Intraspecific variation in movement patterns: modeling individual behaviour in a large marine predator

In large marine predators, foraging entails movement. Quantitative models reveal how behaviours can mediate individual movement, such that deviations from a random pattern may reveal specific search tactics or behaviour. Using locations for 52 grey seals fitted with satellite-linked recorders on Sable Island; we modeled movement as a correlated random walk (CRW) for individual animals, at two temporal scales. Mean move length, turning angle, and net squared displacement (R 2 n: the rate of change in area over time) at successive moves over 3 to 10 months were calculated. The distribution of move lengths of individual animals was compared to a Levy distribution to determine if grey seals use a Levy flight search tactic. Grey seals exhibited three types of movement as determined by CRW model fit: directed movers - animals displaying directed long distance travel that were significantly underpredicted by the CRW (23% of animals); residents - animals remaining in the area surrounding Sable Island that were overpredicted by the model (29% of animals); and correlated random walkers those (48% of animals) in which movement was predicted by the CRW model. Kernel home range size differed significantly among all three movement types. as did travel speed, mean move length, mean R 2 n and total distance traveled. Sex and season of deployment were significant predictors of movement type, with directed movers more likely to be male and residents more likely to be female. Only 30% of grey seals fit a Levy distribution, which suggests that food patches used by the majority of seals are not randomly distributed. Intraspecific variation in movement behaviour is an important characteristic in grey seal foraging ecology, underscoring the need to account for such variability in developing models of habitat use and predation.

[1]  M. Shlesinger,et al.  Beyond Brownian motion , 1996 .

[2]  J. Godsell The relative influence of age and weight on the reproductive behaviour of male grey seals Halichoerus grypus , 1991 .

[3]  I. Newton Population Ecology of Raptors , 1981 .

[4]  I. McLAREN GROWTH IN PINNIPEDS , 1993, Biological reviews of the Cambridge Philosophical Society.

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

[6]  Michael L. Cain,et al.  Random Search by Herbivorous Insects: A Simulation Model , 1985 .

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

[8]  Andrew J. Read,et al.  Monitoring the movements of harbour porpoises (Phocoena phocoena) with satellite telemetry , 1997 .

[9]  A. Hofgaard,et al.  Foraging and movement paths of female reindeer: insights from fractal analysis, correlated random walks, and Lévy flights , 2002 .

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

[11]  W. Bowen,et al.  Variation in Milk Production and Lactation Performance in Grey Seals and Consequences for Pup Growth and Weaning Characteristics , 1999, Physiological and Biochemical Zoology.

[12]  S. L. Lima,et al.  SEARCH STRATEGIES FOR LANDSCAPE‐LEVEL INTERPATCH MOVEMENTS , 1999 .

[13]  W. Bowen,et al.  Grey seal predation on the eastern Scotian Shelf: modelling the impact on Atlantic cod , 1996 .

[14]  L. Barrett,et al.  Random walks and the gas model: spacing behaviour of Grey-Cheeked Mangabeys , 1998 .

[15]  W. Bowen Home Range and Spatial Organization of Coyotes in Jasper National Park, Alberta , 1982 .

[16]  M. Stanton Searching in a Patchy Environment: Foodplant Selection by Colis P. Eriphyle Butterflies , 1982 .

[17]  B. Worton Using Monte Carlo simulation to evaluate kernel-based home range estimators , 1995 .

[18]  D. Kobayashi,et al.  Turtles on the edge: movement of loggerhead turtles (Caretta caretta) along oceanic fronts, spanning longline fishing grounds in the central North Pacific, 1997–1998 , 2000 .

[19]  E. L. Core Plant Ecology of Spruce Mountain, West Virginia , 1929 .

[20]  Martha H. Winsor,et al.  Satellite-monitored movements of radio-tagged bowhead whales in the Beaufort and Chukchi seas during the late-summer feeding season and fall migration , 2000 .

[21]  Deborah Austin,et al.  A THREE‐STAGE ALGORITHM FOR FILTERING ERRONEOUS ARGOS SATELLITE LOCATIONS , 2003 .

[22]  B. McConnell,et al.  Comparative distribution, movements and diet of harbour and grey seals from the Moray Firth, N.E. Scotland , 1996 .

[23]  W. Bowen,et al.  Sex differences in diving at multiple temporal scales in a size‐dimorphic capital breeder , 2003 .

[24]  Sara J. Iverson,et al.  Sex differences in the diving behaviour of a size-dimorphic capital breeder: the grey seal , 2003, Animal Behaviour.

[25]  Bernie J. McConnell,et al.  Movements and foraging areas of grey seals in the North Sea , 1999 .

[26]  Wolf M. Mooij,et al.  Search paths of swans foraging on spatially autocorrelated tubers , 2002 .

[27]  W. Bowen,et al.  Mass and Energy Transfer during Lactation in a Small Phocid, the Harbor Seal (Phoca vitulina) , 1992, Physiological Zoology.

[28]  D. Macdonald,et al.  Scale‐free dynamics in the movement patterns of jackals , 2002 .

[29]  Robert Gisiner,et al.  Local and migratory movements of Hawaiian humpback whales tracked by satellite telemetry , 1998 .

[30]  Simon Benhamou,et al.  Optimal sinuosity in central place foraging movements , 1991, Animal Behaviour.

[31]  R. Powell,et al.  An Evaluation of the Accuracy of Kernel Density Estimators for Home Range Analysis , 1996 .

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

[33]  O. Vesakoski,et al.  Why does herbivore sex matter? Sexual differences in utilization of Fucus vesiculosus by the isopod Idotea baltica , 2001 .

[34]  M. Shlesinger,et al.  Lévy Walks Versus Lévy Flights , 1986 .

[35]  D. Levin,et al.  POLLINATOR FLIGHT DIRECTIONALITY AND ITS EFFECT ON POLLEN FLOW , 1971, Evolution; international journal of organic evolution.

[36]  D. E. Andersen,et al.  Home-Range Estimates of Red-Tailed Hawks Based on Random and Systematic Relocations , 1989 .

[37]  J. Damuth Home range, home range overlap, and species energy use among herbivorous mammals , 1981 .

[38]  W. Bowen,et al.  Role of marine mammals in aquatic ecosystems , 1997 .

[39]  J. A. Schaefer,et al.  Caribou movement as a correlated random walk , 2000, Oecologia.

[40]  H. Stanley,et al.  Optimizing the success of random searches , 1999, Nature.

[41]  K. Ruckstuhl,et al.  Foraging behaviour and sexual segregation in bighorn sheep , 1998, Animal Behaviour.

[42]  K. E. Hodges,et al.  Does body condition affect fecundity in a cyclic population of snowshoe hares , 1999 .

[43]  Eric J. Gustafson,et al.  The Effect of Landscape Heterogeneity on the Probability of Patch Colonization , 1996 .

[44]  S. Benhamou,et al.  Spatial analysis of animals' movements using a correlated random walk model* , 1988 .

[45]  R. Alford,et al.  Nomadic movement in tropical toads , 2002 .

[46]  F. Pérez-Barbería,et al.  Body size dimorphism and sexual segregation in polygynous ungulates: an experimental test with Soay sheep , 1999, Oecologia.

[47]  Yvan Simard,et al.  The rich krill aggregation of the Saguenay - St. Lawrence Marine Park: hydroacoustic and geostatistical biomass estimates, structure, variability, and significance for whales , 1999 .

[48]  J. P. Ball,et al.  Partial migration by large ungulates: characteristics of seasonal moose Alces alces ranges in northern Sweden , 2001, Wildlife Biology.

[49]  P. Kareiva,et al.  Analyzing insect movement as a correlated random walk , 1983, Oecologia.

[50]  Y. Watanuki,et al.  Sex and individual differences in foraging behavior of Japanese cormorants in years of different prey availability , 2002, Journal of Ethology.

[51]  O P Judson,et al.  The rise of the individual-based model in ecology. , 1994, Trends in ecology & evolution.

[52]  I. Gordon,et al.  The relative roles of phylogeny, body size and feeding style on the activity time of temperate ruminants: a reanalysis , 1999, Oecologia.

[53]  P. A. Prince,et al.  Lévy flight search patterns of wandering albatrosses , 1996, Nature.

[54]  Peter Turchin,et al.  Translating Foraging Movements in Heterogeneous Environments into the Spatial Distribution of Foragers , 1991 .

[55]  L. L. Wolf,et al.  Non-Random Foraging by Hummingbirds: Patterns of Movement Between Ipomopsis aggregata (Pursch) V. Grant Inflorescences , 1990 .

[56]  S. Ferguson,et al.  Space use by polar bears in and around Auyuittuq National Park, Northwest Territories, during the ice-free period , 1997 .

[57]  B. Cole Fractal time in animal behaviour: the movement activity of Drosophila , 1995, Animal Behaviour.

[58]  J. Ouellet,et al.  Seasonal home range size and philopatry in two northern white-tailed deer populations , 2000 .

[59]  A. S. Blix,et al.  Annual distribution of hooded seals (Cystophora cristata) in the Greenland and Norwegian seas , 1996, Polar Biology.

[60]  J. Estes The influence of large, mobile predators in aquatic food webs: examples from sea otters and kelp forests , 1996 .

[61]  Michael F. Shlesinger Physics in the noise , 2001, Nature.

[62]  P. Boersma,et al.  Satellite telemetry of the winter migration of Adélie penguins (Pygoscelis adeliae) , 1996, Polar Biology.

[63]  John P. Y. Arnould,et al.  Foraging behaviour of antarctic fur seals during periods of contrasting prey abundance , 1994 .

[64]  W. Bowen,et al.  Sex differences in the seasonal patterns of energy storage and expenditure in a phocid seal , 2003 .

[65]  Peter Kareiva,et al.  THE SEARCH FOR RESOURCES BY CABBAGE BUTTERFLIES (PIERIS RAPAE): ECOLOGICAL CONSEQUENCES AND ADAPTIVE SIGNIFICANCE OF MARKOVIAN MOVEMENTS IN A PATCHY ENVIRONMENT' , 1984 .

[66]  E. Batschelet Circular statistics in biology , 1981 .

[67]  Petr E. Komers,et al.  Behavioural plasticity in variable environments , 1997 .

[68]  W. B. Scott,et al.  Atlantic fishes of Canada , 1988 .

[69]  D. Bryant,et al.  The Searching Speeds of Foraging Shorebirds: Redshank (Tringa totanus) and Oystercatcher (Haematopus ostralegus) , 1993, The American Naturalist.

[70]  C. M. Lessells,et al.  THE NATURAL REGULATION OF GIANT TORTOISE POPULATIONS ON ALDABRA ATOLL. MOVEMENT POLYMORPHISM, REPRODUCTIVE SUCCESS AND MORTALITY , 1979 .

[71]  A. E. Punt,et al.  The effects of future consumption by the Cape fur seal on catches and catch rates of the Cape hakes. 4. Modelling the biological interaction between Cape fur seals Arctocephalus pusillus pusillus and the Cape hakes Merluccius capensis and M. paradoxus , 1995 .

[72]  I. Swingland,et al.  Movement patterns and morphometrics in giant tortoises , 1989 .

[73]  J. Reunanen Search Strategies , 2021, International Journal of Obesity.

[74]  J. P. Ball,et al.  Grey seal, Halichoerus grypus, habitat selection around haulout sites in the Baltic Sea: bathymetry or central-place foraging? , 2000 .

[75]  T. A. Hanley,et al.  ANALYSIS OF THE FUNCTIONAL RESPONSE IN FORAGING IN THE SITKA BLACK-TAILED DEER' , 1988 .

[76]  G. Pyke Optimal foraging: movement patterns of bumblebees between inflorescences. , 1978, Theoretical population biology.

[77]  L. Mech,et al.  Home-Range Formation and Dispersal of Deer in Northeastern Minnesota , 1984 .

[78]  G. Lincoln Biology of Seasonal Breeding in Deer , 1992 .

[79]  Daniel P. Costa,et al.  FORAGING ECOLOGY OF NORTHERN ELEPHANT SEALS , 2000 .

[80]  John E. Gross,et al.  The dynamics and scaling of foraging velocity and encounter rate in mammalian herbivores , 1996 .

[81]  W. Bowen,et al.  Sustained exponential population growth of grey seals at Sable Island, Nova Scotia , 2003 .

[82]  W. H. Burt Territoriality and Home Range Concepts as Applied to Mammals , 1943 .

[83]  M. Hindell,et al.  Foraging zones of royal penguins during the breeding season, and their association with oceanographic features , 1997 .

[84]  B. Worton Kernel methods for estimating the utilization distribution in home-range studies , 1989 .

[85]  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 .

[86]  M. Demment,et al.  Sex differences in giraffe foraging behavior at two spatial scales , 1997, Oecologia.