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No. 304, pp. 1-9, 4 figs.
By John E. Heyning and Marilyn E. Dahlheim
Published 15 January 1988 by The American Society of Mammalogists Orcinus Fitzinger, 1860
There are no comprehensive comparative accounts describing the skulls of large delphinids. The large skull size (condylbasallength to 100 em), the dental formula (10 to 1411 0 to 14) and large teeth distinguish O. orca skulls from those of other species, except large false killer whale (Pseudorca crassidens) skulls (dental formula 8 to 11/8 to 11; Leatherwood et aI., 1982). Skulls of O. orca can be distinguished from those of P. crassidens by the width across the premaxillaries being less than 50% of rostral width just anterior of antorbital notches and the lateral border of the premaxillaries being slightly more sigmoid in dorsal view and wider distally (Fig. 2). Pterygoids widely separated and teeth compressed anteroposteriorly at the roots are two characteristics often cited as diagnostic for O. orca (Glass, 1974), but do not always separate 0. orca and P. crassidens. The tympanic and periotic bone complex of O. orca (Fig. 3)
Orca Gray, 1846. Generic name preoccupied by Orca Wagler, 1830 (=Hyperoodon). Orcinus Fitzinger, 1860. Type species Delphinus orca Linnaeus, 1758. Ophysia Gray, 1868. Type species Orca capensis (=Delphinus orca). Gladiator Gray, 1870. Type species Orca stenorhyncha t=Delphinus orca). Grampus Iredale and Troughton, 1933 (not Gray). Type species Delphinus grampus (=Delphinus orca). CONTEXT AND CONTENT. Order Cetacea, Suborder Odontoceti (considered an order by some authors), Superfamily Delphinoidea, Family Delphinidae. The genus Orcinus has been placed in the subfamily Orcininae by several workers. This taxon is not entirely stable in its membership because, in addition to Orcinus, it may include Pseudorca, Globicephala, Orcaella, and Feresa (Fraser and Purves, 1960; Mead, 1975; Slipjer, 1936) or just Pseudorca (Kasuya, 1973).
Orcinus orca (Linnaeus, 1758)
[Delphinus] orca Linnaeus, 1758:77. Type locality European seas. Delphinus serra Borowski, 1780:38. Type locality Spitzbergen. IX"elphinus] Gladiator Bonnaterre, 1789:23. Type locality Spitz. bergen. Delphinus Duhameli Lacepede, 1804:314. Type is description of animal from France. Delphinus grampus Blainville, 1817:168. Type locality North Atlantic. Orca Capensis Gray, 1846:34. Type locality Cape of Good Hope. Delphinus victorini Grill, 1858:21. Type locality Capetown, South Africa. Clrca] Schlegelii Lilljeborg, 1866:235. Type locality Norway. Orca magellanica Burmeister, 1866:99. Type locality south of Buenos Aires, Argentina. Orca Eschrichtii Reinhardt in Eschricht, 1866: 188. Type locality Faeroe Islands. Orca ater Cope in Scammon, 1869:22. Type based on description of animals from Oregon to Aleutian Islands. Orca rectipinna Cope in Scammon, 1869:22. Type based on description of animals from California. Orca stenorhyncha Gray, 1870:71. Type locality English coast. Orca latirostris Gray, 1870:76. Type locality coast of Essex, North Sea. Ophysia pacifica Gray, 1870:71. Type locality North Pacific? O[rca] pacifica Gray, 1870:394. Type locality coast of Chile. Orca africana Gray, 1871:91. Type locality Cape of Good Hope. Orca tasmanica Gray, 1871:92. Type locality Tasmania. Orca minor Maim, 1871:81. Type locality Sweden. Orca antarctica Fischer, 1876:146. Based on drawing. Orcinus nanus Mikhalev et aI., 1981:563. Antarctic waters. Orcinus glacialis Berzin and Vladimirov, 1983:288. Type locality Indian Ocean sector of Antarctic. CONTEXT AND CONTENT. Context as above. There are no widely accepted subspecies of O. orca. DIAGNOSIS. Orcinus orca is the largest member of the family Delphinidae, with adults ranging in length from 5 to 9 m. The large size, robust body, postocular white spot, and ovate flippers are diagnostic (Fig. 1). Killer whales have proportionately higher dorsal fins than other large delphinids, ranging from one- tenth to one- fifth of the total body length.
FIG. 1. Lateral and ventral views of a typical adult male (A, B) and female (C, D) of Orcinus orca. Illustration by Robin AgoesMakowski.
usually jet black. closure of the elliptical foramen. In some areas they occur seasonally. and off Japan (Kasuya. Killer whales are one of the most strikingly pigmented cetaceans (Fig. juveniles tend to be more yellowish than adults.000 kg for a 6. Mitchell. 1984). 1957). 1982). 1978). Among adult males.04 m male (Hoyt. Orcinus occurs frequently along the Soviet coast in the Bering Sea. 1970) that expands dorsoposteriorly above the urogenital region. Upper: dorsal and ventral views of the tympanic bulla. sightings have been noted in the Labrador Sea. There are several records of O. 1874. Scheffer and Slipp. Although reported from tropical waters and the open ocean. Height of the dorsal fin is a useful character for distinguishing adult males within pods. 1976). 1948). Light-colored areas often are yellowish.5 m for females and 9. constricting medially between the flippers. in the Barents and White seas. 1978) and off South Africa . Sexes of free-living adult females and subadult males are difficult to distinguish.2 MAMMALIAN SPECIES 304 FIG. 1) making field identification easy. In the western North Atlantic. Few animals have been weighed but a maximum recorded mass was 3.35·m female and 4. 1974. Killer whales are robust. Continuous with the ventral white area is a lateral flank patch (Mitchell. orca has been observed off Greenland.8 m in height. Evans et aI. Iceland. they seem to be most numerous in colder waters of both hemispheres. Animals seem to be abundant off the coast of Alaska with a population estimate of 286 for the Prince William Sound and southeast waters (Leath. The head is blunt with virtually no distinguishable beak. Great Britain. Sightings decrease southward along the continental United States (True.. Female killer whales generally attain a body length of 7. and Ireland (Fraser. There is a postocular white patch. the dorsal fin is triangular and may reach 1. 4. Sea of Okhotsk (Tomilin.0 m and males 8. with the greatest abundance within 800 km of major continents (Dahlheim. FIG. The large (both relatively and absolutely) ovate flippers are positioned about one. and lateral views of the skull and lateral view of mandible from a 484·cm male Orcinus orca (LACM 52480). 1982).100 kg for a 6. and massive anterior and posterior processes of the periotic (Kasuya. and off Novaya Zemlya (Tomilin. 1983. Dorsal. 1960. In the southern oceans. maximum lengths of 8. Among males.9 m tall and distinctly falcate. erwood et aI. 1975). however. Canada (Sergeant and Fisher. 1975). 1957). GENERAL CHARACTERS. 1946. Sightings near Hawaii are uncommon (Richards. Population estimates are not available for the remainder of the North Pacific. The ventral aspect of the flukes also are white or light gray. A highly variable gray or white saddle is usually present posterior to the dorsal fin.2 m (Mitchell. 1948) have been seen in the North Pacific. Harmer. There are no population estimates for the North Atlantic. 1874. whereas in adult females and young males it is less than 0. In the North Atlantic Ocean. In the western North Pacific. Scheffer and Slipp. then widening slightly and ending just beyond the urogenital region. The DISTRIBUTION. 1970). flippers may measure 2 m (Harmer. but in other areas they are apparently year-round residents. killer whales occur to the tip of Tierra del Fuego.. Killer whales have been observed in all oceans and seas of the world (Fig.fourth the distance from the snout to flukes and contrast with the sickle-shaped flippers of most delphinids. lower: dorsal and ventral views of the periotic of Orcinus orca (LACM 52480). but yellowish coloration also has been reported from the North Pacific (Scammon. ventral.8 m for males have been reported (Perrin and Reilly. Newfoundland. 1948) and partially albinistic animals (Carl.. 1982) and have been documented as far north as the Chukchi and Beaufort seas. 1927) and attain 20% of the body length. 1904). 1971). among females the flippers attain 11 to 13% of the body length (Eschricht. 1982. Leatherwood and Dahlheim. O. 1984). Year-round occurrence has been documented for the intracoastal waterways of British Columbia and Washington State where an estimated 260 whales comprising 30 pods have been reported (Bigg. 1957). white region typically extends from the entire lower jaw posteriorly. 3. 1952). 1927). In the northeastern Pacific Ocean. 2. Individual and geographical variation in the pigmentation pattern exists (Carl. 1971). 1981). 1866). is characterized by a lack of a ventral keel. Scheffer and Slipp. and off Nova Scotia. 1981. Regular occurrence also is documented off Norway (Ionsgard and Lyshoel. 1982). especially in animals from the Antarctic (Berzin and Vladimirov. dorsally with a well-demarked white venter.. 1973). Evans et aI. killer whales occur in the eastern Bering Sea (Braham and Dahlheim. Killer whales are dark. Reports from the Mediterranean are sporadic (Casinos and Veri cad. South America (Goodall. orca from Caribbean waters (Caldwell et aI. Melanistic individuals (Scammon.
The mass of the brain from a 521-cm (curvilinear) specimen was 4. Ends of phalanges and most carpal elements were composed of cartilage in an adult male examined by Eschricht (1866). orca are arranged in groups of four that are contiguous. Orcinus has been recorded as far south as the Ross Sea (Brown et aI. New Zealand (Baker.500 g (Caldwell and Brown.2 m long (Eschricht. An encephalization quotient of 2. 1967). The venous return from the kidney differs from that of Hyperoodon as the renicular vein receives directly the intrarenicular and centripetal tributaries and no peripheral venous plexus is formed. Nishiwaki's (1972) higher counts may include the morphologically similar metacarpals as the proximal phalanx of each digit. 1977). The reported vertebral count is 7 C. and true keratinization (Harrison and Thurley. Tomilin. The entire skeleton is basically built on the typical delphinid plan. The electrocardiogram of one captive animal showed no P wave. movements. and V 2. but is more robust in all aspects. III 3 to 4. Older animals can have extensive wear on the teeth (Caldwell and Brown. but reports decrease northward along the coasts of both continents.. In Antarctic waters. Wood and Evans. citoniensis has a slightly higher tooth count (14114) and proportionately smaller teeth than the extant species. 1974. IV 2 to 3. Shaded areas indicate records of sightings or strandings. Cave (1977) found that the reniculi of the kidney in O. 1974). 1982). The structure of the epidermis of several delphinids including O.MAMMALIAN SPECIES 304 3 FIG.. The intestines of one animal were 54. an inverted T wave. 1981). Harmer (1927) hypothesized that the accelerated secondary growth of the flippers in maturing males was related to the continued growth of these cartilages. 1974). Rib counts range from 11 to 13 per side (Eschricht..7 on the left (Ling. Human and bovine antibodies cross-react with similar pituitary hormones of Orcinus in- . 1969). The teeth are relatively large. Mean numbers of facial vibrissae (n = 7) were 3. Nishiwaki. Ridgway and Brownson (1984) estimated an average brain mass of 5. citoniensis (Capellini. and 20 to 24 Ca. Numerous large delphinoid teeth. indicating a large and powerful temporalis muscle for jaw closure. 1964). 1967). also has been reported from South Africa (Barnard. whereas Nishiwaki (1972) reported the count as I 2. II 7. The amino acid sequence of the myoglobin of O. II 4 to 6. The tongue reportedly is protrusible in contrast to the tongue in Tursiops truncatus (Donaldson. O. citoniensis (Lydekker. Ness (1967) stated that the degree of skull asymmetry in Orcinus is low compared with that of other large delphinids. details of distribution. 1954). 1957). primarily from the Pliocene. A dredged fossil jaw of Orcinus sp.. 1972) and their apices curve inward. orca lack the stratum granulosum and stratum lucidum layers.617 ± 968 g from animals averaging 555 ern in length. 10 to 12 L. It appears to be a smaller species than O. the mitotic division rate is rapid and is 290 times that of epidermis from the human forearm (Harrison and Thurley. the temporal fossa in noticeably large.6 on the right side and 3. 0. orca (Pilleri and Pilleri. 1866). 1964). The digestive system is similar to that of other delphinids. 1974). O. IV 4. may be part of the normal range with no sightings documented. total 50 to 54 (Eschricht. The 0. FORM. Despite numerous reports of killer whales in the Southern Hemisphere. but Wood and Evans (1980) believe this low number may be biased because of the large size of killer whales and their high body weight caused by blubber. 1981). Geographic distribution of Orcinus orca. This en cephalization quotient is relatively low for odontocetes. semseyi (Biickh. 1866) with the anterior six or seven having both capitular and tubercular attachments to the vertebrae and the remainder attached only by the tuberculum. 1980). orca facial anatomy differed little from the typical delphinid plan of asymmetrical nasal sacs except that several structures were proportionately smaller in comparison to 14 other species. killer whales are recorded from Australia (Bryden. They occur at wide intervals in the Indian Ocean. orca is more similar to that of Globicephala sp. and a simple R wave with a surface-breathing heart rate of 60 beats/min that declined to 30 beats/min when submerged (Spencer et aI. capacity of the blood is reported to be moderate for cetaceans in comparison with the much greater 0.2 I and a tidal flow of 1291/s were calculated for a 4. 1866. Ribs 1 through 6 attach directly to the sternum. orca with an estimated total length of less than 4 m. 1984). 1977).3-m female (Spencer et aI. have been attributed to Orcinus orca or a closely related species. III 5.9 for Orcinus has been estimated (Ridgway and Brownson. 1972). 1984. up to 13 cm in length (Nishiwaki. 1983). 1983) and off the Galapagos Islands (Robinson et aI. Such teeth have been reported from the Pliocene of Italy (Sarra. 11 to 13 T. capacity of Kogia blood (Lenfant. but possessing most of the postcranial skeleton. 1933) and Japan (Matsumoto.. in general. The specimen consists of a partial skull lacking the posterior and left side of the cranium. Because the epidermis is sloughed rapidly during swimming. 0. and abundance are not known (Dahlheim. FUNCTION. In the South Pacific Ocean. FOSSIL RECORD. 1978). 1899) from the Lower Miocene strata of Hungary actually is a physeterid. Both the mandibular and maxillary aveoli are deep. Eschricht (1866) listed the phalangeal formula as I 1. 1883) from the Pliocene of Italy. Mead (1975) found that. citoniensis seems to be intermediate between more typical delphinids and O. The teeth are oval in cross-section at the base. 4. 1887). In these characters. than to the myoglobin of other small delphinids and phocoenids examined (Meuth et aI. 1937). On the skull. A lung tidal volume of 46. An isolated tooth and periotic bone from the Suffork Crag of England has been referred to O. and V 3. One of the few fossils represented by a good skull is 0.. Their presence has been observed along the edge of the pack ice throughout Antarctic waters (Brown et aI. however unshaded areas (Ross.
1866. Trigonocotyle spasskyi.2 to 6. killer whales are known to prey upon other marine mammals and seabirds. Orcinus meat is used only for animal consumption (Jonsgard and Lyshoel. a few unsuccessful attacks have been reported (di Sciara.400 g (R) and 12. Some of this variation is geographic with northeastern Atlantic animals representing the low end and Antarctic animals the high end (Perrin and Reilly.. Gaskin. Estimates of the gestation period differ with 15 months the best current estimate (Perrin and Reilly. Killer whales are relatively free of external parasites but barnacles (Xenobalanus and unidentified) have been observed on the rostrum and trailing edge of the flukes (LACM No. and 9% with squid (Ivashin. Harrison et al. 1982. 1982). Killer whale oil in Japan is equal in price to sperm whale (Physeter catodon) oil. and 227 ern for the Southern Hemisphere (Ross. 1970). 1984). orca was captured in this region. 1981). Eschricht (1866: 159) stated that partially digested remains of these animals were found and later these were misinterpreted as whole animals. 1984). in order of occurrence. The average size of males at sexual maturity ranges from 5. but old meat and viscera are used as fertilizer or bait (Nishiwaki and Handa. orca can be found in Hoyt (1981). 1872). Anasakis simplex (Dailey and Brownell. ECOLOGY. 1958). The size of ovaries from mature killer whales are about 10 to 12 cm by 5 to 7 ern (Mikhalev et al. Breeding cycles seem not to be fixed worldwide with mating and calving seasons often spanning several months. Pliyilobothrium sp. and 250 em for the Antarctic (Mikhalev et al.4 dicating the presence of both lactotrophs and somatotrophs (Schneyer and Odell.. fish. The fresh meat of Orcinus is used for human consumption in Japan. only one live O. Estimates of annual pregnancy rates range from 13. The following endoparasites are known from O. 1972). squid.1 m. 55 were kept for public display in oceanaria.2 to 0.9. The maximum size of fetuses differs regionally and has been documented as 255 cm for the North Atlantic (Perrin and Reilly. The most common disease reported for killer whales is caused by the wearing of teeth resulting in exposure of the pulp cavity.4 (Nishiwaki and Handa.2 m. Off Japan. 9% pinnipeds. Jonsgard. 1968. An examination of 57 mature males from the Antarctic indicated an average testis length of 55 ern and a width of 22 ern.4%).83: 1 for the northeast Pacific (Balcomb et al. Eschricht.7 to 39. killer whales have been hunted for oil and meat.270 g (L) was not mature. 1982).2% with the lower estimates probably more reliable. Killer whales have no significant predators other than man. Useful tables of data on captive O. 1973). 1874. The ectoparasite Cyamus orcini has been described from killer whales (Leung. and miscellaneous (Nishiwaki and Handa. 1981). The average testis mass was calculated at 10. Simpson and Gardner (1972) reported frequent abscessed follicles of the vestigial hair on the rostrum of captive animals that eventually spread over the entire skin surface. There are photographs of a remora (Echeneididae) attached to a killer whale (Lockyer. Attacks and predation on large whales has given killer whales their vernacular name and incidents are well documented (Andrews. Before this. 1981). 1958) with lactation lasting 12 months (Bryden. cetaceans.4 m (Perrin and Reilly. 1984). The only killer whales to breed in captivity produced calves 19 months apart (Hoyt. 1894.48: 1 and 0. Fasciola skriabini. ONTOGENY AND REPRODUCTION. The life span of killer whales was estimated to be 25 years (Jonsgard and Oynes.5 to 6. orca has been described for several fetuses (Guldberg and Nansen. but may be approximately 5% per year for the total population based on information concerning life spans. An adolescent growth spurt is reported in males from 5. orca as between 750 and 950 kg per animal and Eschricht (1866) noted that 205 I was yielded from a 6. 1980). 1984). or killed as a potential competitor by fishermen (Dahlheim. However. Turner. and an iodine value of 86.632 g (R) and 2. (1972) determined from histological examination that a 656-cm individual with testes masses of 3. but may be as long as 35 to 40 years (Mitchell and Baker. 1978). Tomilin (1957) listed the oil yield for O. One often cited misconception is that one O. Kasuya (1971) estimated a mean pod size of 6. and there may be some social control of reproduction. The smallest neonates recorded are 183 em for the North Atlantic. 1981). Distinct layering of dentine is present in tooth sections from 0. some evidence suggests that the birth rate may be density dependent (Fowler. and Nematoda. 1984). Beginning in 1976. Sergeant and Fisher (1957) believed movements off eastern Canada were associated with migrations of seals and rorquals. the growth curves from males and females are identical. and Hoyt (1981). movements of killer whales seem to be related to movements of their food supply. The age of most odontocetes is determined by sectioning teeth and counting the dentine or cementum layers. Tarpy.. The natural mortality rate of killer whales is not known. systemic mycosis (22%).. 1965).000 g with a maximum mass of 23.4 ng/ml has been recorded for one female (Kirby and Ridgway. 1952). 1984). Detailed embryology and placenta morphology of O.. 228 ern for the North Pacific (Perrin and Reilly. Sex ratios at birth appear to be 1: 1. orca was found with 13 whole porpoises and 14 seals in its stomach.6 to 5. In the warm temperate and tropical eastern Pacific. Hancock. Jonsgard and Lyshoel (1970) concluded that the distribution and migration of killer whales in the northeastern Atlantic seem to be dependent upon migration of herring. 1980). and a range of 1 to 30. (1984) found high levels of PCBs and DDT in the tissues of killer whales from Washington State waters.63. estimates of annual birth rate range from 4 to 5% (Dahlheim.4-m animal. Although there are no BEHAVIOR_ Killer whales usually occur in small pods of fewer than 40 individuals.6%). 1977).6 by 5. Infection may penetrate through the pulp cavity causing a jaw abscess. 1958). 1968. but numerous accessory layers make them difficult to interpret (Perrin and Myrick. 1935). 72550).1 cm (Turner. Bigg. Dahlheim et al. Scammon. 1970). Useful summary tables of Orcinus stomach contents by geographical region are provided in Anon. 1970). The cause of death for captive killer whales (n = 32) was reported by Greenwood and Taylor (1985) as pneumonia (25%). calves may be dependent for at least 2 years. MAMMALIAN SPECIES 304 documented instances of killer whales eating man. 1958). of which 10 died. orca: Trematoda. Killer whales are top-level marine carnivores and opportunistic feeders with diets that differ seasonally and regionally. though Mitchell (1975) estimated that killer whales consume 4% of their body weight per day. In Norway. 1985). 1965. The rare genetic Chidiak-Higashi syndrome has been documented in a captive killer whale (Haley. with only 1% of these groups containing 20 or more individuals (Braham and Dahlheim.200 g (L) testes was sexually mature. 1985). Whitehead and Glass. 1979). 1984).' orca. Cestoda. Since 1976.. Rice. From 1962 to 1976.100 g (Mikhalev et al. and 237 were released (Asper and Cornell.. The daily food intake is unknown. Examination of contents from killer whale stomachs from the North Pacific yielded. and 1. Diatoms were reported on the skin of Antarctic animals (Hart. In various parts of the world. Calambokidis et al. In the northeast Atlantic. Bigg and Wolman (1975) provided data on the live-capture fishery in this area from 1962 to 1973 with slightly different numbers. A female with a 91-cm fetus had a corpus luteum measuring 7. 1981). Atherosclerosis was found in one stranded animal (Roberts et al. Estimates of calving intervals range from 3 to 8 years with observational data indicating that the higher estimates are more typical. the time of sexual maturity (Christensen. Of 362 stomach contents of O. Kasuya and Marsh. and undiagnosed (28%). By 1976.34: 1 for the Marion Islands (Condy et al. 1914. A progesterone level of 0. Iceland became active in the live-capture fishery (Greenwood and Taylor. orca examined from the Antarctic region. However. Weaning is thought to occur when a calf reaches a length of 4. 1984). In Alaskan waters. (1982). The relationship of dentine layers to absolute time for O. 1978). 1979. Effects of environmental pollutants are not known for cetaceans. Females attain sexual maturity between a length of 4. 1984). however.3 m (Nishiwaki and Handa. 1872). 1972). In most geographical regions. but overall ratio of males to females has been reported at 0. and a 724-cm animal with 11. 1958). Polygamy undoubtedly exists in killer whales. Primarily fish eaters. 1981). saponification value of 211. 60% contained only fish with 31 % also containing minke whale (Balaenoptera acutorostrata). pinnipeds. 1984. 1982. mating occurs from late autumn to midwinter (Jonsgard and Lyshoel. pod size ranged from 1 to 100 animals. Orcinus oil has an acid value of 0. orca is unknown at present. . 1981). 274 ern for the North Pacific (Nishiwaki and Handa. 1984). mediastinal abscess (9. 302 animals were captured. a live-capture fishery was active in Washington State and British Columbia. other bacterial infections (15. Gaskin (1982).
but a worldwide systematic review is needed. Haug and Sandnes.. Ford and Fisher. van Heel. 0. Rice. The tongue. N. 1984). Popper (1980) suggested that killer whales have a diminished use of echolocation and rely more on passive listening. orca has been documented to bowride ships (Dahlheim. 1978). Christensen.. Pod composition in Alaskan waters was 19% adult males. flipper slapping. and females and young in the middle (Norris and Dohl. AWBREY. A. Killer whales are capable of producing two different sounds simultaneously indicating dual sound sources as recorded for Tursiops GENETICS. Approximately 19% of animals within pods were adult males. 1971). 55% adult females or subadult males. Rept.. killer whales were observed apparently to peer onto ice floes for seals and sometimes hit the ice from below to knock pinnipeds or penguins (Spheniscidae) into the water (Fraser. LEATHERWOOD. Foraging behavior varies. In the Pacific Northwest. In the Antarctic. Balcomb. 1866.. 1980). Killer whales seem to maintain a social hierarchical system. The functional significance of dialects is not known.. 1983). 1981). T. Many large delphinids such as Globicephala sp. killer whales have the best absolute acoustical sensitivity. considerably larger than the minimum size of a 2-mm object detected by a Lagenorhynchus obliquidens (Popper. Because of conspicuous C:band polymorphism. EVANS. and lob tailing are common for killer whales. O. 1982. Report of the workshop on identity. Eschricht. the composition of pods seems to remain consistent through time (Bigg. and 64% involved killer whales partially stranding themselves temporarily to capture pinnipeds in the surf (Lopez and Lopez. however killer whales can achieve speeds of at least 40 km/h (Lang. usually adult males. Swimming speeds usually are 6 to 10 km/h. the karyotype of each specimen was unique (Arnason et aI. 1966. 1949). 1985). 1974). 1982. Internat. 1980). and type of prey. R. The C-banding pattern differs slightly from other delphinids because of the accumulation of heterochromatin. 1980. 1:227-287.F. The vernacular name orca is used commonly.. and genital region of baleen whales seem to be the primary target of 0. 1966). and Pseudorca crassidens are well known to mass strand. REMARKS.MAMMALIAN SPECIES 304 5 truncatus. SANDHOLT.5 h during which time the pod retained its structure with the adult males on the wings of a rank-shaped formation.. Care-giving behavior is well documented (Caldwell and Caldwell. 32:667670. Aquatic Mamm. Pods that associate tend to share certain signals. Mus. Killer whale pods were reported to alert the whalers of the presence of a nearby whale. 1957). Whales and dolphins of New Zealand and .. Most activities seem to be group oriented (Martinez and Klinghammer. Nat. killer whales attack in a pack and tear at the whale from several angles. 1980). 1983. Agoes-Makowski drew Fig. 94% involved cooperative hunting with a 31 % success rate. 1961). in general. orca are not as common (Baker. 1958. and those that do not associate or travel together emit different call repertoires. The G-banded karyotypes of O. The genus Orcinus currently is considered monotypic. Dearden. 1982). and call patterns were noted in the Antarctic (Thomas et aI.ANDS. Off the coast of Argentina. 1985). 1983. 1983). 1. After killing the whale.. 1982). but the biological significance of these large congregations is not understood clearly. Cytogenet. and vital rates of killer whale populations. Large groups may reflect a temporary joining of a number of pods possibly related to seasonal peaks in prey availability or possibly social activities. seemingly. E. and the North Pacific (Awbrey et aI.. Washington. Parallel scars caused by the teeth marks of other killer whales indicate that some intraspecific aggression occurs (Greenwood et aI. 1980). Goodall. 1978. Evans and Bastian (1969) reported that killer whales circle a school of fish then one or two whales break rank and dash through the middle of the school to feed. ARNASON. and British Columbia. 5:21-27.. Hall and Johnson (1971) presented an audiogram of Orcinus with a sensitivity range from 500 Hz to 31 kHz with the greatest sensitivity at 15 kHz (-70 ± 5 dB re 1 dyne/ ern"). Large aggregations also were reported from Antarctic and Norwegian waters (Budylenko. R. Internat. North Atlantic (Jehl et al. possibly a recently evolved secondary characteristic (Amason et al. Lenfant (1969) reported that length of dives ranged from 1 to 10 min. 1982). structure. AND B. Recordings from captive animals of known pod origin indicate that pod repertoires are stable over time (Ford and Fisher. Killer whales were observed to chase gray whales (Eschrichtius robustus) in the Bering Sea. 1981) and O. They approached the gray whales initially in a linear formation. Groups of several hundred animals were observed in Alaskan waters (Braham and Dahlheim. Underwater visual acuity is reported to be as good as that of pinnipeds and most teleost fish (Dawson. E. 32:617-631. Ross Sea killer whale vocalizations: preliminary description and comparison with those of some Northern Hemisphere killer whales. Cell Genet. Individual and group differences in signals have been well documented (Dahlheim and Awbrey. and 27% juveniles and calves (Leatherwood et aI. Young killer whales are believed to be taught this technique of capturing pinnipeds (Lopez and Lopez. glacialis (Berzin and Vladimirov. 1962). then gave pursuit in a loose cresent-shaped formation with the individuals spaced approximately 300 m apart (Ljungblad and Moore. orca are similar to the karyotypes of Tursiops and Stenella sp. 1980. Whaling Comm.. a conservative view of recognizing only one highly variable species probably is warranted.W. in California. In the late 1800's and early 1900's an unusual cooperative relationship between killer whales and shore-based whalers occurred in Australia. Regional differences in frequency.. Duffield-Kulu. (Arnason et al. Banding studies on six killer whales: an account of C-band polymorphism and G-band patterns. Cameron.. it is dependent on the size. 1980). occur infrequently (Norris and Prescott. The karyotype of the killer whale is 2n = 44. 1982. individual pods can be identified based solely on dialectal variation (Ford and Fisher. but with a reduced audible frequency range. and 41 % immature whales of both sexes. BAKER.by most authorities with geographical variation noted in size and color pattern. ASPER.. A. the whalers allowed the killer whales to feed upon the tongues of the dead whales (Wellings. 1941. Brownell.3. 1977.. (1982) noted that pods contained a maximum of 75 animals with a mean of 5. typical for most cetaceans (Arnason et aI. 1980). Mitchell. Tomilin. lips.. both seem to refer to the same population of smaller individuals that have more yellowish-pigmented light areas. LITERATURE CITED ANDREWS. J. D. Hist. Two recently described Antarctic species. The smallest object that could be discriminated by use of echolocation by a killer whale in captivity was a 10-mm plastic ring. Killer whales have an extensive repertoire of sounds with most phonations in the range of 4 to 5 kHz (Dahlheim and Awbrey. A pod of 15 killer whales was followed and harassed by the RV Alpha Helix for 6. R. E.R. Chovan. Amer. orca behavior in captivity is provided by Defran and Pryor (1980). National Marine Fisheries Service drew Fig. Known multipod gatherings were recorded for waters off southern Vancouver Island. H. and D. spyhopping. LUTLEY. of 568 observations of hunting killer whales. about one-half of the pods had 5 to 10 animals but pods of as many as 100 were observed (Mikhalev et aI. Orcinus nanus (Mikhalev et aI. 40% adult females. THOMAS.. I. 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. NATIONAL MARINE MAMMAL LABORATORY. Editors of this account were B. (R. DAHLHEIM. AND MARILYN E. Busnel York. Fish. SYDNEY ANDERSON. NATURAL HISTORY MUSEUM OF Los ANGELES COUNTY. SEATTLE. SECTION OF BIRDS AND MAMMALS. 1135 pp. 7600 SAND POINT WAY N E. J. C. eds. SPECIES systems New 304 and 9 1. Los ANGELES. CALIFORNIA 90007.MAMMALIAN sonar Press. HEYNING. Plenum JOHN E.). VERTS and Managing editor was CARLETON J. WASHINGTON 98115.. NATIONAL MARINE FISHERIES SERVICE. F. PHILLIPS. 900 EXPOSITION BLVD.