Modeling haul-out behavior of walruses in Bering Sea ice

Understanding haul-out behavior of ice-associated pinnipeds is essential for designing and interpreting popula-tion surveys and for assessing effects of potential changes in their ice environments. We used satellite-linked transmitters to obtain sequential information about location and haul-out state for Pacific walruses, Odobenus rosmarus divergens (Il-liger, 1815), in the Bering Sea during April of 2004, 2005, and 2006. We used these data in a generalized mixed model of haul-out bout durations and a hierarchical Bayesian model of haul-out probabilities to assess factors related to walrus haul-out behavior, and provide the first predictive model of walrus haul-out behavior in sea ice habitat. Average haul-out bout duration was 9 h, but durations of haul-out bouts tended to increase with durations of preceding in-water bouts. On aver-age, tagged walruses spent only about 17% of their time hauled out on sea ice. Probability of being hauled out decreased with wind speed, increased with temperature, and followed a diurnal cycle with the highest values in the evening. Our haul-out probability model can be used to estimate the proportion of the population that is unavailable for detection in spring surveys of Pacific walruses on sea ice.