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Home , History of whaling, Physeter, Spermaceti, Sperm oil, Sperm whale, Sperm whaling

The Sperm , macrocephalus

MERRILL E. GOSHO, DALE W. RICE, and JEFFREY M. BREIWICK

Introduction hole asymmetrically situated on the Sperm have a strong school­ left side of the head near the tip. The ing instinct, forming schools of The Physeter Y-shaped lower jaw on the underside females and young, young males, and 1 macrocephalus Linnaeus, 1758 , is of the head contains two rows of mixed ages and sexes. The older adult the largest of the toothed whales 20-30 erupted teeth. males are often solitary and tend to (Odontoceti). In the past, males as The interior of the mouth and the migrate to higher latitudes than do the large as 20 m (65 feet) in length have surrounding area are often white, in females and younger . been recorded; today, however, males contrast to the rest of the dark body, Sperm whales are noted for their larger than 18 m (60 feet) are rare. which has been variously described as ability to make prolonged deep dives. The maximum length of female sperm black, dark bluish-gray, slate gray, Large adult males can remain whales is 12 m (40 feet)(Berzin, 1971). iron gray, purplish brown, grayish­ submerged for over an hour, while The average size of a at is 4 brown, or blackish-brown. The sperm females and younger animals usually m (13 feet) (Ohsumi, 1%5). Gaskin whale has no dorsal ; however, a surface after 15-20 minutes (Rice, (1972) reported that sperm whales can hump or a series of humps are present 1978). Clarke (1976) observed two reach a maximum age of 50 years. along the dorsal surface of the tail­ large adult male sperm whales diving The sperm whale is distinguished stock. The of the trunk is cor­ off Durban, , in water by an unusually large head that is rugated into many series of over 3,193 m (10,476 feet) deep. The from about one-fourth to one-third of longitudinal ripples. whales made dives lasting 82 and 83 its total body length. It is the only liv­ minutes; when captured later their The authors are with the National Marine ing cetacean that has a single blow Laboratory, Northwest and Alaska stomachs contained bottom-dwelling Center, National Marine Fisheries Ser­ . 'This name is used instead of Physeler calodon vice, NOAA, 7600 Sand Point Way N.E., Bin The head of the sperm whale con­ according to Husson and Holthius, 1974. CI5700, Seattle, WA 98115. tains the organ, which is a large reservoir for spermaceti oil. Sperm whales are hunted for this oil as well as for the lower grade oil con­ tained in the . The remainder of the whale is processed into feed, fertilizer, and, to a limited ex­ tent, food and pharma­ ceuticals.

Distribution and Migration Habitat Sperm whales inhabit all of the world. Their distribution is de­ pendent on their food source and A sperm whale surfaces in the Southern . Note the characteristic wrinkl­ suitable conditions for breeding, and ed appearance of the body surface and the obtusely rounded dorsal hump. varies with the sex and age composi­ Photo by G. Joyce. tion of the groups.

54 Marine Fisheries Review These whales usually inhabit the During the summer, sperm whales around the southern coasts of Africa offshore waters. Berzin (1971) re­ migrate to higher latitudes; the and , and of males south of ported that sperm whales are re­ mature males migrate much farther Cape Horn (Mackintosh, 1%5). It is stricted to waters deeper than 300 m poleward than do the females and not clear if the mature males which (1,000 feet), while Watkins (1977) re­ younger males. The females and migrate to higher latitudes always ported that they are usually not found younger animals may be restricted in return to the same breeding herds in waters less than 1,000 m (3,300 their migrations by an intolerance to (Gaskin, 1972). feet) deep. When found close to land, low water temperatures. Nishiwaki the sperm whale concentrations are (1967) reported that female sperm usually associated with a sharp drop whales in the North Pacific do not Pacific Ocean in depth of the bottom where upwell­ migrate into waters having a ing occurs. In these areas, organic temperature lower than lOoC. During the summer, sperm whales production is high, implying the Gilmore (1959) reported that harem are widely distributed throughout the presence of a good food supply herds with mixed sexes and ages are entire Pacific Ocean. The females and (Clarke, 1956; Berzin and Rovnin, restricted by the 20°C isotherm, young sperm whales usually remain in 1966). although Gaskin (1972) observed tropical and temperate waters be­ Deep-water () nursery schools with very small tween lat. 45°N and lat. 45°S (Rice, are the major food of sperm whales. in waters to the east of 1978), while the males continue north Global distribution is in­ with a temperature of 14°-16°C. to the , Aleutian fluenced by such factors as their in­ Because breeding herds are confined Islands, and Bering , and south to ability to withstand salinities below 30 almost exclusively to the warmer the . The northernmost 0/00 and a temperature requirement of waters, many of the larger male sperm boundary for male sperm whales is 10°-20°C which is necessary for their whales return in the winter to the from Cape Navarin Oat. 62°N) to the eggs to develop (Akimushkin, 1963). lower latitudes to breed. Pribilof Islands (Omura, 1955). Berzin (1971) reported that the occur­ In the Northern Hemisphere, Sperm whales are usually distributed rence of sperm whales is rare in the sperm whale stocks are believed to be below lat. 40 0 N during the winter. Yellow and East China , the Sea segregated units. However, in the In the spring, sperm whales in the of , and the Baltic Sea; these Southern Hemisphere (Fig. la, b), western North Pacific begin to seas are relatively shallow and have a there could be a gradual mixing of migrate from the Philippines to the low salinity. both male and female sperm whales southern coast of Japan, along the

An immature male sperm whale lies on the slipway of the former station at Coal Harbour, Vancouver Island, B.c. Note the large barrel-like head and the long, narrow lower jaw. Photo by D. W. Rice.

46(4),1984 55 Figure la. - Pacific Ocean distribution of sperm whales based on past commer­ cial catches (Gilmore, 1959).

Japan coast to the Kurile Islands, and present in midsummer. Two annual November to April (Rice, 1974). Dur­ up to Kamchatka (Berzin, 1971). peaks of abundance occur off central ing the summer, sperm whales are Many of the mature males leave their California in mid-May and mid­ found in the Gulf of Alaska, Aleutian herds to continue northward to the September, which suggest a north­ Islands, and southeastern . Aleutian Islands, Bering Sea, and the ward migration in the spring and a grounds in the Gulf of Alaska (Ohsumi, 1966; southward migration in the fall. Dur­ Pacific south of lat. 40 0 N (Fig. 1a) Ohsumi and Masaki, 1977). ing the winter, breeding schools of were historically located around the In the eastern North Pacific, sperm sperm whales are frequently sighted Hawaiian Islands, from the Bonin whales are commonly found off cen­ over the continental slope off Califor­ Islands to Midway Island, from the tral California, although very few are nia from lat. 33°N to lat. 38°N from Philippines and Borneo along the

56 Marine Fisheries Review Figure lb. - Atlantic and distribution of sperm whales based on past commercial catches (Gilmore, 1959).

Equator to South America, along the 70 0 -75°N in the Ocean (Berzin, restricted to latitudes less than lat. western coast of South America, and 1971). Sperm whales have been 40 0 N (Slijper, 1962). around the Society Islands, the Mar­ reported as as , Past sperm whaling grounds in the quesas, , Samoa, New Zealand, west of Island, the Gulf of Mexico, , and and in the Tasman Sea (Townsend, southeastern off the Mur­ off the southeastern United States are 1935). man Coast, and the Kanin Peninsula. shown in Figure la, while those in the Female sperm whale sightings have rest of the are shown Atlantic and Arctic Oceans occurred as far north as lat. 54°N in in Figure lb. In addition, small The northernmost limit of male the North Atlantic, although female numbers of sperm whales were taken sperm whales is approximately lat. and young sperm whales are generally during the summer in , off

46(4),1984 57 110 110 of Sp~rmwholes.

0 1 - 50. 120 • 100. • 200. ... 1,01- 1.50 13 •

Figure 2. - Total catch of sperm whales in the Antarctic and average yield of sperm whales per effective catcher's day's work (c.d.w.) for seasons 1950-51, 1951-52, 1954-55, and 1955-56 apportioned between squares of 5 degrees latitude and 10 degrees longitude (Holm and ]onsgard, 1959).

Labrador, , Green­ Stream and West currents whales would summer in the northern land, , , and around along the North American coast as far areas and winter around the , the Faroe and Shetland Islands; they as Davis Strait in the summer and , Canary, and Cape Verde frequently appear off the coasts of returned to the Gulf of Mexico, An­ Islands. Martin (1982) reported that a , , , and the tilles, and Bermuda in the winter to 14 m (46-foot) male sperm whale cap­ Netherlands, and strandings often oc­ breed. However, Mitchell (1975) tured off Iceland in August 1981 had cur there. Sperm whales also occur in reported that a male sperm whale, previously been harpooned in the the , off the west coasts tagged in 1966 off Nova Scotia, was Azores in August 1980. of Spain and , and in the captured over 7 years later off Spain Sperm whales occur year-round off (Berzin, 1971). in August 1973. the coasts of southern Africa and Tomilin (1957) believed that sperm Tomilin (1957) also believed that in South America (Berzin, 1971). The whales followed the warm Gulf the eastern North Atlantic sperm southernmost boundary of harem

58 Marine Fisheries Review animals in the South Atlantic appears to be around lat. 50 o -54°S. Indian Ocean In the 18th and 19th centuries, sperm whaling in the Indian Ocean was conducted in all seasons. However, most whaling (Fig. 1b) oc­ curred in the western half of the In­ dian Ocean (Townsend, 1935). Gambell (1967, 1972) reported that sperm whales in the region around Durban were present year-round, moving into and out of the area at different times of the year; there was a net northward movement up the coast in the early part of the year. Bannister and Gambell (1965) believed that schools of sperm whales drifted south in the austral spring. In the central In­ Figure 3. - Sperm whale stock divisions of the International Whaling Commission. dian Ocean, Soviet surveys found large concentrations of sperm whales to the north of St. Paul and Amster­ Antarctic, the males travel to their into Eastern and Western Divisions dam Islands in December and wintering grounds. Best (1969) (Fig. 3) (Allen, 1980; IWC, I980a). January (Berzin, 1971). reported the presence of a skin-film In the North Atlantic Ocean, there Bannister (1969) reported that, in containing Antarctic diatoms on male is no indication that more than one the summer, sperm whales follow the sperm whales taken off the southwest sperm whale stock exists. For this coastlines of Australia, moving coast of South Africa in April and reason, the IWC treats sperm whales southward along the western coast May. This diatom is rarely present in in the North Atlantic as a single stock and westward or southwestward warmer waters and indicates that for management purposes (IWC, along the southern coast. Bannister these animals had recently arrived 1981). (1969) also hypothesized that three from the Antarctic. This skin film of At present, there is no clear-cut stocks inhabited the Indian Ocean: Antarctic diatoms was also found on evidence that sperm whales in the In­ One off the southeast coast of Africa, sperm whales in Albany (southwest dian Ocean north of the Equator are an oceanic stock around Amsterdam Australia) and New Zealand by Ban­ from the same stocks as those south and St. Paul Islands, and an eastern nister (1968; 1969) and in South of the Equator (IWC, 1980b). The stock off western Australia. Georgia by Matthews (1938). latter are treated as Southern Antarctic IWe Sperm Whale Hemisphere sperm whale stocks for Stocks and Divisions management purposes. Southern Male sperm whales travel up to the Hemisphere sperm whales are ice edge in the Antarctic (Berzin, To make population assessments separated into nine divisions extend­ 1971). Sperm whales are more evenly and manage the world sperm whale ing from the Antarctic ice edge to the distributed in the Antarctic than at stocks, the International Whaling Equator and between the meridians of lower latitudes and travel from one Commission (IWC) has divided the longitude shown in Figure 3. region to another around the Antarc­ world oceans into the following areas: tic in search of food (Kirpichnikov, North Pacific, North Atlantic, Indian 1950). Holm and Jonsg§.rd (1959) Ocean, and Southern Hemisphere Life History and reported that sperm whales were par­ (Fig. 3). Feeding ticularly abundant from long. 70 0 W The IWC reported the possibility eastward to long. l700 W (Fig. 2). that more than two stocks of sperm Sperm whales feed mainly on Because female sperm whales do whales may inhabit the North Pacific; medium- to large-sized mesopelagic not enter Antarctic waters, the 2Oth­ however, because of a lack of data squids, including the giant squids Ar­ century there was exclusively precisely delineating the geographical chiteuthis and Moroteuthis; adult for males. Sperm whaling in the Ant­ boundaries, and because the mature males generally take larger squids arctic took place mainly during the males from these different stocks may than do the females and immature austral summer (December to intermingle in the Bering Sea, the males. Sperm whales-especially March). After summering in the North Pacific Ocean was divided only mature males in higher latitude

46(4),1984 59 waters - also take a significant quanti­ ing schools only during the mating vessels visited various parts of the ty of large demersal and mesopelagic season. Atlantic (off Newfoundland, Cape sharks, skates, and (Berzin, Verde, Brazil, and the Falkland 1971; Clarke, 1980). Natural Mortality Islands), the Caribbean Sea (off the West Indies), and the Gulf of Mexico Important natural mortality factors are unknown. Although the sperm (Harmer, 1928; Scammon, 1874). Sperm whale populations are whale is renowned for occasional England began sperm whaling in organized into two types of groupings mass strandings of entire schools, this 1785; France followed in 1790. Al­ (Best 1979): 1) Breeding schools (also factor is insignificant in terms of though France had 40 vessels engaged called harem schools or mixed overall mortality. Estimates of total in sperm whaling, the French Revolu­ schools) and 2) bachelor schools. natural mortality rates in different tion forced an end to its whaling Older males are often solitary. areas range from 0.05 to 0.09 per year (Harmer, 1928). By the end of the Breeding schools consist of females (IWC, 1971). 18th century, Portugal had also begun of all ages and juvenile males, and sperm whaling (Slijper, 1962). usually contain about 20-40 in­ Between 1787 and 1791, some ships dividuals (larger groups of up to Exploitation and Population Size began venturing around Cape Horn several hundred appear to be History of Exploitation to the Pacific Ocean, while others assemblages of two or more schools). moved into the Indian Ocean. The mature females ovulate in spring The abundance of whales along the Around 1800, U.S. were and early summer- April through eastern coast of America was one of operating off , Chile, and the August in the Northern Hemisphere the main reasons for the early settle­ Galapagos Islands. In 1802, they and October through February in the ment of by the English. traveled to New Zealand and the Southern Hemisphere (the breeding During the 17th century, the animals Molucca Islands. In 1822, more than season of equatorial populations is were hunted close to shore and 30 ships were for sperm not known). During this season, one catches consisted primarily of hump­ whales off the coast of Japan; this or more large mature temporari­ back, Megaptera novaeangliae, and number increased to 100 by 1835 ly join each breeding school. right, Balaena glacialis, whales. (Harmer, 1928; Scammon, 1874). The calves, which average about 4 Sperm whaling began on the New Harmer (1928) reported that by 1842 m (13 feet) long at birth, are born England coast around 1712 (Harmer, there were 594 U.S. whaling vessels after a gestation period of about 15 1928; Starbuck, 1878). and 230 from other nations. months. The period lasts 1-2 Humpback, right, and sperm By 1846, Hawaii had become an years, but the calves probably begin whales are relatively slow-swimming important whaling center, harboring taking solid food long before wean­ whales and were accessible to early more than 600 ships. In addition, ing. Females attain sexual maturity at whalers using open boats. Further­ numerous merchants and a mean age of 9 years, when they more, these whales floated after ship's chandlers had moved their average 9 m (30 feet) long. They re­ death. During the 17th and first half businesses there (Slijper, 1962). How­ main in the breeding schools and pro­ of the 18th centuries, the whales were ever, from 1846 Yankee whaling duce a calf every 3-6 years. In males, killed and towed back to shore sta­ began to decline. puberty is prolonged; beginning at tions for processing. The whale oil The number of U.S. vessels engag­ about 9 years of age and a body which was produced by this process ed in all whaling had increased from length of 9 m (30 feet), it reaches com­ was used for lighting, lubrication, and 500 in 1835 to 736 in 1846 (Table 1). pletion when the testes are fully sper­ softer kinds of (Ts:'lnnessen and However, the number of U.S. whal­ matogenic at about 20 years of age Johnsen, 1982). ing vessels had dropped to 514 by and a body length of 12 m (39 feet). The on-board tryworks and the 1861 and to 124 by 1886. Further­ Bachelor schools consist entirely of spermaceti were developed more, not all of these vessels were males, which, as they approach about 1750. The on-board tryworks engaged solely in sperm whaling. physiological sexual maturity, leave enabled the whaling ships to process Harmer (1928) reported that the the breeding schools and aggregate in the whales on the whaling grounds sperm whaling fleet consisted of 315 loose groups of up to about 40 without having to return to port with U.S. vessels in 1844, 231 vessels in animals. Males continue to grow long each catch. The spermaceti candle 1870, and dropped to 134 vessels in after sexual maturity. As they grow was a smokeless candle which increas­ 1875. older, they become less gregarious, ed the market for spermaceti oil The estimated annual sperm whale and by the time they are about 30 (Kugler, 1981). catch also declined during this period, years old and 13 m (43 feet) long, they In 1762, about 78 U.S. vessels were fluctuating between 6,000 and 8,000 remain solitary most of the year; at engaged in whaling; this number in­ sperm whales from 1835 to 1845 and this age, they have attained "social" creased to 125 by 1770 (Scammon, between 3,000 and 5,000 sperm maturity and consort with the breed- 1874). Searching for whales, these whales from 1846 to 1861. By 1886,

60 Marine Fisheries Review Table 1.-Number of U.S. whaling vessels, barrels of , and estimated sperm whale catch', (1835-72 data from during and after the 1870's occurred Scammon, 1874; 1877-86 data from Clark, 1887). when the effort decreased after the U.S. whaling Barrels of Est. sperm U.S. whaling Barrels of Est. sperm Year vessels 2 sperm oil whale catch Year vessels 2 sperm oil whale catch 1860's (IWC 1969). Labor also became a problem for 1835 500 172,683 7,598 1859 625 91,408 4,022 1836 507 132,130 5,814 1860 569 73,708 3,243 the U.S. whaling industry. In the mid­ 1837 509 181,724 7,996 1861 514 68,932 3,033 dle of the 19th century, labor and 1836 550 131,856 5,801 1862 423 55,641 2,449 1839 676 150,000 6,600 1883 353 65,ffi5 2,862 capital were shifted away from whal­ 1840 559 157,791 6,943 1864 307 64,372 2,833 ing and shipping and into agricultural 1841 585 159,304 7,Wi! 1865 276 33,242 1,483 and mineral development, where 1842 594 165,837 7,288 1886 283 36,883 1,614 greater profits existed. A skilled 1843 627 186,985 7,347 1867 312 43,433 1,911 1844 647 139,594 6,142 1886 329 47,174 2,076 worker could earn two or three times 1845 696 157,917 6,949 1869 336 47,936 2,109 1846 736 95,217 4,190 1870 321 55,183 2,428 as much onshore as he could in whal­ ing (T~nnessen and Johnsen, 1982). 1847 723 120,753 5,313 1871 288 41,534 1,827 1848 659 107,976 4,751 1872 218 44,881 1,975 Manpower was lost to the developing 1849 614 100,944 4,442 cotton industry (1846) and to the 1850 543 92,892 4,088 1877 183 41,119 1,810 1851 553 99,591 4,382 1878 179 43,508 1,914 California gold rush (1849) (Slijper, 1852 620 78,872 3,471 1879 178 41,308 1,817 1962). Whaling crews were initially 1880 173 37,614 1,856 1853 661 103,077 4,535 composed almost entirely of 1854 688 73,696 3,243 1881 177 30,600 1,346 1855 638 72,649 3,197 1882 161 29,864 1,315 Americans, but were later recruited 1856 635 80,941 3,562 1883 147 24,595 1,082 from foreign countries (Starbuck, 1857 655 78,440 3,452 1864 144 22,670 998 1856 854 81,941 3,606 1885 133 24,203 1,065 1878). 1886 124 23,312 1,025 The (1861-65)

'Sperm whale catch estimated using 25 barrels per sperm whale plus 10 percent for mortally wounded and lost whales not only further reduced the labor not landed. force, but also destroyed a con­ 'All U.S. whaling vessels, not solely sperm whaling vessels. siderable portion of the whaling fleet. Confederate raider ships, such as the Alabama (McClung, 1978) and the Table 2.-Estimated catch' and ellort for sperm whaling, 1800-1909 (lWC, 1969~ Shenandoah (Starbuck, 1878) cap­ U.S. sperm whaling Worldwide tured and sank many Yankee whaling

Gross Avg. no. of Add 15 ships. tonnage whales per Avg. Average Yearly Avg. no. of percent During this period, sperm oil was Yearly of Barrels vessel voyage barrels total whales wounded barrels registered per ton per length per catch in per + ship- receiving increasing competition from Decade of oil whalers of boat year 2 (yr)' whale! barrels year wreck other means of illumination, notably 1800-09 11,910 5,810 2.ffi 37.0 1.0 (24.0)est. 19,850 827 951 from natural gas but also from 1810-19 14,726 6,127 2.40 19.7 1.2 (24.0)est. 22,655 944 1,201 1820-29 64,951 35,896 1.81 30.6 2.0 24.0 92,787 3,886 4,446 vegetable oils, animal , 1830-39 126,334 88,039 1.43 23.3 2.0 33.0 147,414 4,467 5,137 whale oil, and ultimately from 1840-49 130,628 145,557 0.90 15.6 2.2 36.2 145,790 4,027 4,631 1850-59 85,333 145,726 0.59 10.7 2.2 37.3 94,814 2,542 2,923 (Kugler, 1981). 186Q-69 53,616 90,903 0.59 10.5 2.1 32.9 59,573 1,811 2,083 The sperm whale catch and effort 1870-79 42,454 46,258 0.92 11.4 2.4 32.1 47,171 1,470 1,692 188D-89 24,617 25,744' 0.96 13.5 2.2 (30.2)net 27,352 906 1,042 continued to decline in the early part 1890-99 14,321 9,781' 1.46 24.0 2.0 (28.3)net 15,912 562 646 1900-09 16,362 5,711' 2.87 38.4 1.6 26.4 18,202 689 792 of the 20th century. In 1925, when the 1910-19 42.5 1.5 whaling vessel John R. Manta return­ 1920-29 24.3 1.0 ed to New Bedford for the last time, 'The total catches, in numbers, were estimated from the U.S. production of oil, divided by the average production per the era of Yankee sperm whale hunt­ whale, increased by the proportion of foreign vessels in the fishery, and by 15 percent to allow for whales wounded and dying, but not processed, and whales caught by vessels wrecked before returning to home port. ing was over (Hegarty, 1959). 'Data from Townsend (1935). Modern mechanized whaling devel­ 'For these years, tonnage of steam whalers (introduced in 1881 and specializing in Arctic whaling) subtracted from total. oped in Norway in the latter half of the 19th century and was initially used to exploit the faster swimming ror­ the estimated catch had dropped to (CPUE) (measured as the sperm oil quais (blue, fin, sei, and Bryde's 1,025 sperm whales. production per ton of boat, or per whales). The techniques of modern Many factors contributed to the vessel per year) for the 19th century whaling involved the use of steam­ decline of the U.S. sperm whale (Table 2) shows a marked decline driven (later, diesel-driven) whaling fishery. Starbuck (1878) reported that from the early years to a minimum in vessels, fired from cannons, whaling as a business declined because the 1850's and 1860's. This would be and grenades, attached to the har­ of the scarcity and shyness of whales, consistent with a decline in abundance poon, which exploded in the whale. In requiring longer and more expensive of sperm whales as a result of heavy 1863, the first steam-powered whaling voyages. The catch per unit of effort exploitation. An apparent recovery vessel which had separate guns for

46(4),1984 61 shots and grenade shots was From 1883 to 1924, the average an­ vessel size, horsepower, captain and introduced. In 1868, Sven Foyn per­ nual worldwide catch was less than gunner experience, and technical ad­ fected a cannon which combined the 1,000 sperm whales (Tables 2 and 3). vances are important considerations harpoon and grenade into one instru­ After 1924, the annual catch increas­ in standardizing effort. ment (T16nnessen and Johnsen, 1982). ed, but until 1951 did not exceed The most important technical ad­ In 1903, the first modern factory ship 10,000 sperm whales. However, as the vance has been the introduction of was used (Slijper, 1962). abundance of the larger echo-locating equipment such as Although some attempts were decreased, greater attention was paid ASDIC (Anti- Detection made to modernize American whaling to the smaller rorquals and the sperm Investigation Committee), a type of methods, these were largely unsuc­ whales. Furthermore, following . In the fishery the cessful. The United States was never a World War II, sperm whale oil once use of ASDIC tends to frighten major participant In modern again came into demand as an ex­ whales, causing them to remain on the mechanized whaling. treme pressure and in­ surface and thus become easier to dustrial (Frost, 1978). The catch catch. In catching sperm whales, of sperm whales reached 29,000 in ASDIC has been used to track whales Table 3.-World catch of sperm whales (catch data for 1964; the average annual catch underwater, so that the catcher boat 1910-37 from Clar1

1925 1,475 1950 8,219 1975 21,045 cess. The assumption is that CPUE, time) (Allen and Kirkwood, 1977) and 1926 1,775 1951 18,281 1976 17,134 where effort is usually defined as the a length-specific model which is based 1927 1,441 1952 11,558 1977 12,279 1928 1,989 1953 9,577 1978 11,065 number of boat-days spent whaling or on the length composition of sperm 1929 2,074 1954 13,543 1979 8,536 the number of hours spent searching, whale catches (Beddington and 1930 1,311 1955 15,593 1980 2,091 is proportional to population abun­ Cooke, 1981). Models incorporating 1931 597 1956 18,590 1981 1,456 dance. A major problem in assessing biological parameters such as changes 1932 811 1957 19,156 1982 526 1933 1,423 1956 21,646 whale stocks using a time series of in survival and pregnancy rate, harem 1934 1,999 1959 21,298 catch and effort data is standardizing size, and age at sexual and social lCalendar years for all areas outside Antarctic. Antarctic effort so that units of effort in each maturity are also used in assessing seasons are 19091910, etc. year are equivalent. Factors such as sperm whale stocks. Tables 4-6 give

Table 4.-Estimates 01 initial (1910) and current (1982) sperm whale stock size lor the North Pacific Ocean. Table 5.-Estimates of initial (1905) and current (1981) sperm whale stock size (on thousands) Exploitation for the North Atlantic Ocean.' Reference population Area year Sex and age size Spain l Population Reference component year Sex Iceland Azores AB Western Initial Males (age 11 + ) 128,500 North (1910) Femaies (age 10+) 180,900 Exploitable Initial Males 28.6 13.6 12.6 12.9 Pacific' (1905) Females NA' 30.2 28.0 28.6

Current Males (age 11 +) 61,000 Current Males 19.1 3.8 2.3 2.7 (1982) Females (age 10+) 137,100 (1981) Females NA 18.5 15.2 15.9

Eastern Initial Males (age 13+ )' 142,700 Mature Initial Females 43.0 35.7 33.1 33.7 North (1910) Females (mature) 168,300 (1905) Pacific 2 Current Females 38.9 19.7 15.7 16.7 Current Males (age 13 + ) 111,400 (1981) (1982) Females (mature) 162,600 'Data from IWC (1982). Separate estimates of the North Atlantic stock using a variant of the 'Data from IWC (1983). Based on length·specific model. length·specific method. 'Data from IWC (1979). Based on catch and effort data. 'Two series of Spanish catch and effort data have been used. lAverage age at recruitment. 'NA = no data available.

62 Marine Fisheries Review the most recent pre-exploitation and nent. Most relevant data required by tic, and northern Indian Ocean. In the current stock size estimates for sperm the models are lacking or are not well Western division of the North Pacific, whales in the North Pacific, and understood. The length-specific the catch limits for the 1982 and 1983 North Atlantic, and in the Southern model of Beddington and Cooke coastal seasons (taken by Japan) are Hemisphere. (1981) requires knowledge of the 450 and 400 whales, respectively (in­ Caution must be used in inter­ growth curve as well as other cluding an 11.5 percent of preting the population estimates in parameters and assumptions. The females). Tables 4-6. Although they are the CPUE data are confounded by most current estimates available, new geographical variations in density of Literature Cited analyses, using different data sets and whales, problems in interpretation Akimushkin, I. I. 1963. Golovonogie mol­ biological parameters, may produce (with possibly two or more "real" lyuski morei SSSR (Cephalopods of the seas of the U.S.S.R.). Izd. Akad. Nauk SSSR, much different estimates. The populations being fished as one), . [In Russ., transl. by Isr. Program estimates in Table 5 (North Atlantic) seasonal and annual changes in avail­ Sci. Transl., 1965, 223 p.; avail. U.S. Dep. may be especially unreliable due to ability, and the shifting of whaling Commer., Natl. Tech. Inf. Serv., Springfield, Va., as TT65-50013.l concern about the validity of certain operations with time. Allen, K. R. 1980. Conservation and man­ assumptions of the estimation techni­ Additionally, there are inconsisten­ agement of whales. Butterworth and Co., Lond., 107 p. que. There is also uncertainty in the cies in the available population ____, and G. Kirkwood. 1977. A assumption of a single stock. The estimates (e.g., as reported by the sperm whale population model based on estimates do suggest, however, that Scientific Committee of the IWC) as cohorts (SPCOH). Rep. Int. Whaling Comm. 27:268-271. there has been a decline in abundance, well as difficulties in establishing Bannister, J. L. 1968. An aerial survey for more so in males than females. stock identity. Inconsistencies be­ sperm whales off the coast of Western tween population estimates result Australia 1963-65. Aust. J. Mar. Fresh­ from using different population water Res. 19:31-51. Management _---,--,._. 1969. The biology and status models, each requiring different types of the sperm whale off Western Sperm whale populations or stocks of data and often different assump­ Australia - an extended summary of results of recent work. Appendix 7. Rep. Int. have been difficult to assess because tions. There is also a lack of data for Whaling Comm. 19:70-76. of the number of factors required by the more recent years, and, because of ---c-' and R. Gambell. 1965. The succession and abundance of fin, sei, and the assessment models. These factors the IWC whaling moratorium, future other whales off Durban. Nor. Hvalfangst­ include: Effort, effort modifiers, data will probably be minimal. Tidende 54(3):45-60. pregnancy rates, natural mortality At present there are zero quotas (no Beddington, J. R., and J. G. Cooke. 1981. Development of an assessment technique for rate, age at recruitment, reserve male catch allowed) for sperm whales in the male sperm whales based on the use of length ratio, and a density-dependent expo- Southern Hemisphere, North Atlan- data from the catches, with special reference

Table 6.-Estimates of stock size for Southern Hemi· sphere sperm whales.

Initial (1946) Current stock stock size size Stock Sex Division A' S' 1977' 1979'

Males 1 12,400 4.600 2 33,000 23.000 16.400 10,600 3 38,400 25,700 13,700 6.600 4 23.100 15,800 12,100 7.600 5 21,900 15,800 7,400 4,200 6 11.300 5,100 7 15,900 4,900 8 37,700 33,600 9 38,200 48,400 12,800 16.300 -- Total 229.900 111.300

Females 1 20,300 17,800 2 54,100 49,100 50,800 41.100 3 63,000 55,000 52.600 42,900 4 37.900 33,900 21.600 32.300 5 35.900 33,800 34,100 30,800 6 18,500 17,300 7 26,100 21,000 8 58,500 57,400 9 46,400 25,700 Total 360,700 299,400

'Data from IWC (1978). Males 20 + years old (15 + years old for Div. 9); Females 10+ years old. A sperm whale surfaces off California. Note the single, open on the 'Data from Iwe (1980b). left side of the snout. Photo by D. W. Rice.

46(4), /984 63 to the North-west Pacific stock. Rep. Int. vessels sailing from American ports-A con­ the sea. William Morrow and Co., N.Y., Whaling Comm. 31:747-760. tinuation of Alexander Starbuck's "History of 192 p. Berzin, A. A. 1971. Kashalot (The sperm the American whale fishery" 1876-1928. Mitchell, E. 1975. Preliminary report on whale). Izd. "Pishchevaya Promyshlennost", Reynolds Print., Inc., New Bedford, Mass., Nova Scotia fishery for sperm whales Moscow. [In Russ., trans!. by Isr. Program 58 p. (Physeter catodon). Rep. Int. Whaling Sci. Trans!., 1972, 394 p., avai!. U.S. Dep. Holm, J. L., and A. Jonsgilrd. 1959. Occur­ Comm. 25:226-235. Commer., Nat!. Tech. Inf. Serv., Springfield, rence of the sperm whale in the Antarctic and Nishiwaki, M. 1967. Distribution and Va., as TT71-50152.] the possible influence of the moon. Nor. migration of marine in the North __--,--,---, and A. A. Rovnin. 1966. Ras­ Hvalfangst-Tidende 48(4):161-182. Pacific area. Bul!. Ocean Res. Inst. Univ. predelenie i migratsii kitov v severo­ Husson, A. M., and L. B. Holthius. 1974. Tokyo I, 64 p. vostochnoi chasti Tikhojgo okeana, v Ber­ Physeter macrocephalus Linnaeus, 1758, the Ohsumi, S. 1965. Reproduction of the ingovom i Chukotskom moryakh (Distribu­ valid name for the sperm whale. !. sperm whale in the North Pacific. Sci. Rep. tion and migration of whales in the nor­ Meded. (Leiden) 48:205-217. Whales Res. Inst., Tokyo 19:1-35. theastern part of the Pacific Ocean, Bering IWe. 1969. Appendix VI: Report of the 1966. Sexual segregation of and Chukchee Seas). Izv. Tikhookean. IWC-FAO working group on sperm whale the sperm whale in the North Pacific. Sci. Nauchno-Issled. Inst. Rybn. Khoz. stock assessment. Rep. Int. Whaling Rep. Whales Res. Inst., Tokyo 20:1-16. Okeanogr. (T1NRO) 58: 179-207. [In Russ., Comm. 19:39-84. _----:-__, and Y. Masaki. 1977. Stocks trans!' by U.S. Dep. Inter., Bur. Commer. 1971. Report of the Special and trends of abundance of the sperm whale ., Seattle, Wash., 1966, p. 103-136 In K. Meeting on Sperm Whale Biology and Stock in the North Pacific. Rep. Int. Whaling I. Panin (editor), Soviet research on marine Assessments. Rep. Int. Whaling Comm. Comm.27:167-175. mammals of the Far East.] 21 :40--50. Omura, H. 1955. Whales in the northern Best, P. B. 1969. The sperm whale (Physeter 1978. Report of Scientific part of the North Pacific. Nor. Hvalfangst­ catodon) off the west coast of South Africa. Committee: Sperm whales. Rep. Int. Whal­ Tidende 44(6):323-345. (4) Distribution and movements. S. Afr. ing Comm. 28:57-62. Rice, D. 1974. Whales and whale research in Div. Sea Fish. Invest. Rep. 78:1-12. 1979. Annex E: Report of the the eastern North Pacific. In W. E. Schevill 1979. Social organization in subcommittee on sperm whales. Rep. Int. (editor), The whale problem; a status report, sperm whales, Physeter macrocephalus. In Whaling Comm. 29:65-74. p. 170-195. Harvard Univ. Press, Cam­ H. E. Winn and B. L. Olla (editors), 1980a. Report of the special bridge, Mass. Behavior of marine animals, volume 3: ceta­ meeting on sperm whale assessments 27 1978. Sperm whales. In ceans, p. 227-289. Plenum Press, N.Y. November to 8 December 1978. Rep. Int. D. Haley (editor), Marine mammals of the Clark, A. H. 1887. The American whale­ Whaling Comm., Spec. Issue 2:107-136. eastern North Pacific and Arctic waters, p. fishery. Science (Wash., D.C.) 9(217): 1980b. Annex F: Report of the 82-87. Pac. Search Press., Seattle. 321-324. sub-committee on sperm whales. Rep. Int. Scammon, C. M. 1874. The marine mam­ Clarke, M. R. 1976. Observation on sperm Whaling Comm. 30:79-96. mals of the north-western coast of North whale diving. J. Mar. Bio!. Assoc. U.K. 1981. Annex D: Report of the America together with an account of the 56:809-810. sub-committee on sperm whales. Rep. Int. American whale-fishery. John H. Carmany 1980. Cephalopoda in the diet Whaling Comm. 31:78-102. and Co., San Franc. (Repub!. in 1968 by of sperm whales of the Southern Hemisphere ____ 1982. Annex D. Report of the Dover Pub!., Inc., N.Y., 319 p.) and their bearing on sperm whale sub-committee on sperm whales. Rep. Int. Slijper, E. J. 1962. Whales. [Trans!. from biology. Discovery Rep. 37:1-324. Whaling Comm. 32:68-86. Dutch by A. J. Pomerans.] Basic Books, Clarke, R. 1954. Open boat whaling in the 1983. Annex D. Report of the Inc., N.Y., 475 p. Azores. Discovery Rep. 26:281-354. sub-committee on sperm whales. Rep. Int. Starbuck, A. 1878. History of the American Whaling Comm. 33:74-90. 1956. Sperm whales of the whale fishery from its earliest inception to the Kirpichnikov, A. A. 1950. 0 sovremennom year 1876. U.S. Comm. Fish Fish., Part 4, Azores. Discovery Rep. 28:237-98. rasprostranenii kashalotov v mirovom Committee for Whaling Statistics. 1959-1983. Rep. Comm. 1875-1876, append. 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64 Marine Fisheries Review