Haul-Out Behaviour of the World's Northernmost Population of Harbour Seals (Phoca vitulina) throughout the Year

The harbour seal population in Svalbard occurs at the northernmost limit of the species' range. It experiences environmental extremes far beyond the norm for this species, including an extended period of polar night and extensive sea ice cover. In 2009 and 2010, 60 harbour seals (30 pups + 30 immature/mature seals) from this population were equipped with Satellite-Relay Data Loggers (SRDLs) to study their haul-out behaviour, with a special focus on the winter period. Using a combination of Generalized Additive Mixed Models and Cox Proportional Hazard models, the influences of sex, maturity, temporal, spatial and environmental factors on haul-out behaviour were explored. All of the seals continued to haul out even through the coldest periods during the polar night, though clear seasonality in the time spent hauled out daily was displayed by both immature and mature seals. Time spent hauled out daily decreased from ∼5.2 hrs in September to ∼1.2 hrs in February in these age groups, while pups displayed less seasonality (∼2.4 hrs/day throughout most of the year). The average at-sea period also exhibited seasonality, increasing to a maximum of ∼1.6 days in February (monthly maxima for individual animals ranged from 7 to 19 days). The seals showed a strong preference to haul out at low tide when hauling out on land but not when using sea ice as a haul-out platform. A diel rhythm in haul-out behaviour was present during the months with day–night cycling and midnight sun but not during the polar night. Haul-out behaviour was impacted to a greater extent by air pressure, through its effect on wind speed, than by absolute temperature values. The extreme environment in Svalbard likely causes some physiological challenges that might impact survival rates negatively, particularly among pups. Climate warming is likely to have positive effects on Svalbard's harbour seal population.

[1]  K. Kovacs,et al.  The World’s Northernmost Harbour Seal Population–How Many Are There? , 2013, PloS one.

[2]  I. Stirling,et al.  Habitat use by harbour seals (Phoca vitulina) in a seasonally ice-covered region, the western Hudson Bay , 2013, Polar Biology.

[3]  Robert J. Small,et al.  Sex- and age-specific survival of harbor seals (Phoca vitulina) from Tugidak Island, Alaska , 2012 .

[4]  J. V. Ver Hoef,et al.  Haul-Out Behavior of Harbor Seals (Phoca vitulina) in Hood Canal, Washington , 2012, PloS one.

[5]  Toby A. Patterson,et al.  Spatial Variation in Foraging Behaviour of a Marine Top Predator (Phoca vitulina) Determined by a Large-Scale Satellite Tagging Program , 2012, PloS one.

[6]  L. Bachmann,et al.  A missing piece in the Arctic food web puzzle? Stomach contents of Greenland sharks sampled in Svalbard, Norway , 2012, Polar Biology.

[7]  G. Pendleton,et al.  Use of glacial and terrestrial habitats by harbor seals in Glacier Bay, Alaska: costs and benefits , 2011 .

[8]  K. Kovacs,et al.  A population on the edge: genetic diversity and population structure of the world's northernmost harbour seals (Phoca vitulina) , 2011 .

[9]  Sanford Weisberg,et al.  An R Companion to Applied Regression , 2010 .

[10]  B. McConnell,et al.  Winter habitat use of harbour seals ( Phoca vitulina ) fitted with Fastloc™GPS/GSM tags in two tidal bays in France , 2010 .

[11]  K. Kovacs,et al.  Status and biology of harbour seals ( Phoca vitulina ) in Svalbard , 2010 .

[12]  A. S. Blix,et al.  Remarkable development of diving performance and migrations of hooded seals (Cystophora cristata) during their first year of life , 2010, Polar Biology.

[13]  Alain F. Zuur,et al.  A protocol for data exploration to avoid common statistical problems , 2010 .

[14]  Mark A. Hindell,et al.  Factors influencing the winter haulout behaviour of Weddell seals: Consequences for satellite telemetry , 2010 .

[15]  Aaron Christ,et al.  Mixed Effects Models and Extensions in Ecology with R , 2009 .

[16]  Paul M. Thompson,et al.  Guidelines for the treatment of marine mammals in field research , 2009 .

[17]  C. D. Duck,et al.  Harbour seal movements and haul-out patterns: implications for monitoring and management , 2009 .

[18]  Monique MacKenzie,et al.  Estimating seasonal abundance of a central place forager using counts and telemetry data , 2009 .

[19]  B. McConnell,et al.  Using mobile phone telemetry to investigate the haul-out behaviour of harbour seals Phoca vitulina vitulina , 2009 .

[20]  K. Kovacs,et al.  Climate Change Impacts on Seals and Whales in the North Atlantic Arctic and Adjacent Shelf Seas , 2008, Science progress.

[21]  Alan Y. Chiang,et al.  Generalized Additive Models: An Introduction With R , 2007, Technometrics.

[22]  A. S. Blix,et al.  Ross seal (Ommatophoca rossii) annual distribution, diving behaviour, breeding and moulting, off Queen Maud Land, Antarctica , 2007, Polar Biology.

[23]  Robert P Freckleton,et al.  Why do we still use stepwise modelling in ecology and behaviour? , 2006, The Journal of animal ecology.

[24]  S. Wood Generalized Additive Models: An Introduction with R , 2006 .

[25]  A. S. Blix,et al.  Circadian organization in reindeer , 2005, Nature.

[26]  R. Small,et al.  Regional differences in diving behavior of harbor seals in the Gulf of Alaska , 2004 .

[27]  K. Kovacs,et al.  Long-distance movements of harbour seals (Phoca vitulina) from a seasonally ice-covered area, the St. Lawrence River estuary, Canada , 2004 .

[28]  K. Kovacs,et al.  Haulout behaviour of High Arctic harbour seals (Phoca vitulina vitulina) in Svalbard, Norway , 2003, Polar Biology.

[29]  P. Boveng,et al.  STABILITY IN THE PROPORTION OF HARBOR SEALS HAULED OUT UNDER LOCALLY IDEAL CONDITIONS , 2003 .

[30]  David R. Anderson,et al.  Model selection and multimodel inference : a practical information-theoretic approach , 2003 .

[31]  T. Bekkby,et al.  DISPERSAL and BYCATCH MORTALITY IN GRAY, HALICHOERUS GRYPUS, AND HARBOR, PHOCA VITULINA, SEALS TAGGED AT THE NORWEGIAN COAST , 2002 .

[32]  K. Kovacs,et al.  Diving behaviour of sub-adult harbour seals (Phoca vitulina) at Prins Karls Forland, Svalbard , 2002, Polar Biology.

[33]  Mike Fedak,et al.  Overcoming the Constraints of Long Range Radio Telemetry from Animals: Getting More Useful Data from Smaller Packages1 , 2002, Integrative and comparative biology.

[34]  K. Kovacs,et al.  Diving development in nursing harbour seal pups. , 2001, The Journal of experimental biology.

[35]  Ø. Wiig,et al.  Distribution and diving of harbour seals (Phoca vitulina) in Svalbard , 2001, Polar Biology.

[36]  P. Thompson,et al.  Display-area size, tenure length, and site fidelity in the aquatically mating male harbour seal, Phoca vitulina , 2000 .

[37]  F. Trillmich,et al.  Lunar cycles in diel prey migrations exert a stronger effect on the diving of juveniles than adult Gal pagos fur seals , 1999, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[38]  E. Reierth,et al.  Activity rhythm in High Arctic Svalbard ptarmigan (Lagopus mutus hyperboreus) , 1998 .

[39]  D. Lavigne,et al.  Ontogeny of the Thermal Limits in the Harbor Seal (Phoca vitulina) , 1997, Physiological Zoology.

[40]  K. Kovacs,et al.  AGE ESTIMATION OF GREY SEALS (HALICHOERUS GRYPUS)USING INCISORS , 1996 .

[41]  S. Innes,et al.  Energy Metabolism and Thermoregulation in Juvenile Harbor Seals (Phoca vitulina) in Air , 1995, Physiological Zoology.

[42]  R. Barry,et al.  Atmosphere, Weather and Climate (Sixth Edition) , 1994 .

[43]  D. Lavigne,et al.  Models of Heat Loss by Marine Mammals: Thermoregulation Below the Zone of Irrelevance , 1993 .

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

[45]  K. Kovacs,et al.  Maternal investment in otariid seals and walruses , 1992 .

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

[47]  D. Boness,et al.  The Pinnipeds: Seals, Sea Lions, and Walruses., Marianne Riedman (Ed.). University of California Press, Berkeley (1990), xxiii , 1991 .

[48]  P. Thompson,et al.  Winter Foraging by Common Seals (Phoca Vitulina) in Relation to Food Availability in the Inner Moray Firth, N.E. Scotland , 1991 .

[49]  T. Vinje,et al.  Barents Sea drift ice characteristics , 1991 .

[50]  I. Gjertz,et al.  THE MOST NORTHERLY HARBOR SEAL, PHOCA VITULINA, AT PRINS KARLS FORLAND, SVALBARD , 1990 .

[51]  B. McConnell,et al.  SEASONAL AND SEX-RELATED VARIATION IN THE ACTIVITY PATTERNS OF COMMON SEALS (PHOCA VITULINA) , 1989 .

[52]  Ø. Wiig A DESCRIPTION OF COMMON SEALS, PHOCA VITULINA L. 1758, FROM SVALBARD , 1989 .

[53]  J. Godsell Herd formation and haul-out behaviour in harbour seals (Phoca vitulina) , 1988 .

[54]  D. Demaster,et al.  DIEL HAUL‐OUT PATTERNS AND SITE FIDELITY OF HARBOR SEALS (PHOCA VITULINA RICHARDSI) ON SAN MIGUEL ISLAND, CALIFORNIA, IN AUTUMN , 1987 .

[55]  P. Thompson,et al.  Age and sex differences in the timing of moult in the common seal, Phoca vitulina , 1987 .

[56]  B. Taylor,et al.  Distribution and haul-out behavior of harbor seals in Glacier Bay, Alaska , 1987 .

[57]  K. Kovacs,et al.  Maternal investment and neonatal growth in phocid seals , 1986 .

[58]  K. Pitcher Variation in Blubber Thickness of Harbor Seals in Southern Alaska , 1986 .

[59]  B. Stewart Diurnal hauling patterns of harbor seals at San Miguel Island, California , 1984 .

[60]  P. Wadhams The Ice Cover in the Greenland and Norwegian Seas (Paper 1R0463) , 1981 .

[61]  L. Irving,et al.  Metabolism and temperature regulation in young harbor seals Phoca vitulina richardi. , 1975, The American journal of physiology.

[62]  W. Hamilton Geometry for the selfish herd. , 1971, Journal of theoretical biology.

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

[64]  J. Ling Pelage and Molting in Wild Mammals with Special Reference to Aquatic Forms , 1970, The Quarterly Review of Biology.

[65]  R. Barry,et al.  Atmosphere, Weather and Climate , 1968 .

[66]  A. Mansfield DISTRIBUTION OF THE HARBOR SEAL, PHOCA VITULINA LINNAEUS, IN CANADIAN ARCTIC WATERS , 1967 .

[67]  E. Feltz,et al.  Thermal requirements in vitro of epidermal cells from seals. , 1966, Cryobiology.

[68]  J. Hart,et al.  THE METABOLISM AND INSULATION OF SEALS AS BARE-SKINNED MAMMALS IN COLD WATER , 1957 .

[69]  A. Dreher Modeling Survival Data Extending The Cox Model , 2016 .

[70]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[71]  S. Gerland,et al.  Variability of fast-ice thickness in Spitsbergen fjords , 2006, Annals of Glaciology.

[72]  K. Kovacs,et al.  Growth and population parameters of the world’s northernmost harbour seals Phoca vitulina residing in Svalbard, Norway , 2004, Polar Biology.

[73]  John J. Burns,et al.  THE ABUNDANCE OF HARBOR SEALS IN THE GULF OF ALASKA , 2003 .

[74]  I. Gjertz,et al.  A REVIEW OF THE DISTRIBUTION AND ABUNDANCE OF HARBOR SEALS, PHOCA VITULINA, ON SVALBARD, NORWAY, AND IN THE BARENTS SEA , 1997 .

[75]  M. Fedak,et al.  Habitat use and diving behaviour of harbour seals in a coastal archipelago in Norway , 1995 .

[76]  A. Bjørge,et al.  Haul-out behaviour of the Norwegian harbour seal during summer , 1995 .

[77]  A. S. Blix,et al.  Whales, seals, fish, and man : proceedings of the International Symposium on the Biology of Marine Mammals in the North East Atlantic, Tromsø, Norway, 29 November-1 December 1994 , 1995 .

[78]  P. Grambsch,et al.  A Package for Survival Analysis in S , 1994 .

[79]  P. M. Thompson Harbour seal movement patterns , 1993 .

[80]  J. F. R. McIlveen,et al.  Fundamentals of weather and climate , 1991 .

[81]  J. Boulva,et al.  Biology of the harbor seal, Phoca vitulina, in eastern Canada , 1979 .

[82]  A. Thompson,et al.  Physical Oceanography , 1961, Nature.

[83]  E. Heske,et al.  American Society of Mammalogists , 1919 .

[84]  D.,et al.  Regression Models and Life-Tables , 2022 .