Climate change and control of the southeastern Bering Sea pelagic ecosystem

Abstract

We propose a new hypothesis, the Oscillating Control Hypothesis (OCH), which predicts that pelagic ecosystem function in the southeastern Bering Sea will alternate between primarily bottom-up control in cold regimes and primarily top-down control in warm regimes. The timing of spring primary production is determined predominately by the timing of ice retreat. Late ice retreat (late March or later) leads to an early, ice-associated bloom in cold water (e.g., 1995, 1997, 1999), whereas no ice, or early ice retreat before mid-March, leads to an open-water bloom in May or June in warm water (e.g., 1996, 1998, 2000). Zooplankton populations are not closely coupled to the spring bloom, but are sensitive to water temperature. In years when the spring bloom occurs in cold water, low temperatures limit the production of zooplankton, the survival of larval/juvenile fish, and their recruitment into the populations of species of large piscivorous fish, such as walleye pollock (Theragra chalcogramma), Pacific cod (Gadus macrocephalus) and arrowtooth flounder (Atheresthes stomias). When continued over decadal scales, this will lead to bottom-up limitation and a decreased biomass of piscivorous fish. Alternatively, in periods when the bloom occurs in warm water, zooplankton populations should grow rapidly, providing plentiful prey for larval and juvenile fish. Abundant zooplankton will support strong recruitment of fish and will lead to abundant predatory fish that control forage fish, including, in the case of pollock, their own juveniles. Piscivorous marine birds and pinnipeds may achieve higher production of young and survival in cold regimes, when there is less competition from large piscivorous fish for coldwater forage fish such as capelin (Mallotus villosus). Piscivorous seabirds and pinnipeds also may be expected to have high productivity in periods of transition from cold regimes to warm regimes, when young of large predatory species of fish are numerous enough to provide forage. The OCH predicts that the ability of large predatory fish populations to sustain fishing pressure will vary between warm and cold regimes. The OCH points to the importance of the timing of ice retreat and water temperatures during the spring bloom for the productivity of zooplankton, and the degree and direction of coupling between zooplankton and forage fish. Forage fish (e.g., juvenile pollock, capelin, Pacific herring [Clupea pallasii]) are key prey for adult pollock and other apex predators. In the southeastern Bering Sea, important changes in the biota since the mid-1970s include a marked increase