Winter mortality has been documented in a large number of freshwater fish populations, and a smaller, but increasing, number of marine and estuarine fishes. The impacted populations include a number of important North American and European resource species, yet the sources of winter mortality remain unidentified in most populations where it has been documented. Among the potential sources, thermal stress and starvation have received the most research attention. Other sources including predation and pathogens have significant impacts but have received insufficient attention to date. Designs of more recent laboratory experiments have reflected recognition of the potential for interactions among these co‐occurring stressors. Geographic patterns in winter mortality are, in some cases, linked to latitudinal clines in winter severity and variability. However, for many freshwater species in particular, the effects of local community structure (predators and prey) may overwhelm latitudinal patterns. Marine (and estuarine) systems differ from freshwater systems in several aspects important to overwintering fishes, the most important being the lack of isolating barriers in the ocean. While open population boundaries allow fish to adopt migration strategies minimizing exposure to thermal stresses, they may retard rates of evolution to local environments. Geographic patterns in the occurrence and causes of winter mortality are ultimately determined by the interaction of regional and local factors. Winter mortality impacts population dynamics through episodic depressions in stock size and regulation of annual cohort strength. While the former tends to act in a density‐independent manner, the latter can be density dependent, as most sources of mortality tend to select against the smallest members of the cohort and population. Most stock assessment and management regimes have yet to explicitly incorporate the variability in winter mortality. Potential management responses include postponement of cohort evaluation (to after first winter of life), harvest restrictions following mortality events and habitat enhancement. Future research should place more emphasis on the ecological aspects of winter mortality including the influences of food‐web structure on starvation and predation. Beyond illuminating an understudied life‐history phase, studies of overwintering ecology are integral to contemporary issues in fisheries ecology including ecosystem management, habitat evaluation, and impacts of climate change.
Pacific halibut (Hippoglossus stenolepis) are an ecologically, commercially, and culturally important Alaskan groundfish species. Commercial harvest of halibut dates back to the late 19th century and has been managed by the International Pacific Halibut Commission (IPHC) since 1921. IPHC surveys have revealed declining trends in survey biomass in multiple regions and region‐specific declines in mean size‐at‐age (size‐at‐age) over the past two decades (>50% in some areas). Changes in size‐at‐age can arise from a variety of physical, ecological, sampling, and fishery effects, including size‐dependent fishery or predation mortality, alteration in growth from variability in prey quality or quantity, and changes in temperature‐dependent metabolic demands. Here, we develop and apply a bioenergetics model for halibut using survey‐based diet and temperature data for Alaska to evaluate potential environmental drivers of size‐at‐age. In general, juvenile (<40 cm fork length) foraging rates were highest in the Gulf of Alaska concomitant with higher potential growth and elevated basal metabolic demands during warm summer conditions. In contrast, adult (40–120 cm FL) potential growth was highest in the Eastern Bering Sea, potentially reflecting lower metabolic costs and higher rates of prey consumption in that region. We additionally find evidence for interannual variation in potential growth, with a higher frequency of reduced growth potential in the last decade, particularly in the Eastern Bering Sea in 2015 and 2016 for both juvenile and adult halibut. These results suggest the potential for patterns in size‐at‐age to arise from trophic and environmental constraints that collectively limit growth in some regions and years.
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