The Sperm Whale

Home , Ambergris, Southern fin whale, Whaling in Chile

The Great Whales: History and Status of Six Species Listed as Endangered Under the U.S. Endangered Species Act of 1973

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Authors Perry , Simona L.; DeMaster, Douglas P.; Silber , Gregory K.

Download date 24/09/2021 09:18:16

Link to Item http://hdl.handle.net/1834/26411 The Sperm Whale

Introduction whales with calves have an average dive Northern and Southern Hemisphere time of 8 minutes, while mixed groups populations at some time in their evo- The sperm whale, Physeter macro- without calves average 25 minutes lutionary history (Dufault et al.86). cephalus Linnaeus, 175885, is the larg- (Leatherwood et al., 1982; Mano, 1990; Large-scale oceanographic events, est and most sexually dimorphic mem- Gordon and Steiner, 1992). Body size, such as El Niño, also seem to affect the ber of the odontoceti or toothed whales. degree of social cohesion, oceano- distribution and movements of sperm Males reach a maximum length of 18.5 graphic features or bottom topography, whales (Whitehead et al.87), creating an- m, while the maximum length of fe- and prey species availability are all fac- nual and seasonal geographic variability. males is 12.5 m (Odell, 1992). Calves tors that may affect variation in dive North Pacific at birth have an average length of 4 m. behavior between individuals and areas. The most distinctive physical feature of In 1981, the IWC’s Scientific Com- Distribution and Migration the sperm whale is the exceptionally mittee designated two sperm whale large head to body ratio: about one-third Sperm whales inhabit all ocean ba- stocks in the North Pacific, western and of the total body length. The enormous sins, from equatorial waters to the po- eastern, based on whaling records, the head contains a reservoir of spermaceti lar regions. In general, their distribution results of Discovery tagging38, and the oil produced by the spermaceti organ. varies by gender and age composition intermingling of males in the northern And a black or grayish fatty substance, of groups and is related to prey avail- latitudes (Donovan, 1991). The bound- known as ambergris, is produced in the ability and certain oceanic conditions ary delineating western and eastern sperm whale’s intestines. Near the an- (Fig. 35). Mature females, calves, and North Pacific stocks became known as terior of the head and to the left is a immature whales of both sexes are the “Cambridge Line” (Fig. 36), and has single blow hole (Fig. 34). The narrow found in social groups in temperate and been much debated since its acceptance bottom jaw holds 17Ð29 pairs of func- tropical waters year round. Very rarely by the IWC’s Scientific Committee. tional teeth that align with indentations are female/immature groups found There is evidence from historical whal- in the upper jaw (Rice, 1989). higher than lat. 50°N and lat. 50°S ing data, ship-based surveys, and These toothed whales are rarely (Reeves and Whitehead, 1997). Male oceanographic features, that three or found in waters less than 300 m deep. sperm whales lead a mostly solitary life more North Pacific stocks of sperm They are often concentrated around oce- after reaching sexual maturity between whales should be recognized for man- anic islands in areas of upwelling and 9 and 20 years of age and travel into agement purposes (Bannister and along the outer continental shelf and regions as high as lat. 70°N in the North Mitchell, 1980; Kasuya, 1991). mid ocean waters (Rice, 1989). Atlantic and lat. 70°S in the Southern Sperm whales occur throughout the Sperm whales are deep divers. Soli- Ocean (Reeves and Whitehead, 1997). North Pacific. Female and immature tary adult males can stay submerged for General migration patterns vary be- whales are found year round in temper- over 60 minutes at recorded depths of tween males and females. In summer, ate and tropical waters from the Equa- over 2,000 m (Watkins et al., 1993). all sperm whales can be found at the tor to around lat. 45% N. During sum- Mixed groups of females and young highest latitudes of their range. In win- mer, mature male sperm whales are ter, female/immature groups migrate thought to move north into waters off closer to equatorial waters in both hemi- the Aleutian Islands, Gulf of Alaska, and 85 Both Physeter macrocephalus and Physeter spheres, possibly following warmer sea- catodon are scientific names used in the literature to refer to the sperm whale. However, P. surface temperatures (Kasuya and 86 Dufault, S., H. Whitehead, and M. Dillon. macrocephalus has been designated as the pre- Miyashita, 1988; Waring et al., 1993). 1997. An examination of current knowledge on ferred name by the International Code of Zoo- the stock structure of sperm whales (Physeter mac- logical Nomenclature’s Article 24(a) naming pro- Sexually mature males join these fe- rocephalus) world-wide. Unpubl. doc. SC/49/07 cedure (Rice, 1998). P. macrocephalus has never male/immature groups throughout the submitted to Rep. Int. Whal. Comm., 19 p. been used to refer to another species, while P. winter. The genetic homogeneity of 87 Whitehead, H., V. Papastavrou, and S. Smith. catadon has been used in reference to other ce- 1990. Sperm whales and El Niño off the taceans (e.g. pilot whales) (see Husson and sperm whales worldwide, suggests that Galapagos Islands. Unpubl. doc. SC/40/Sp4 sub- Holthius,1974). genetic exchange occurred between mitted to Rep. Int. Whal. Comm., 6 p.

61(1), 1999 59 the southern Bering Sea (Fig. 4). How- purposes: 1) Alaska, 2) California/Or- abundance in this region. Preliminary ever, Discovery tag38 data revealed con- egon/Washington, and 3) Hawaii (Bar- results of this survey (new abundance siderable east-west movement between low et al., 1997; Hill et al., 1997). These estimates are still being determined) Alaska and the western North Pacific, stock designations are based on distri- along with the results of previous sur- with little evidence of north-south butional data from sightings and catches veys in the same area, suggest that sea- movement in the eastern North Pacific only, and there are no reliable pheno- sonal sperm whale distribution in these (Ohsumi and Masaki, 1977; Wada, typic or genotypic data available for any waters varies annually, and brings into 1980; Taylor88). of the sperm whale “centers” in U.S. question the belief that sperm whales In the U.S. EEZ of the North Pacific, waters (Small and DeMaster, 1995; Hill are abundant during February off cen- three discrete population “centers” have et al., 1997). tral California (Bannister and Mitchell, been identified for stock assessment A survey conducted by the NMFS in 1980). More work is necessary in the the winter of 1997 (Sperm Whale Abun- temperate eastern Pacific before reliable 88 Taylor, B. 1997. Appendix 3. In D. P. DeMaster dance and Population Structure, or stock structure designations can be (Editor), Minutes from fifth meeting of the SWAPS) was designed to census these made (Taylor88; Taylor et al.89). Alaska Scientific Review Group, 7-9 May 1997, three “centers” and the area south of the Sperm whales in the western North Seattle, Wash., 21 p. Avail. from D. P. DeMaster, Director, Natl. Mar. Mammal Lab., 7600 Sand U.S./Mexican border to more clearly Pacific stock occur from the Equator, Point Way, N.E., Seattle, WA 98115. determine sperm whale distribution and along the Philippines, and up to the Kuril Islands and Kamchatka Peninsula, Russia (Fig. 5) (Kasuya and Miyashita, 1988; Shuntov, 1994). Based on Japa- nese coastal whaling data, tag returns, blood typing, and whale distribution associated with oceanographic current systems (i.e. Kuroshio Current and Oyashio Front), Kasuya (1991) pro- posed that there are three distinct sperm whale stocks in the North Pacific: 1) northwest North Pacific, 2) southwest North Pacific, and 3) eastern North Pa- cific. However, distinction between the three is confounded by the overlap in male distribution in northern latitudes (Kasuya and Miyashita, 1988). North Atlantic There is evidence from the harvest of Discovery tagged38 individuals that North Atlantic sperm whales are one geo- graphically continuous stock. One sperm whale tagged on the Scotian Shelf was killed over 7 years later off Spain (Mitchell, 1975c). From five to six hand- held harpoons from the Azore sperm whale fishery were recovered from whales killed off northwest Spain (Donovan, 1991), with an additional Azorean har- poon recovered from a male sperm whale killed off Iceland (Fig. 7) (Martin, 1982). Consequently, the IWC recognized the North Atlantic sperm whale population as one management stock.

89 Taylor, B. L., S. L. Mesnick, and A. E. Dizon. 1998. Progress report and suggested future research on using genetic data to define sperm whale stock structure in the North Pacific. Figure 34.—Aerial view of an adult sperm whale. Note the angled blow. NMML Unpubl. doc. SC/50/CAWS19 submitted to Rep. Collection. Int. Whal. Comm.

60 Marine Fisheries Review Figure 35.—Worldwide sperm whale distribution by gender. Adapted from Gosho et al. (1984).

Female and immature animals stay in Atlantic temperate or tropical waters year round. In the western North Atlan- tic, concentrations of female/immature groups are found in the Caribbean Sea (Gosho et al., 1984) and south of New England in continental-slope and deep- ocean waters along the eastern United States (Blaylock et al., 1995). In eastern Atlantic waters, female/immature groups aggregate in waters off the Azores, Madeira, Canary, and Cape Verde Islands (Fig. 7) (Tomilin, 1967). Mature male sperm whales have been recorded as far north as Spitsbergen (Fig. 18) (¯ien, 1990). All recent sightings and strandings from the eastern North Atlan- tic suggest a predominance of solitary and paired mature male sperm whales in waters off Iceland, the Faroe Islands, and the Figure 36.—The “Cambridge Line”; North Pacific sperm whale stock boundary Norwegian Sea (Gunnlaugsson and recognized by the IWC (Donovan, 1991). Sigurjónsson, 1990; Øien,1990; Chris- tensen et al., 1992a). Nine southern cepha- the coast of Iceland (Martin and Clarke, high latitudes. In the western North At- lopod species (known only from south 1986). This suggests that these male lantic, the Scotian Shelf’s Gully and of lat. 45°N) have been found in stom- whales are feeding in low latitudes of Shortland Canyon regions are fre- achs of male sperm whales killed off the North Atlantic before traveling to quented by male sperm whales during

61(1), 1999 61 summer (Whitehead et al., 1992; Mullins et al.90). Schmidly (1981), Fritts91, and Hansen et al.92 suggested that there is a distinct stock of sperm whales in the northern Gulf of Mexico. The NMFS recognizes these Gulf of Mexico sperm whales as one distinct stock (Waring et al., 1998). However, these whales are currently recognized as part of the entire North Atlantic stock by the IWC (IWC, 1980c). In U.S. waters of the Atlantic Ocean, sperm whales occur over the continen- Figure 37.—IWC Southern Hemisphere stock “Division” designations for sperm tal shelf edge (CeTAP70) and well into whales (Donovan, 1991). the continental slope and mid ocean regions during all seasons (Waring et al., either the warm core ring or the shelf- region into nine sperm whale “Divi- 1998). The NMFS has recognized these edge exists alone. This suggests that the sions” (Fig. 37). Donovan (1991) noted western North Atlantic whales as one interaction of variables such as sea sur- that these divisions are based more on stock (Waring et al., 1998). There is face temperature (SST), bottom topog- manipulating available data from com- seasonal variability in the latitudinal raphy, and associated primary produc- mercial whaling than on actual biologi- distribution of sperm whale concentra- tivity are all important to sperm whale cal information. They are: tions in this area. In winter, concentra- distribution in the Atlantic Ocean. tions exist east and northeast of Cape Division 1 Western Atlantic Northern Indian Ocean Hatteras (Fig. 6), and as spring ap- (long. 60°WÐ30°W), proaches, these concentrations shift There is no biological evidence to Division 2 Eastern Atlantic northward to waters off Delaware, suggest genetic interchange between (long. 30°WÐ20°E), Maryland, and Virginia, across the cen- sperm whales in the northern Indian Division 3 Western Indian tral portion of the Mid Atlantic Bight, Ocean and sperm whales south of the (long. 20°EÐ60°E), and into southern Georges Bank. Equator. This has led the IWC’s Scien- Division 4 Central Indian Throughout summer these concentra- tific Committee to assign separate stock (long. 60°EÐ90°E), tions are distributed as in the spring, but identity to Northern and Southern Division 5 Eastern Indian also include areas east and north of Hemisphere populations in the Indian (long. 90°EÐ130°E), Georges Bank, the Northeast Channel, Ocean (IWC, 1995b). Little is known Division 6 Eastern Australia and the continental shelf south of New of the northern Indian Ocean sperm (long. 130°EÐ160°E), England. In fall, the highest concentra- whale; however, distinct concentrations Division 7 New Zealand tions occur over the continental shelf of whales have been documented off the (long. 160°EÐ170°W), south of New England with some coast of Somalia and in Sri Lanka’s Gulf Division 8 Central Pacific (long. groups found in the Mid Atlantic Bight of Mannar (Fig. 12, 14) (James and 170°WÐ100°W), and (Winn et al., 1987; Waring et al., 1998). Soundararajan, 1979; Gordon, 1991; Division 9 Eastern Pacific Waring et al. (1993) reported signifi- Eyre, 1995). These concentrations are (long. 100°WÐ60°W). cantly more sperm whales in areas usually females and immature whales, where warm core rings of the Gulf male sightings being rare in the northern Male sperm whales are widely dis- Stream interact with continental shelf- Indian Ocean (Gordon, 1991). Stranding persed along the Antarctic ice edge from edge bathymetric features than where reports are uncommon along the Indian December to March (austral summer) coast for all whale species; the five sperm (Gosho et al., 1984). In contrast, mixed whale strandings reported between 1748 groups of females and immature whales 90 Mullins, J., H. Whitehead, and L. S. Weilgart. and 1980, were all males (James and have a southern limit in the South At- In press. Behaviour and vocalizations of two single sperm whales, Physeter macrocephalus, Soundararajan, 1979). lantic of lat. 50Ð54°S (Gosho et al., off Nova Scotia, 15 p. Can. J. Fish. Aquat. Sci. 1984; Tynan, 1998). Southern Hemisphere 91 Fritts, T. H. 1983. Turtles, birds, and mam- The Indian Ocean Sanctuary was cre- mals in the northern Gulf of Mexico and nearby Atlantic waters. Rep. prep. for U.S. Dep. Inter., For the purposes of worldwide popu- ated in 1979, under Article v(1)(c) of FWS/OBS-82/65, 455 p. lation assessment and management, the the ICRW, and all commercial whaling 92 Hansen, L. J., K. D. Mullin, and C. L. Roden. IWC recognizes the area south of the was prohibited within its boundaries. 1995. Estimates of cetacean abundance in the Equator as one biogeographical region, This boundary extends from the Antarc- northern Gulf of Mexico from vessel surveys. ° Contrib. MIA-94/95-25, NMFS Southeast Fish- the Southern Hemisphere. The Com- tic continent to lat. 55 S and from long. eries Science Center, Miami, Fla., 9 p. mission has divided this circumpolar 20°E to long. 130°E. In the western In-

62 Marine Fisheries Review dian Ocean (Division 3), there is evi- Table 17.—Recent abundance estimates for North Atlantic sperm whales. dence that concentrations of mixed fe- Population Coefficient of 1 male/immature whale groups exist Area estimate variation Source south of the Seychelles (Fig. 12) (James Western North Atlantic U.S. EEZ2 (1991) 337 0.50 Blaylock et al., 1995 and Soundararajan, 1979; Kasuya and Summer 1995 2,698 0.67 Waring et al., 1998 Wada, 1991; Kahn et al., 1993; Eyre, Spring and summer 1982 219 0.36 CeTAP70 Northern Gulf of Mexico 1995). In the central Indian Ocean (Di- 92 vision 4), concentrations of sperm Spring and summer 1991Ð94 530 0.31 Hansen et al. whales have been recorded to the north Eastern North Atlantic Iceland 1,234 0.17 Gunnlaugsson and Sigurjonsson, 1990 of St. Paul and Amsterdam Islands in Faroe Islands 308 0.38 Gunnlaugsson and Sigurjonsson, 1990 the austral summer (Fig. 14) (Gosho et Norwegian Sea 5,231 0.31 Christensen et al., 1992a al., 1984). Northern Norway to Spitsbergen 2,548 0.27 ¯ien, 1990 Rice (1977b), Wade and Gerrodette 1 Source footnote numbers refer to text footnote numbers. (1993), and Dufault and Whitehead 2 U.S. EEZ = United States Exclusive Economic Zone (200 n.mi from nearest land). (1995) suggested that a separate equatorial Pacific sperm whale population exists. Photo-identification studies off line transect data from 1991 to 1993 and eral seasons, are much smaller than those the Galapagos Islands and mainland 1996 ship-based surveys off California, summarized by Gosho et al. (1984). The Ecuador and North Peru indicate that Oregon, and Washington, Barlow74 es- small sample sizes, limited coverage, and there may also be a geographical separa- timated a weighted average of 1,191 probable undercounting of whales in the tion between Galapagos and Ecuador/ (CV = 0.22) and a minimum popula- areas result in statistical biases within 93 North Peru whales, although their genetic tion estimate (Nmin) of 995 sperm these estimates (Barlow and Sexton ). discreteness has yet to be verified (Fig. whales for those years. Historical abundance estimates for 13) (Dufault and Whitehead, 1995). From a ship-based line transect sur- Icelandic, Azorean, and Spanish North vey in the eastern tropical Pacific be- Atlantic stocks are provided in Table 4. Current and Historical Abundance tween lat. 10°N and 10°S, Wade and An initial abundance estimate of North Pacific Gerrodette (1993) provided a sperm 224,800 animals was calculated for all whale abundance estimate of 22,700 North Atlantic stocks in 1905 (Gosho All current abundance estimates for (CV = 0.22, 95% C.I. 14,800Ð34,600); et al., 1984), although this estimate is the number of sperm whales in the en- although, as noted earlier, this estimate no longer considered reliable by the tire North Pacific are considered unre- of abundance may include animals from IWC Scientific Committee. liable. As a result, caution should be more than one stock. No current abun- used when interpreting published esti- Northern Indian Ocean dance estimates are available for the mates. Rice (1989) estimated 930,000 entire western North Pacific. There are no current population abun- (no CV) sperm whales in the North Pa- Historical abundance estimates for dance estimates available for the north- cific Ocean. Gosho et al. (1984) esti- the western and eastern North Pacific ern half of the Indian Ocean. mated a total North Pacific population stocks are provided in Table 4. An abun- of 472,100 (no CV). Both of these esti- dance estimate of 620,400 animals was Southern Hemisphere mates are statistically unreliable be- calculated for the entire North Pacific in The current estimate of 299,400 (no cause they are based primarily on 22 1910 (Gosho et al., 1984), although this CV) sperm whales from the Equator to CPUE data collected during whaling estimate is no longer considered reliable lat. 70°S dates from 1977 and is statis- operations and do not take into account by the IWC Scientific Committee. tically unreliable (IWC, 1988). This any possible decline in abundance that estimate was calculated on the basis of occurred after whaling ceased (IWC, North Atlantic historical whaling records and CPUE22 1988). Based on historical whaling records data from whaling operations (Odell, Barlow (1994a) provided a current and CPUE22 data from modern whal- 1992). sperm whale abundance estimate for ing operations, there are an estimated Utilizing JSV and IWC/IDCR survey U.S. waters off central California of 756 190,000 (no CV) sperm whales inhab- data, Butterworth et al.47 estimated individuals (CV = 0.49, 95% C.I. 303Ð iting the entire North Atlantic (Odell, sperm whale abundances south of lat. 1,886). Limited trend data from 1979 1992). However, this estimate is con- 60°S (3,200Ð14,000; CV = 0.39Ð0.19) to 1991 showed a relatively stable abun- sidered unreliable by the IWC’s Scien- and south of lat. 30°S (128,000Ð290,000; dance of sperm whales in California tific Commitee (IWC, 1988). coastal waters (Barlow, 1994b). Pre- Recent regional abundance estimates liminary results from the winter of 1997 93 Barlow, J., and S. Sexton. 1996. The effect of have been calculated for both the west- SWAPS survey indicated that the esti- diving and searching behavior on the probability ern and eastern North Atlantic (Table of detecting track-line groups, g0, of long-diving mate of 756 whales may have actually 17). These numbers, from ship-based whales during line-transect surveys. NMFS been an overestimate (Taylor88). Using Southwest Fisheries Science Center, La Jolla, and aerial surveys conducted over sev- Calif., Admin Rep. LJ-96-14.

61(1), 1999 63 CV = 0.44Ð0.46), respectively (Table 18). of oil produced per whale rather than males) taken from the Bering Sea (Fig. Given the Antarctic latitudes surveyed, the actual number of whales caught. 38). Although there were no estimates these numbers most likely represent a Extrapolation from these types of data of whales caught in this region, a sig- large proportion of male whales. has led to only rough estimates of the nificant decline in CPUE22 north of lat. In South Pacific waters, Childerhouse number of whales killed per year 50°N led Kasuya (1991) to conclude et al. (1995) determined, using photo- (Gosho et al., 1984). In addition, newly that the Bering Sea sperm whale popu- identification and an “open” mark-re- revealed Soviet whaling catch data from lation had been greatly depleted. capture model, that between 60 and 180 Southern Hemisphere factory ships in- North Atlantic (no CV) male sperm whales occur off dicate considerable underreporting of Kaikoura, New Zealand (Division 7), sperm whale catches (Zemsky et al., In the North Atlantic, the hunting of each winter. In the equatorial Pacific, 1995; Zemsky et al., 1996). According sperm whales occurred off the west the total population of sperm whales to these “new” catch data, approxi- coast of Iceland, Norway (coastal and between the Galapagos and Ecuador mately 14,700 harvested sperm whales pelagic), the Faeroe Islands, Britain and North Peru was estimated at 3,891 went unreported in the original Soviet (coastal), West Greenland, Nova Scotia, (95% C.I. 2,600Ð5,300) (Whitehead et catch data between 1947 and 1987. As Newfoundland/Labrador, New En- al., 1992). more of these Soviet data are made gland, the Azores, Madeira, Spain, and Historical abundance estimates for available, catch-based population esti- Spanish Morocco (Waring et al., 1998) the nine Southern Hemisphere divisions mates will need to be revised. (Fig. 6, 7). Some whales were taken off are provided in Table 19. An abundance the U.S. Mid Atlantic coast, although estimate for the year 1946 of 547,600 Historic Exploitation Patterns the number of whales actually caught sperm whales (no CV) for the entire North Pacific is unclear in the literature (Townsend, Southern Hemisphere was calculated 1935; Reeves and Mitchell, 1988). from these division-based estimates. All Large-scale pelagic whaling in the Commercial whaling operations for North Pacific Ocean ceased in 1980, but of these estimates are statistically un- sperm whales were also conducted in U.S. fleets found few whales and there- reliable due to their use of historical the northern Gulf of Mexico during the whaling catch data and CPUE22 from fore had stopped whaling by 1979 late 1700’s to the early 1900’s (Townsend, (T¿nnessen and Johnsen, 1982). In whaling operations. It is important to 1935). There are no catch estimates avail- 1988, the IWC banned the killing of note that sperm whale catches from the able for the number of sperm whales early 19th century through the early sperm whales. Table 20 summarizes caught during the U.S. operations. The current estimates of sperm whale 20th century were calculated on barrels numbers caught in Norway and off catches from the North Pacific stocks Canada are summarized in Table 20. by whaling operations from 1911 to 94 Table 18.—Ship-based line transect abundance esti- 1987 . It must be noted that these num- Southern Hemisphere mates of Antarctic circumpolar sperm whales (Butter- bers, especially those from Japanese worth et al., text footnote 47). The average annual catch of sperm whaling operations, may lead to under- whales in the Southern Hemisphere Population estimation of historic abundance due to Area by year estimates CV from 1956 to 1976 was over 20,000 underreporting to the IWC (Kasuya and whales. Gosho et al. (1984) provided South of lat. 60°S Miyashita, 1988). The total estimated 1978/79Ð1983/84 113,200 0.39 summaries of those and worldwide post World War II take from the entire 1985/86Ð1990/91 114,000 0.19 sperm whale catch levels. South of lat. 30°S North Pacific is 258,000 animals. Not 1965/66Ð1977/78 290,0001 0.46 specifically indicated in Table 20, but Current Exploitation 1978/79Ð1987/88 128,0001 0.44 of significance to North Pacific stocks, 1 IWC worldwide catch limits (quotas) Extrapolated from Japanese Survey Vessel (JSV) data are the number of whales (mostly (Jan.ÐFeb. only). for all sperm whale stocks are currently 94 Based on barrels of oil produced per whale. set at zero (IWC, 1995b). As a result of

Table 19.—Estimated historical abundance of sperm whales in the Southern Hemisphere for the year 1946 Table 20.—Recorded sperm whale catch numbers from North Pacific and North Atlantic whaling operations. (Gosho et al., 1984) based on 1978 and 1980 IWC catch data. Area Years No. of animals Source

IWC stock division Population estimate North Pacific Western North Pacific (Kurils, Hokkaido, 1 32,700 Sanriku, central & south Japan) 1911Ð45 19,989 Kasuya, 1991 2 72,100 Eastern North Pacific pre-WWII 3,699 Kasuya, 1991 3 80,700 Total from Japanese operations 1955Ð86 62,033 Kasuya and Miyashita, 1988 4 49,700 Total North Pacific 1947Ð87 258,000 Barlow et al., 1995b 5 49,600 6 29,800 North Atlantic 7 42,000 Western Norway 1925Ð69 374 Christensen et al., 1992a 8 96,200 Western Norway 1948Ð71 1,088 Christensen et al., 1992a 9 94,800 Newfoundland/Labrador 1904Ð72 424 Blaylock et al., 1995 Total 547,600 Nova Scotia 1964Ð72 109 Blaylock et al., 1995

64 Marine Fisheries Review Figure 38.—Three sperm whales on a flensing platform in Alaska. The right animal’s spermaceti organ has been removed. Uni- versity of Washington Special Collections, Lagen Collection, negative UW17505. this prohibition on whaling, the number of reported sperm whale kills for either commercial or aboriginal harvest has been zero in recent years (IWC, 1995a). The only known subsistence harvest of sperm whales occurs in Lomlem, Indonesia, where a few whales are taken per year using primitive methods (Barnes, 1991). Today the threat of commercial overharvest is greatly reduced compared to the potentially more long-term threats of habitat degradation, marine pollution, human exploitation of potential prey, and commercial fisheries interactions.

Life History and Ecology Feeding In general, the sperm whale’s primary prey consists of larger mesopelagic cephalopod and fish species, including the giant squid, Architeuthis sp. Ap- proximately 40 species of cephalopods are consumed by sperm whales worldwide. In the North Pacific, the four most Figure 39.—Showing its square-shaped head, a sperm whale breaches in the dis- common prey items of sperm whales off tance. S. Leatherwood, NMML Collection. central California are all cephalopod species (i.e. Moroteuthis, Gonatopsis, et al., 1989). In the Indian Ocean, the eaten by sperm whales are of the Histio- Histioteuthis, and Galiteuthis) (Fiscus cephalopod species most commonly teuthid family (Gordon, 1991). Sperm

61(1), 1999 65 whales in the high latitudes of the North rate of 20% in unexploited populations Human-related Mortality Atlantic (i.e. Norwegian Sea and Ice- to 25% in exploited populations (IWC, Fisheries Interactions land) feed on deep-dwelling fish spe- 1980c). cies of the genus Cyclopterus (lump- In U.S. waters of the Pacific, inciden- suckers) and Sebastes (redfishes). Fish Natural Mortality tal take of sperm whales has been docu- prey comprises almost half of the total Serological studies on North Pacific mented in drift gillnet operations. The biomass eaten by sperm whales in this and North Atlantic sperm whales indi- average annual rate of mortality and region, while the other half is comprised cate that these whales are carriers of and serious injury from the offshore Cali- of cephalopods (Martin and Clarke, are infected by calciviruses and papil- fornia drift gillnet fishery from 1991 to 1986; Christensen et al., 1992b). lomavirus (Smith and Latham, 1978; 1995 is nine sperm whales (Barlow et Lambertsen et al., 1987). For example, al., 1997). Observers aboard Alaska Reproduction there was evidence of papillomavirus- sablefish, Anoplopoma fimbria; and Pa- One of the most recognizable features associated disease in 10% of a popula- cific halibut, Hippoglossus stenolepis, of sperm whale social structure is the tion sample taken from Iceland (Lam- longline vessels have documented sperm “nursery school” that contains between bertsen et al., 1987). whales feeding on longline-caught fish in 20 and 30 individuals, including fe- Killer whales, false killer whales, the Gulf of Alaska (Hill and Mitchell96). males, calves, and juveniles (Whitehead Pseudorca crassidens; and short-finned In 1997, the first entanglement of a sperm and Arnbom, 1987; Whitehead, 1996; pilot whales, Globicephala melana, whale in Alaska’s longline fishery was Richard et al., 1996). At around 6 years have all been observed in what appears recorded, although the whale was not se- of age, juvenile males (and possibly fe- to be harassment of sperm whale groups riously injured (Hill and DeMaster97). males) leave these nursery schools to (Arnbom et al., 1987; Palacios and There is no evidence that mortality or se- form juvenile or bachelor schools (Ri- Mate, 1996; Weller et al., 1996). Sperm rious injury occurs as a result of interac- chard et al., 1996). Juvenile schools whale defensive maneuvers in these in- tions with this fishery; however, the na- contain individuals between 4 and 20 teractions seem to be based on protec- ture and extent of these longline fishery- years of age and are less cohesive and tion of the youngest and most vulner- sperm whale interactions is not yet clear. smaller than the nursery schools (White- able of the group (Nishiwaki, 1962; The first observed incidental take of head and Arnbom, 1987). By the time Weller et al., 1996). Bleeding wounds a sperm whale in U.S. waters of the males reach 30 years of age, they are have been observed on sperm whale North Atlantic was in 1989 in a drift mostly solitary. heads and tail flukes after such events gillnet. The estimated fishery-related Breeding among sperm whales takes (Arnbom et al., 1987; Dufault and mortality and serious injury rate was 2.2 place in spring and early summer in Whitehead, 1995). The most recent sperm whales in 1989, 4.4 in 1990, and both hemispheres (from April to August documented incident of killer whales zero from 1991 to 1996 (Waring et al., in the Northern Hemisphere and from attacking sperm whales occurred off 1998), although in 1995 one sperm October to February in the Southern Point Conception, Calif., in October whale was observed entangled in a pe- Hemisphere). Pairings take place in the 1997 (Roberts95). During the attack, lagic drift gillnet. This entangled whale lower latitudes, where males join nurs- approximately 25 killer whales mortally was released alive, although gear re- ery schools for days at a time. However, wounded at least one of nine sperm mained wrapped around several body photo-identification studies suggest that whales. The incident was unusual because parts. Waring et al. (1998) describe three not all males breed each year (Martin, it apparently involved only large, mature known instances of sperm whale en- 1980; Whitehead and Arnbom, 1987). sperm whales. The attacked whales tanglements in the northwest Atlantic; Females reach sexual maturity at 9 showed no defensive actions except in two in net gear and one in longline gear. years of age and a length of approxi- protecting injured members of their pod. There is little information available mately 9 m. The calving interval is one Estimated natural mortality rates for regarding fishery interactions with of the longest for mammals, between sperm whales are age-specific. Juve- sperm whales outside U.S. waters. 4.8 and 6 years (Kasuya, 1991). Gesta- niles (age 0Ð2) have an estimated an- However, behavior similar to that ob- tion lasts from 14 to 16 months, and lac- nual mortality of 0.09, while mature served in the Alaska longline fishery has tation is between 1 and 2 years (Kasuya, (age 2 and above) whales have an esti- also been documented during longline 1991). Males reach sexual maturity at 20 mated annual mortality of 0.05 (IWC, operations off South America (South years of age and a length of 12 m. At 1971). Because of the lack of informa- Georgia, Kerguelen, and southern around 30 years of age, males reach “so- tion on the causes of natural mortality cial” maturity and begin living the previ- in sperm whales, these rates are no 96 Hill, P. S., and E. Mitchell. 1998. Sperm whale ously mentioned solitary life. longer considered statistically reliable interactions with longline vessels in Alaska waters during 1997. Unpubl. rep. presented at 6th Pregnancy rates vary between ex- by the IWC (IWC, 1980c). Pacific Scientific Review Group Meeting, March ploited and unexploited stocks. Exploi- 30, 1998, Honolulu, Hawaii. Avail. from National tation of sperm whale stocks may have 95 Roberts, K. 1998. NOAA Corps Officer, Pa- Marine Mammal Laboratory, 7600 Sand Point cific Marine Environmental Laboratory-EDD, Way, N.E., Seattle, WA 98115. caused a density-dependent response, 7600 Sand Point Way, N.E., Seattle, WA 98115. 97 Citation updated in proof: see Hill and DeMaster, which increased the average pregnancy Personal commun. 1999 in literature cited.

66 Marine Fisheries Review Table 21.—Factors possibly influencing the recovery of sperm whale stocks under the ESA (1973) ¤4(a)(1)1992 Amend.

Factor North Pacific North Atlantic Gulf of Mexico Indian Ocean Southern Hemisphere

1.Present or threatened Pollution Pollution (e.g. plastics, heavy Oil and gas development (e.g. Pollution Pollution destruction or modifica- metals) noise disturbance, oil spills) tion of habitat

2. Overutilization for com- Unknown Whale watching and associ- Scientific research and asso- Unknown Whale watching, scientific re- mercial, recreational, ated vessel traffic ciated vessel traffic search, photography, and as- scientific, or educational sociated vessel traffic purposes

3. Disease or predation Papilloma and calcivirus; Papilloma and calcivirus Papilloma and calcivirus; Orci- Papilloma and Orcinus and Pseudorca attacks Orcinus attacks nus, Pseudorca, and Globi- calcivirus cephala attacks

4.Other natural or man- Entanglement in fishing gear Entanglement in fishing gear Unknown Unknown Entanglement in fishing gear made factors (e.g. longline, drift gillnets) (e.g. drift gillnets) (e.g. longline)

Chile) where sperm whales have become et al., 1992). It is unknown whether an- enous and widespread distribution, entangled in longline gear, have been ob- thropogenic noise has biological signifi- which is apparently gender- and age- served feeding on fish caught in the gear, cance for sperm whales. related. Table 21 summarizes informa- and have been reported following longline tion on potential threats affecting the vessels for days (CCAMLR, 1994; Pollution status of sperm whales. Therefore, any Ashford et al., 1996; Capdeville, 1997). Relatively high mercury levels have reevaluation of sperm whale classifica- These observations, combined with an- been found in breeding females cap- tion status awaits the collection of more ecdotal reports suggest that interactions tured off southern Australia. It is unclear reliable information on distribution, between sperm whales and longline op- whether these mercury levels affect the migration patterns, abundance, and erations may be widespread in the waters whale’s health (Cannella and Kitchener, trends in abundance on a stock-specific off South America (Hill and Mitchell96). 1992). Plastic debris is probably in- basis, as well as the development of gested quite frequently by sperm whales objective delisting criteria. Nonetheless, Noise Disturbance at sea. For example, a 15 m male sperm if the accuracy of abundance estimates In recent years, many studies on the whale captured in nearshore waters off and stock determinations for North At- effect of noise on the behavior of whales Iceland had a 3-gal plastic bucket lantic and North Pacific sperm whale have been done (Richardson et al., lodged in his intestinal tract (Lambert- populations can be made more reliable 1995). A resident population of sperm sen and Kohn, 1987). with additional survey data, and if hu- whales occurs in the northern Gulf of man-related sources of mortality and Classification Status Mexico, an area of intensive oil and gas serious injury remain low, some stocks exploration and development activities. The sperm whale was listed as en- might be candidates for downlisting. Oil production platforms and their as- dangered under the ESA in 1973 and is Acknowledgments sociated vessels have unknown effects protected under the MMPA. Endan- on sperm whales (Odell, 1992). Stud- gered status is applied to all sperm Thanks to those who took the time to ies of whale reactions to seismic sur- whale stocks utilizing U.S. waters comment on earlier drafts (Willis veys in the Gulf of Mexico indicated (Anonymous, 1994b). The western Hobart, Jim Carretta, Kim Shelden, that sperm whales reacted to seismic North Pacific stock is the only sperm Leah Gerber, Paul Wade, Michael pulses by moving away 50 km or more whale stock designated as a “Protected Payne, Dave Rugh, and Scott Hill) and (Mate et al., 1994). In the southern In- Stock” by the IWC. Under this desig- to those who commented on later drafts dian Ocean, most sperm whales stopped nation, the IWC recognizes that these (Phillip Clapham, Jay Barlow, Robert vocalizing when exposed to seismic whales are 10% or more below their Brownell, Kevin Chu, Katherine Wang, pulses as much as 300 km away (Bowles maximum sustainable yield (MSY) and Jeffrey Breiwick). Thanks also go et al., 1994). Sperm whales have also level, or 54% of carrying capacity (K) to Kristin Laidre for her attention to been observed exhibiting startle re- (IWC, 1995b). Although without trend detail and map-making skills. And, sponses to a closely approaching ves- data or information on status relative to many thanks to Pieter Folkens for per- sel (Whitehead et al., 1990). Observed K for this stock, the validity of this des- mission to use his illustrations. This reactions of sperm whales in the pres- ignation is questionable. manuscript was supported in part by ence of vessels include more erratic Since Braham’s 1991 status review3, the NMFS National Marine Mammal surface movements, reduced surface there has been little new information to Laboratory (Contracts 40ABNF80 time, fewer blows per surfacing, shorter improve the accuracy of population es- 0705, 40ABNF701925, and 43ABNF intervals between successive blows, and timates or stock identity. One of the 701085) and through ESA/MMPA increased frequency of dives without major difficulties in identifying distinct funds from the NMFS Office of Pro- raised flukes (Cawthorn, 1992; Gordon sperm whale stocks is their heterog- tected Resources.

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