FORAGING ECOLOGY OF NORTHERN ELEPHANT SEALS

Sexual segregation in foraging is predicted from the great size disparity of male and female northern elephant seals, Mirounga angustirostris. Our aim was to test this prediction by measuring diving and foraging behavior, foraging locations, and distribution of the sexes during biannual migrations in the northeastern Pacific Ocean. Daily movements of 27 adult males and 20 adult females, during 56 migrations from Afio Nuevo, California, USA, were determined by Argos satellite telemetry via head-mounted platform transmitter terminals. Diving records were obtained with archival time-depth-speed recorders attached to the backs of seals that were recovered when the seals returned to the rookery. Pronounced sex differences were found in foraging location and foraging pattern, as reflected by hor- izontal transit speed and diving behavior. Males moved directly north or northwest at a mean speed of 90 ? 27 km/d to focal foraging areas along the continental margin ranging from coastal Oregon (534 km away) to the western Aleutian Islands (4775 km away). Males remained in these areas (mean size = 7892 ki2) for 21-84% of their 4-mo stays at sea. The predominance of flat-bottom dives in these areas suggests concentrated feeding on benthic prey. Migration distance and estimated mass gain were positively correlated with male size, and individual males returned to the same area to forage on subsequent migrations. In contrast, females ranged across a wider area of the northeastern Pacific, from 380 to 600 N and from the coast to 172.5? E. Focal foraging areas, indicated by a reduction in swim speed to <0.4 m/s, were distributed over deep water along the migratory path, with females remaining on them a mean of 3.5 d before moving to another one. Jagged-bottom dives that tracked the deep scattering layer prevailed in these areas, suggesting that females were feeding on pelagic prey in the water column. Females took roughly similar initial paths in subsequent migrations, but large deviations from the previous route were observed. We conclude that there is habitat segregation between the sexes. Females range widely over deep water, apparently foraging on patchily distributed, vertically migrating, pelagic prey, whereas males forage along the continental margin at the distal end of their migration in a manner consistent with feeding on benthic prey.

[1]  C. A. Hui Undersea topography and distribution of dolphins of the genus Delphinus in the southern California bight , 1979 .

[2]  Stephen Harris,et al.  Home‐range analysis using radio‐tracking data–a review of problems and techniques particularly as applied to the study of mammals , 1990 .

[3]  P. F. Brodie,et al.  Cetacean Energetics, an Overview of Intraspecific Size Variation , 1975 .

[4]  Brent S. Stewart,et al.  DIVING PATTERNS OF NORTHERN ELEPHANT SEAL BULLS , 1991 .

[5]  R. M. Laws,et al.  Elephant seals : population ecology, behavior, and physiology , 1994 .

[6]  Henrickson Rv,et al.  Ketamine immobilization of northern elephant seals. , 1975 .

[7]  J. Berger,et al.  Pregnancy incentives, predation constraints and habitat shifts: experimental and field evidence for wild bighorn sheep , 1991, Animal Behaviour.

[8]  K. Hobson,et al.  STABLE ISOTOPE ANALYSES OF TOOTH ANNULI REVEAL TEMPORAL DIETARY RECORDS: AN EXAMPLE USING STELLER SEA LIONS , 1998 .

[9]  Daniel P. Costa,et al.  Continuous, deep diving in female northern elephant seals, Mirounga angustirostris , 1988 .

[10]  P. Pyle,et al.  Spatiotemporal Patterns of White Shark (Carcharodon carcharias) Predation at the South Farallon Islands, California , 1992 .

[11]  F. Weckerly,et al.  Sexual Segregation in Ungulates: New Directions for Research , 1996 .

[12]  D. Costa,et al.  A programmable acoustic recording tag and first results from free-ranging northern elephant seals , 1998 .

[13]  P. Stabeno,et al.  Long-term measurements of flow near the Aleutian Islands , 1997 .

[14]  M. Dahlheim A photographic catalog of killer whales, Orcinus orca, from the central Gulf of Alaska to the southeastern Bering Sea , 1997 .

[15]  E. Nordøy,et al.  Energy sources in fasting grey seal pups evaluated with computed tomography. , 1985, The American journal of physiology.

[16]  Fred J. Brenner,et al.  Animals in Migration , 1970 .

[17]  B. McConnell,et al.  Movements of southern elephant seals , 1996 .

[18]  J. French Wildlife telemetry by satellite , 1994 .

[19]  Yasuhiko Naito,et al.  Diving behavior in relation to ambient water temperature in northern elephant seals , 1994 .

[20]  D. Costa,et al.  The energetics of lactation in the northern elephant seal , 1986 .

[21]  Daniel P. Costa,et al.  Swim speed in a female northern elephant seal: metabolic and foraging implications , 1992 .

[22]  M. C. Macaulay,et al.  Cetaceans in the Great South Channel, 1979–1989: right whale (Eubalaena glacialis) , 1995 .

[23]  D. Crocker,et al.  DIVING BEHAVIOR OF JUVENILE NORTHERN ELEPHANT SEALS , 1996 .

[24]  H. Dingle Migration: The Biology of Life on the Move , 1996 .

[25]  M. Bowers,et al.  Differential Habitat Utilization by Sexes of the Deermouse, Permomyscus Maniculatus , 1979 .

[26]  S. Reilly,et al.  Seasonal changes in distribution and habitat differences among dolphins in the eastern tropical Pacific , 1990 .

[27]  G. Bartholomew A MODEL FOR THE EVOLUTION OF PINNIPED POLYGYNY , 1970, Evolution; international journal of organic evolution.

[28]  S. Iverson,et al.  Fatty acid signatures reveal fine scale structure of foraging distribution of harbor seals and their prey in Prince William Sound, Alaska , 1997 .

[29]  Gerald L. Kooyman,et al.  Diverse Divers: Physiology and behavior , 1989 .

[30]  Distribution and behaviour of male sperm whales on the Scotian Shelf, Canada , 1992 .

[31]  Donald J. Hall,et al.  Ontogenetic Habitat Shifts in Bluegill: The Foraging Rate‐Predation Risk Trade‐off , 1988 .

[32]  B. L. Boeuf,et al.  Size, dominance and copulatory success in male northern elephant seals, Mirounga angustirostris , 1994, Animal Behaviour.

[33]  V. Bleich Mountain Sheep and Coyotes: Patterns of Predator Evasion in a Mountain Ungulate , 1999 .

[34]  David C. Douglas,et al.  Satellite Telemetry: A New Tool for Wildlife Research and Management, , 1988 .

[35]  B. L. Boeuf,et al.  Feeding habits and feeding grounds of the Northern elephant seal , 1984 .

[36]  J. Wehausen,et al.  Sexual Segregation in Mountain Sheep: Resources or Predation? , 1997 .

[37]  K. Kenyon The sea otter in the eastern Pacific Ocean , 1975 .

[38]  P. Sherman,et al.  Sexual dimorphisms and breeding systems in pinnipeds, ungulates, primates, and humans , 1979 .

[39]  A. Gautier-hion Seasonal Variations of Diet Related to Species and Sex in a Community of Cercopithecus Monkeys , 1980 .

[40]  Yasuhiko Naito,et al.  Prolonged, continuous, deep diving by northern elephant seals , 1989 .

[41]  E. Born,et al.  Haul‐out and diving activity of male Atlantic walruses (Odobenus rosmarus rosmarus) in NE Greenland , 1997 .

[42]  T. Clutton‐Brock,et al.  Sexual segregation and density-related changes in habitat use in male and female Red deer (Cerrus elaphus) , 1987 .

[43]  S. Newhouse,et al.  Resource Partitioning between Sexes in White-Tailed Deer , 1989 .

[44]  Tinker,et al.  Killer whale predation on sea otters linking oceanic and nearshore ecosystems , 1998, Science.

[45]  R. Dietz,et al.  Movements and swimming speed of narwhals, Monodon monoceros, equipped with satellite transmitters in Melville Bay, northwest Greenland , 1995 .

[46]  Madhav Gadgil,et al.  Male-female differences in foraging on crops by Asian elephants , 1988, Animal Behaviour.

[47]  R. C. Lasiewski,et al.  Heating and cooling rates, heart rate and simulated diving in the Galapagos marine iguana. , 1965, Comparative biochemistry and physiology.

[48]  B. McConnell,et al.  Post-Breeding Distribution and Diving Behavior of Adult Male Southern Elephant Seals from Patagonia , 1999 .

[49]  L. Dill,et al.  A DETERMINATION OF THE ENERGETIC EQUIVALENCE OF THE RISK OF PREDATION , 1989 .

[50]  D. Crocker,et al.  Effects of buoyancy on the diving behavior of northern elephant seals. , 1998, The Journal of experimental biology.

[51]  I. Boyd,et al.  Milk fatty acid signatures indicate both major and minor shifts in the diet of lactating Antarctic fur seals , 1997 .

[52]  D. Demaster,et al.  ASSESSING NORTHERN ELEPHANT SEAL FEEDING HABITS BY STOMACH LAVAGE , 1987 .

[53]  BODY SIZE AND SEXUAL SIZE DIMORPHISM IN MARINE IGUANAS FLUCTUATE AS A RESULT OF OPPOSING NATURAL AND SEXUAL SELECTION: AN ISLAND COMPARISON , 1997 .

[54]  R. Helm Rate of digestion in three species of pinnipeds , 1984 .

[55]  W. Bowen,et al.  Seasonal and Geographic Variation in the Species Composition and Size of Prey Consumed by Grey Seals (Halichoerus grypus) on the Scotian Shelf , 1993 .

[56]  G. Bartholomew A Field Study of Temperature Relations in the Galapagos Marine Iguana , 1966 .

[57]  D. Crocker,et al.  CHAPTER 17 – Diving Behavior of Elephant Seals: Implications for Predator Avoidance , 1996 .

[58]  Bernie J. McConnell,et al.  Foraging ecology of southern elephant seals in relation to the bathymetry and productivity of the Southern Ocean , 1992, Antarctic Science.

[59]  N. Gales,et al.  Ultrasonic measurement of blubber thickness of the southern elephant seal, Mirounga leonina (Linn.) , 1987 .

[60]  Sara J. Iverson,et al.  Fatty acid signatures and classification trees: new tools for investigating the foraging ecology of seals , 1997 .

[61]  Daniel P. Costa,et al.  Drift diving in female northern elephant seals: implications for food processing , 1997 .

[62]  B. Stewart Ontogeny of Differential Migration and Sexual Segregation in Northern Elephant Seals , 1997 .

[63]  Brent S. Stewart,et al.  DOCUMENTING MIGRATIONS OF NORTHERN ELEPHANT SEALS USING DAY LENGTH , 1992 .

[64]  B. L. Boeuf,et al.  A METHOD FOR ESTIMATING MASS OF LARGE PINNIPEDS , 1991 .

[65]  Imants G. Priede,et al.  Tracking of marine animals by satellite , 1991 .

[66]  D. W. Strasburg Distribution, abundance, and habits of pelagic sharks in the central Pacific ocean , 1958 .

[67]  B. McConnell,et al.  Satellite tracking of grey seals (Halichoerus grypus) , 1992 .

[68]  J. A. Litvaitis Differential Habitat Use by Sexes of Snowshoe Hares (Lepus americanus) , 1990 .

[69]  M. Riedman,et al.  The Pinnipeds: Seals, Sea Lions, and Walruses , 1989 .

[70]  Brent S. Stewart,et al.  Double Migrations of the Northern Elephant Seal, Mirounga angustirostris , 1995 .

[71]  Peter W. Sorensen,et al.  The distributional biology of the right whale ( Eubalaena glacialis) in the western North Atlantic. , 1986 .

[72]  S. B. Blackwell,et al.  Onboard acoustic recording from diving northern elephant seals. , 1996, The Journal of the Acoustical Society of America.

[73]  B. L. Boeuf,et al.  Male-male Competition and Reproductive Success in Elephant Seals , 1974 .

[74]  D. Costa,et al.  Moult energetics of the northern elephant seal (Mirounga angustirostris) , 1992 .

[75]  Donald L. Kramer,et al.  The behavioral ecology of air breathing by aquatic animals , 1988 .