population density were used, as well on their migration path. Between fic is at a minimum. From 1974-75 to as various carrying capacity levels, 1972 and 1979, the Mexican Govern­ 1979-80 the count was made at and various levels of prehistoric ment designated three of the five ma­ Granite Canyon, 4 miles south of aboriginal removal rate. The model jor calving lagoons in Baja California Yankee Point. A census of the that produced a population trajectory as gray whale refuges (Reeves, 1977; population leaving the summer in best agreement with the census data Swartz and Jones, 1982). These are feeding grounds was made from 1977 and historical evidence indicated that the lagoons visited by most of the to 1979 at Unimak Pass. the carrying capacity (or maximum U.S. tour boats and private tourists. The 1977 estimate of the popula­ population size historically) may have The number of vessels allowed in the tion leaving the Bering Sea was 15,099 been 24,000, and that the population lagoons at anyone time is limited, ± 2,341 (Rugh and Braham, 1979), had been reduced to below 12,000 by and for the 3 years 1977-79 Rugh the year 1800 as a result of aboriginal (1984) estimates 17,000. The popula- takes which may have averaged 600 'Reilly, S. B. 1984. Future trends in gray whales per year (Reilly, 1981). The whale population size. Unpubl. manu~cr. population trajectory generated by Southwest Fisheries Center, Natl. Mar. FIsh. 4K. W. Kenyon, 11990 Lakeside Place N.E., Serv., NOAA, P.O. Box 271, La Jolla, CA Seattle, WA 98125. Pers. commun. this model for the period 1800 to the 92038.

12 Marine Fisheries Review and entry into certain areas is forbid­ den. Thousands of tourists and a multimillion-dollar industry result from these activities. Oil and gas exploration is con- templated or under way on the con­ tinental shelf from California to the Beaufort Sea, throughout the migra­ tion range of this species. Annually, the gray whale population migrates by or through at least eight oil lease areas in U.S. waters alone. On the winter calving grounds, exploratory areas in­ clude sites within and adjacent to present calving and rearing areas, such as the offshore waters of Viz­ caino Bay, where seismic exploration for gas deposits took place during spring 1981. The effects of oil pollu­ A gray whale, raising its barnacle encrusted head above the surface of Laguna tion on the benthic organisms on Ojo de Liebre in Baja California, reveals its paired blowholes. Photo by D. W. which these whales feed are unknown. Rice. Little is known about what effects, if any, other activities associated with coastal development might have; cer­ tain man-made sounds cause migrat­ virons remains an important conser­ Estudios Economicos y Sociales del Tercer ing whales to deviate from their Mundo, Mexico. [In Span.] vation measure. Fay, F. H., R. A. Dieterich, L. M. Shults, and course (Tyack et al. 6). B. P. Kelly. 1978. Morbidity and mortali­ Past industrial activities have ty of marine mammals. In Environmental shown some impacts. For example, in Literature Cited assessment of the Alaskan continental shelf., Annual Reports 1:39-79. U.S. Dep. Com­ the calving of Guerrero mer., Natl. Oceanic Atmos. Admin., En­ Negro, daily dredging and vessel traf­ Andrews, R. C. 1914. Monographs of the viron. Res. Lab., Boulder, Colo. fic caused the whales to abandon the Pacific . I. The California gray Fraser, F. C. 1970. An early 17th century whale (Rhachianecles glaucus Cope). Mem. record of the California gray whale in Icelan­ area from 1957 to 1967. The whales Am. Mus. Nat. Hist. (New Ser.), 1:227-287. dic waters. Invest. Cetacea 2: 13-20. did not return until 6 years after such Bogoslovskaya, L. S., L. M. Votrogov, and Gard, G. 1974. Aerial census of gray whales operations had ceased (Gard, 1974; T. N. Semenova. 1981. Feeding habits of in Baja California lagoons, 1970 and 1973, the gray whale off Chukotka. Rep. Int. with notes on behavior, mortality, and con­ Bryant and Lafferty, 1980). Current Whaling Comm. 31:507-510. servation. Calif. Fish Game 60:132-143. exploitation of phosphorus near the Bowen, S. L. 1974. Probable extinction of Gilmore, R. M. 1960. A census of the Cali­ calving lagoon of Magdalena Bay in the Korean stock of gray whale (Eschrichlius fornia gray whale. U.S. Fish Wildl. Serv., robuslus). J. Mammal. 55:208-209. Spec. Sci. Rep. Fish. 342, 30 p. southern Baja California may be Braham, H. W. 1984. Migration and Henderson, D. A. 1972. Men and whales at cause for concern (Cordoba, 1981). feeding of gray whales (Eschrichlius Scammon's Lagoon. Dawson's Book Shop, robuslus) in Alaska. In M. L. Jones, S. L. Los Angeles, 313 p. Because of the scarcity of suitable Swartz, and J. S. Leatherwood (editors), The Howell, A. B., and L. M. Huey. 1930. Food isolated calving and nursery areas for gray whale, p. 249-266. Acad. Press, N. Y. of the gray and other whales. J. Mammal. gray whales, and the whales' specializ­ --::--:--c--' C. Fiscus, and D. Rugh. 1977. II :321-322. Marine mammals of the Bering and southern Ivashin, M. V., and V. N. Mineev. 1978. 0 ed feeding habits, future coastal or Chukchi Seas. In Environmental assess­ sastoyanii zapasov serykh kitov (The state of shallow-water development must be ment of the Alaskan continental shelf. An­ the gray whale stock.) Rybn. Khoz. well monitored to determine any ef­ nual Reports 1: I-99. U.s. Dep. Commer., 3:15-17. [In Russ., Transl. by S. Pearson, Natl. Oceanic Atmos. Admin., Environ. Res. Natl. Mar. Mammal Lab., Nat. Mar. Fish. fects on any critical stages of this Lab., Boulder, Colo. Serv., NOAA, Seattle, Wash., 1979,7 p.] whales' life cycle. For these reasons, Brownell, R. 1977. Current status of the gray --::--:-__, and 198 I. The whale. Rep. Int. Whaling Comm. 27:209­ history of gray whale harvesting off habitat protection of the coastal en- 211. Chukotka. Rep. Int. Whaling Comm. Bryant, P., and C. Lafferty. 1980. The gray 31 :503-505. whales of Guerrero Negro. Whalewatcher Kleinenberg, S. E., and T. I. Makarova 6Tyack, P., C. Clark, and C. Malme. 1983. 14(4):3-5. (editors). 1955. Knitboinyi promysel Sovet­ Migrating gray whales alter their motion in Cederlund, B. A. 1939. A subfossil gray skogo Soiuza (The whaling industry of the response to sounds associated with oil develop­ whale discovered in Sweden in 1859. ZooI. ). Izd. "Rybnoe Khozyaistvo", ment. (Abstr.) In Proceedings of the 5th Bien­ Bidr. Upps. 18:269-285. Moscow, 117 p. [In Russ.) nial Conference on the Biology of Marine Cordoba, F. 1981. La ballena gris y la Marquette, W., and H. Braham. 1982. Gray Mammals, November 27-December I, 1983, explotacion de fosfora en whale distribution and catch by Alaskan Boston, Mass., p. 104. (Available from first (The gray whale and the exploitation of Eskimos: A replacement for the bowhead author, Woods Hole Oceanogr. Inst., Woods phosphorus in southern Baja whale? Arctic 35:386-394. Hole, MA 02543.) California). In Ballena Gris. Centro de ____, M. K. Nerini, and

46(4),1984 13 R. V. Miller. 1982. studies, and dynamics of the California gray whale Scammon, C. M. 1874. The marine mam­ Autumn 1980-Spring 1981: Harvest, (Eschrich/ius rabus/us). Ph.D. Thesis, mals of the northwestern coast of North biology and distribution. Rep. Int. Whaling Univ. Wash., Seattle, 265 p. America. John H. Carmany and Co., San Comm. 32:357-370. ____, D. Rice, and A. Wolman. 1980. Franc., 319 p. Mitchell, E. 1979. Comments on magnitude Preliminary population estimate for the Sprague, J. G., N. B. Miller, and J. L. Sumich. of early catch of east Pacific gray whale California gray whale based upon Monterey 1978. Observations of gray whales in La­ (Eschrich/ius rabus/us). Rep. Int. Whaling shore censuses, 1967/68 to 1978/79. Rep. guna de San Quintin, northwestern Baja Cali­ Comm.29:307-314. Int. Whaling Comm. 30:359-368. fornia, Mexico. J. Mammal. 59:425427. Nerini, M. K. 1984. Feeding ecology of the ----:-c:-:---.' , and Sund, P. 1975. Evidence of feeding during gray whale. In M. L. Jones, S. L. Swartz, and 1983. Population assessment of the gray migration and of an early birth of the Califor­ J. S. Leatherwood (editors), The gray whale, Eschrich/ius rabustus, from Califor­ nia gray whale (Eschrichtius rabustus). whale. Acad. Press, N.Y. nia shore censuses, 1967-1980. Fish. Bull., J. Mammal. 56:265-266. ____, and J. S. Oliver. 1983. Gray U.S. 81:267-281. Swartz, S. L., and M. L. Jones. 1982. Demo­ whales and the structure of the Bering Sea Rice, D. W. 1961. Census of the California graphic studies and habitat assessment of benthos. Oecologia (Berl.) 59:224-225. gray whale. Nor. Hvalfangst-Tidende 50: gray whales, Eschrichtius rabustus, in Nishiwaki, M., and T. Kasuya. 1970. Re­ 219-225. Laguna San Ignacio, Baja California Sur, cent record of a gray whale in the adjacent 1983. Gestation period and Mexico. Cetacean Res. Assoc., San Diego, waters of Japan and a consideration on its fetal growth of the gray whale. Rep. Int. Calif., 56 p. (Submitted to U.S. Mar. Mam­ migration. Sci. Rep. Whales Res. Inst. Whaling Comm. 33:539-544. mal Comm., Wash., D.C., as Contract No. Tokyo 22:29-38. ____, and A. A. Wolman. 1971. The MM20792194, avail. U.S. Dep. Commer., Omura, H. 1974. Possible migration route life history and ecology of the gray whale Natl. Tech. Inr. Serv., Springfield, Va., as of the gray whale on the coast of (Eschrich/ius rabus/us). Am. Soc. Mam­ PB82-123373.) Japan. Sci. Rep. Whales Res. Inst., Tokyo mal., Spec. Publ. 3, 142 p. ____, and 1983. Gray 26:1-14. --:---c:-::' D. E. Withrow, and whale (Eschrichtius rabustus) calf production Patten, D. R., and W. F. Samaras. L. A. Fleischer. 1981. Gray whales on the and mortality in the winter range. Rep. Int. 1977. Unseasonable occurrences of gray winter grounds in Baja California. Rep. Whaling Comm. 33:503-507. whales. Bull. South. Calif. Acad. Sci. Int. Whaling Comm. 31:477493. Townsend, C. H. 1887. Present condition of 76:205-208. Rugh, D. 1984. Fall migration and census of the California gray whale fishery. Bull. Pike, G. C. 1962. Migration and feeding of the gray whale at Unimak Pass, Alaska. In U.S. Fish Comm. 6:346-350. the gray whales (Eschrichtius rabus/us). M. L. Jones, S. L. Swartz, and J. S. Leather­ van Deinse, A. B., and G. C. A. Junge. 1937. J. Fish. Res. Board Can. 19:815-838. wood (editors), The gray whale. Acad. Recent and older finds of the California gray Ray, G. c., and W. E. Schevill. 1974. Feed­ Press, N.Y. whale in the Atlantic. Temminckia ing of a captive gray whale, (Eschrich/ius _-,-__' and H. Braham. 1979. Califor­ 2:161-188. rabus/us). Mar. Fish. Rev. 36(4):31-38. nia gray whale (Eschrich/ius rabus/us) fall Zimushko, V. V., and S. A. Lenskaya. 1970. Reeves, R. R. 1977. The problem of gray migration through Unimak Pass, Alaska, o pitanii serogo kita (Eschrichtius gibbasus whale (Eschrich/ius rabus/us) harassment at 1977: a preliminary report. Rep. Int. Whal­ Erx.) na mestakh nagula (Feeding of the gray the breeding lagoons and during migration. ing Comm. 29:315-320. whale (Eschrichtius gibbasus Erx.) at forag­ U.S. Mar. Mammal Comm., Wash., D.C., --:--,-c-:' and M. A. Fraker. 1981. Gray ing grounds). Ekologiya 1(3):26-35. [In Rep. MMC-76/06, 60 p. whale (Eschrich/ius rabustus) sightings in the Russ., Transl. by Consult. Bur., Div. Plenum Reilly, S. B. 1981. Population assessment eastern Beaufort Sea. Arctic 34: 186-187. Publ. Corp., N.Y., 1971, p. 205-212.]

14 Marine Fisheries Review