MARINE MAMMAL SCIENCE, 21(1):121 135 ( January 2005) Ó 2005 by the Society for Marine Mammalogy CLIMATE CHANGE AND RINGED SEAL (PHOCA HISPIDA) RECRUITMENT IN WESTERN HUDSON BAY STEVEN H. FERGUSON Fisheries and Oceans, 501 University Crescent, Winnipeg, Manitoba R3T 2N6, Canada E-mail: fergusonsh@dfo-mpo.gc.ca IAN STIRLING Canadian Wildlife Service, 5320 122 Street, Edmonton, Alberta T6H 3S5, Canada PHILIP MCLOUGHLIN Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2, Canada ABSTRACT Climate warming is predicted to reduce the extent of ice cover in the Arctic and, within the Hudson Bay region, the annual ice may be significantly decreased or entirely lost in the foreseeable future. The ringed seal (Phoca hispida), a key species that depends on sea ice, will likely be among the first marine mammals to show the negative effects of climatic warming. We used 639 ringed seals killed by Inuit hunters from western Hudson Bay (1991 1992, 1999 2001) to assess trends in recruitment relative to snow depth, snowfall, rainfall, temperature in April and May, North Atlantic Oscillation (NAO) from the previous winter, and timing of spring break-up. Snowfall and ringed seal recruitment varied from lower than average in the 1970s, to higher in 1980s and lower in 1990s. Prior to 1990, seal recruitment appeared to be related to timing of spring ice break-up which was correlated with the NAO. However, recent 1990 2001 environmental data indicate less snowfall, lower snow depth, and warmer temperatures in April and May when pups are born and nursed. Decreased snow depth, particularly below 32 cm, corresponded with a significant decrease in ringed seal recruitment as indicated by pups born and surviving to adults that were later harvested. Earlier spring break-up of sea ice together with snow trends suggest continued low pup survival in western Hudson Bay. Key words: Arviat, Inuit, North Atlantic Oscillation, Nunavut, Phoca hispida, pup survival, rainfall, sea ice break-up, snowfall, temperature. 121
122 MARINE MAMMAL SCIENCE, VOL. 21, NO. 1, 2005 Warming of the earth s climate is forecast to be greatest at the poles and, within the Arctic, western Hudson Bay is one of the areas in which the rate and degree of climatic warming is predicted to be greatest (Prinsenberg 1986, Parkinson 2000, Krajick 2001, Gough and Wolfe 2001, Comiso 2003). The challenge for resident species to accommodate such change is increased in the Arctic because of its large scale, the rapid rate at which the warming is predicted to occur, large interannual variation in climate, and the accelerated pace of human development (IPCC 2001). As a result, climate change in the Arctic is expected to have large effects (Hughes 2000, Levitus et al. 2001). Higher ocean temperatures and lower salinities, contraction of seasonal sea ice extent, rising sea levels, and a host of other effects (Munk 2003) are certain to have significant impacts on marine species. For marine mammals adapted to life with sea ice, the effects of reductions in sea ice are likely to be reflected initially by shifts in range and abundance (Tynan and DeMaster 1997). Demographic changes associated with shifts in geographic range will likely be observed as decreased recruitment in areas of reduced sea ice. Northern seal and walrus (Odobenus rosmarus) populations may represent useful indicators of ecological change in northern ecosystems because of their dependence upon annual sea ice (Tynan and DeMaster 1997) which has already undergone significant changes in annual cycles of distribution and abundance and significant further reduction is predicted (Comiso 2003). Although ringed seals (Phoca hispida) are the most abundant seal in the Arctic and are considered a keystone species in the marine ecosystem (Smith et al. 1991), little research has focused on the effect of climate warming on this species (Harwood et al. 2000, Stirling and Smith 2004). The evolutionary adaptations of ringed seals to exploit sea ice habitat for reproduction and overwinter survival makes this species particularly suitable for examining effects of climate warming (Lowry et al. 1980, Ryg et al. 1990, Smith et al. 1991, Moulton et al. 2002). Ringed seal pups are born from mid-march to mid-april, suckled for about six weeks, and weaned prior to spring break-up in June (McLaren 1958, 1993, Hammill and Smith 1991). During lactation pups spend about half their time in subnivean dens on top of the ice, and half underwater diving (Lydersen and Hammill 1993a, Furgal et al. 1996), during which time they and adults are hunted by polar bears (Stirling and Archibald 1977, Smith 1980). Pups in subnivean birth or haul-out lairs with thin snow roofs are more vulnerable to predators than those in lairs with thick ones (Smith and Stirling 1975, Hammill and Smith 1991, Furgal et al. 1996). Although there is considerable information on ringed seals in some other parts of the Arctic, information on their distribution, abundance, breeding habitat, and reproductive rates in western Hudson Bay is relatively limited (Smith 1975, Lunn et al. 1997, Holst et al. 1999). Largely because the sea ice in western Hudson Bay is both relatively unstable and inaccessible from shore-based locations by surface travel in April and May, there are no data on the distribution of birth lairs, habitat preferences, or vulnerability of pups to predation. In the absence of data, it has generally been assumed there are enough ringed seals to satisfy the needs of Inuit hunters and bears and that seal population sizes are large and stable. In a preliminary study however, Holst et al. (1999) reported that the pregnancy rate of ringed seals and the proportion of young-of-the-year in an open water sample from Arviat (Fig. 1) in 1991 1992 were unexpectedly low. Even though there are no direct data on ecological aspects of the birth lair habitat or productivity on the sea ice of western Hudson Bay, we are able to define and
FERGUSON ET AL.: CLIMATE CHANGE AND SEAL RECRUITMENT 123 Figure 1. Study area in western Hudson Bay. Communities from which seals were harvested and from which weather stations provided environmental data for this study are indicated. Box denotes area used to determine sea ice break-up dates and dotted line represents approximate area hunted. evaluate several environmental factors that may affect the quality of natal dens in relation to recruitment of seals into the population. The amount of snowfall in April and May is likely important, and may correlate with factors such as the predation rate (negative) on seal pups by polar bears (Hammill and Smith 1991), depth of drifts with subnivean birth lairs (positive), or rainfall (negative) (Stirling and Smith 2004). In addition to predation risk, survival of pups in April and May may be influenced by the negative effects of exposure, including hypothermia (Smith et al. 1991, Kelly 2001), in the event of den collapse. Our goal was to determine whether annual variation in the survivorship curve was correlated with these environmental factors. METHODS Study Area Our study area encompassed the western Hudson Bay region (Fig. 1). Hudson Bay is a relatively shallow salt-water body with counter clockwise currents (Prinsenberg 1986, Saucier et al. 2004). Hudson Bay is covered by annual ice from mid-november to May and in some years can be almost free of ice by late June (Collin and Dunbar 1964). Although shore leads and polynyas are present throughout the winter, spring break-up generally begins in April with openings observed in the Roes Welcome Sound polynya to the southwest of Southampton Island (Stirling 1997). Ice decays rapidly to the south and southeast because of
134 MARINE MAMMAL SCIENCE, VOL. 21, NO. 1, 2005 KINGSLEY, M. C. S., AND T. J. BYERS. 1998. Failure of reproduction in ringed seals (Phoca hispida) in Amundsen Gulf, Northwest Territories in 1984 1987. Pages 197 210 in M. P. Heide-Jørgensen and C. Lydersen, eds. Ringed seals in the North Atlantic. Volume 1. NAMMCO Scientific Publications, Tromsø, Norway. KRAJICK, K. 2001. Arctic life, on thin ice. Science 291:424 425. LEVITUS, S., J. I. ANTONOV, J. WANG, T. L. DELWORTH, K. W. DIXON AND A. J. BROCCOLI. 2001. Anthropogenic warming of Earth s climate system. Science 292:267 270. LOWRY, L. F., K. J. FROST AND J. J. BURNS. 1980. Variability in the diet of ringed seals, Phoca hispida, in Alaska. Canadian Journal of Fisheries and Aquatic Science 37:2254 2261. LUNN, N. J., I. STIRLING AND S. N. NOWICKI. 1997. Distribution and abundance of ringed (Phoca hispida) and bearded seals (Erignathus barbatus) in western Hudson Bay. Canadian Journal of Fisheries and Aquatic Science 54:914 921. LYDERSEN, C., AND M. O. HAMMILL. 1993a. Diving in ringed seal (Phoca hispida) pups during the nursing period. Canadian Journal of Zoology 71:991 996. LYDERSEN, C., AND M. O. HAMMILL. 1993b. Activity, milk intake and energy consumption in free-living ringed seal (Phoca hispida) pups. Journal of Comparative Physiology B Biochemical Systematic and Environmental Physiology 163:433 438. MCLAREN, I. A. 1958. The biology of the ringed seal (Phoca hispida Schreber) in the eastern Canadian Arctic. Bulletin Fisheries Research Board of Canada 118:1 97. MCLAREN, I. A. 1993. Growth in pinnipeds. Biological Review 6:1 79. MOULTON, V. D., W. J. RICHARDSON, T. L. MCDONALD, R. E. ELLIOTT AND M. T. WILLIAMS. 2002. Factors influencing local abundance and haulout behaviour of ringed seals (Phoca hispida) on landfast ice of the Alaskan Beaufort Sea. Canadian Journal of Zoology 80:1900 1917. MUNK, W. 2003. Ocean freshening, sea level rising. Science 300:2041 2043. MYSAK, L. A., R. G. INGRAM, J.WANG AND A. VAN DER BAAREN. 1996. The anomalous seaice extent in Hudson Bay, Baffin Bay and the Labrador Sea during three simultaneous NAO and ENSO episodes. Atmospheric Ocean 34:313 343. PARKINSON, C. L. 2000. Variability of Arctic sea ice: The view from space, an 18-year record. Arctic 53:341 358. PRINSENBERG, S. J. 1986. Man-made changes in the freshwater input rates of Hudson and James bays. Canadian Journal of Fisheries and Aquatic Science 37:1101 1110. RYG, M., T. G. SMITH AND N. A. ØRITSLAND. 1990. Seasonal changes in body mass and body consumption of ringed seals (Phoca hispida) on Svalbard. Canadian Journal of Zoology 68:470 475. SAUCIER, F. J., S. SENNEVILLE, S. PRINSENBERB, F. ROY, G. SMITH, P. GACHON, D. CAYA, AND R. LAPRISE. 2004. Modelling the sea ice-ocean seasonal cycle in Hudson Bay, Foxe Basin and Hudson Strait, Canada. Climate Dynamics 23:303 326. SKINNER, W. R., R. L. JEFFERIES,T.J.CARLETON,R.F.ROCKWELL AND K. F. ABRAHAM. 1998. Prediction of reproductive success and failure in lesser snow geese based on early season climatic variables. Global Change Biology 4:3 16. SMITH, T. G. 1973. Population dynamics of the ringed seal in the Canadian eastern Arctic. Bulletin Fisheries Research Board of Canada 181:1 55. SMITH, T. G. 1975. Ringed seals in James Bay and Hudson Bay: population estimates and catch statistics. Arctic, 28:170 182. SMITH, T. G. 1980. Polar bear predation of ringed and bearded seals in the land-fast sea ice habitat. Canadian Journal of Zoology 58:2201 2209. SMITH, T. G. 1987. The ringed seal, Phoca hispida, of the Canadian western Arctic. Canadian Bulletin of Fisheries and Aquatic Science 216:1 81. SMITH, T. G., AND L. A. HARWOOD. 2001. Observations of neonate ringed seals, Phoca hispida, after early break-up of the sea ice in Prince Albert Sound, Northwest Territories, Canada, spring 1998. Polar Biology 24:215 219. SMITH, T. G., AND C. LYDERSEN. 1991. Availability of suitable land-fast ice and predation as factors limiting ringed seal populations, Phoca hispida, in Svalbard. Pages 585 594 in
FERGUSON ET AL.: CLIMATE CHANGE AND SEAL RECRUITMENT 135 E. Sakshaug, C. C. E. Hopkins and N. A. Øritsland, eds. Proceedings of the Pro Mare Symposium on Polar Marine Ecology, Trondheim, 12 16 May 1990. SMITH, T. G., AND I. STIRLING. 1975. The breeding habitat of the ringed seal (Phoca hispida). The birth lair and associated structures. Canadian Journal of Zoology 53:1297 1305. SMITH, T. G., M. O. HAMMILL AND G. TAUGBØL. 1991. A review of the developmental, behavioral, and physiological adaptations of the ringed seal, Phoca hispida, to life in the arctic winter. Arctic 44:124 131. SOKAL, R., AND F. ROHLF. 1981. Biometry. W. H. Freeman and Co., New York, NY. STERN, H. L. AND M. P. HEIDE-JORGENSEN. 2003. Trends and variability of sea ice in Baffin Bay and Davis Strait, 1953 2001. Polar Research 22:11 18. STEWART, R. E. A., B. E. STEWART, I.STIRLING AND E. STREET. 1996. Counts of growth layer groups in cementum and dentine in ringed seals (Phoca hispida). Marine Mammal Science 12:383 401. STIRLING, I. 1997. The importance of polynyas, ice edges, and leads to marine mammals and birds. Journal of Marine Systems 10:9 21. STIRLING, I., AND W. R. ARCHIBALD. 1977. Aspects of predation of seals by polar bears in the eastern Beaufort Sea. Journal of Fisheries Research Board of Canada 34:1126 1129. STIRLING, I., N. J. LUNN AND J. IACOZZA. 1999. Long-term trends in the population ecology of polar bears in western Hudson Bay in relation to climate changes. Arctic 52: 294 306. STIRLING, I., AND T. G. SMITH. 2004. Implications of warm temperatures and an unusual rain event on the survival of ringed seals on the coast of southeastern Baffin Island. Arctic 57:59 67. TYNAN, C. T., AND D. P. DEMASTER. 1997. Observations and predictions of Arctic climatic change: Potential effects on marine mammals. Arctic 50:308 322. VINJE, T. 2001. Fram Strait ice fluxes and atmospheric circulation: 1950 2000. Journal of Climate 14:3508 3517. Received: 12 December 2003 Accepted: 20 July 2004