Evolution of age and length at maturation of Alaskan salmon under size-selective harvest.

Spatial and temporal trends and variation in life-history traits, including age and length at maturation, can be influenced by environmental and anthropogenic processes, including size-selective exploitation. Spawning adults in many wild Alaskan sockeye salmon populations have become shorter at a given age over the past half-century, but their age composition has not changed. These fish have been exploited by a gillnet fishery since the late 1800s that has tended to remove the larger fish. Using a rare, long-term dataset, we estimated probabilistic maturation reaction norms (PMRNs) for males and females in nine populations in two basins and correlated these changes with fishery size selection and intensity to determine whether such selection contributed to microevolutionary changes in maturation length. PMRN midpoints decreased in six of nine populations for both sexes, consistent with the harvest. These results support the hypothesis that environmental changes in the ocean (likely from competition) combined with adaptive microevolution (decreased PMRNs) have produced the observed life-history patterns. PMRNs did not decrease in all populations, and we documented differences in magnitude and consistency of size selection and exploitation rates among populations. Incorporating evolutionary considerations and tracking further changes in life-history traits can support continued sustainable exploitation and productivity in these and other exploited natural resources.

Quantifying and comparing size selectivity among Alaskan sockeye salmon fisheries.

Quantifying long-term size-selective harvest patterns is necessary for understanding the potential evolutionary effects on exploited species. The comparison of fishery selection patterns on the same species subject to different gear types, in different areas, and over multi-decadal periods can reveal the factors influencing selection. In this study we quantified and compared size-selective harvest by nine Alaskan sockeye salmon (Oncorhynchus nerka) fisheries to understand overall patterns. We calculated length-specific linear selection differentials (the difference in average length of fish before vs. after fishing), which are produced by different combinations of exploitation rates and length-selectivity values, and nonlinear standardized differentials, describing disruptive selection, across all years for each fishery. Selection differentials varied among years, but larger fish were caught in 73% of years for males and 84% of years for females, leaving smaller fish to spawn. Disruptive selection was observed on female and male fish in 84% and 92% of years, respectively. Linear selection was stronger on females than males in 77% of years examined, and disruptive selection was stronger on males in 71% of years. Selection pressure was influenced by a combination of factors under and beyond management control; analyses using mixed-effects models indicated that fisheries were less size selective in years when fish were larger than average and had lower exploitation rates. The observed harvest of larger than average sockeye salmon is consistent with the hypothesis that size-selective fishing contributes to decreasing age and length at maturation trends over time, but temporal variability in selection and strong disruptive selection suggests that the overall directional pressure is weaker than is often assumed in evolutionary models.

Selection due to nonretention mortality in gillnet fisheries for salmon.

Fisheries often exert selective pressures through elevated mortality on a nonrandom component of exploited stocks. Selective removal of individuals will alter the composition of a given population, with potential consequences for its size structure, stability and evolution. Gillnets are known to harvest fish according to size. It is not known, however, whether delayed mortality due to disentanglement from gillnets exerts selective pressures that reinforce or counteract harvest selection. We examined gillnet disentanglement in exploited populations of sockeye salmon (Oncorhynchus nerka) in Bristol Bay, Alaska, to characterize the length distribution of fish that disentangle from gillnets and determine whether nonretention mortality reinforces harvest selection and exerts common pressures according to sex and age. We also evaluated discrete spawning populations to determine whether nonretention affects populations with different morphologies in distinct ways. In aggregate, nonretention mortality in fish that disentangle from gillnets counters harvest selection but with different effects by sex and age. At the level of individual spawning populations, nonretention mortality may exert stabilizing, disruptive, or directional selection depending on the size distribution of a given population. Our analyses suggest nonretention mortality exerts significant selective pressures and should be explicitly included in analyses of fishery-induced selection.

Quantifying six decades of fishery selection for size and age at maturity in sockeye salmon.

Life history traits of wild animals can be strongly influenced, both phenotypically and evolutionarily, by hunting and fishing. However, few studies have quantified fishery selection over long time periods. We used 57 years of catch and escapement data to document the magnitude of and trends in gillnet selection on age and size at maturity of a commercially and biologically important sockeye salmon stock. Overall, the fishery has caught larger fish than have escaped to spawn, but selection has varied over time, becoming weaker and less consistent recently. Selection patterns were strongly affected by fish age and sex, in addition to extrinsic factors including fish abundance, mesh size regulations, and fish length variability. These results revealed a more complex and changing pattern of selective harvest than the 'larger is more vulnerable' model, emphasizing the need for quantified, multi-year studies before conclusions can be drawn about potential evolutionary and ecological effects of fishery selection. Furthermore, the results indicate that biologically robust escapement goals and prevention of harvest of the largest individuals may help prevent negative effects of size-selective harvest.