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Direction and magnitude of natural selection on body size differ among age-classes of seaward-migrating Pacific salmon.

ABSTRACT: Due to the mediating role of body size in determining fitness, the "bigger-is-better" hypothesis still pervades evolutionary ecology despite evidence that natural selection on phenotypic traits varies in time and space. For Pacific salmon (genus Oncorhynchus), most individual studies quantify selection across a narrow range of sizes and ages; therefore, uncertainties remain concerning how selection on size may differ among diverse life histories. Here, we quantify the direction and magnitude of natural selection on body size among age-classes of multiple marine cohorts of O. nerka (sockeye salmon). Across four cohorts of seaward migrants, we calculated standardized selection differentials by comparing observed size distributions of out-migrating juvenile salmon to back-calculated smolt length from the scales of surviving, returning adults. Results reveal the magnitude of selection on size was very strong (>90th percentile compared to a database of 3,759 linear selection differentials) and consistent among years. However, the direction of selection on size consistently varied among age-classes. Selection was positive for fish migrating to sea after two years in freshwater (age 2) and in their first year of life (age 0), but negative for fish migrating after 1 year in freshwater (age 1). The absolute magnitude of selection was negatively correlated to mean ocean-entry timing, which may underpin negative selection favoring small age-1 fish, given associations between size and timing of seaward migration. Collectively, these results indicate that "bigger is not always better" in terms of survival and emphasize trade-offs that may exist between fitness components for organisms with similarly diverse migratory life histories.

SUBMITTER: Ulaski ME

PROVIDER: S-EPMC7463379 | biostudies-literature | 2020 Sep

REPOSITORIES: biostudies-literature

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Direction and magnitude of natural selection on body size differ among age-classes of seaward-migrating Pacific salmon.

Ulaski Marta E ME Finkle Heather H Westley Peter A H PAH

Evolutionary applications 20200409 8

Due to the mediating role of body size in determining fitness, the "bigger-is-better" hypothesis still pervades evolutionary ecology despite evidence that natural selection on phenotypic traits varies in time and space. For Pacific salmon (genus <i>Oncorhynchus</i>), most individual studies quantify selection across a narrow range of sizes and ages; therefore, uncertainties remain concerning how selection on size may differ among diverse life histories. Here, we quantify the direction and magnit ...[more]

PMID: 32908600

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Project description:Ecologists are pressed to understand how climate constrains the timings of annual biological events (phenology). Climate influences on phenology are likely significant in estuarine watersheds because many watersheds provide seasonal fish nurseries where juvenile presence is synched with favorable conditions. While ecologists have long recognized that estuaries are generally important to juvenile fish, we incompletely understand the specific ecosystem dynamics that contribute to their nursery habitat value, limiting our ability to identify and protect vital habitat components. Here we examined the annual timing of juvenile coldwater fish migrating through a seasonally warm, hydrologically managed watershed. Our goal was to (1) understand how climate constrained the seasonal timing of water conditions necessary for juvenile fish to use nursery habitats and (2) inform management decisions about (a) mitigating climate-mediated stress on nursery habitat function and (b) conserving heat-constrained species in warming environments. Cool, wet winters deposited snow and cold water into mountains and reservoirs, which kept the lower watershed adequately cool for juveniles through the spring despite the region approaching its hot, dry summers. For every 1°C waters in April were colder, the juvenile fish population (1) inhabited the watershed 4-7 d longer and (2) entered marine waters, where survival is size selective, at maximum sizes 2.1 mm larger. Climate therefore appeared to constrain the nursery functions of this system by determining seasonal windows of tolerable rearing conditions, and cold water appeared to be a vital ecosystem component that promoted juvenile rearing. Fish in this system inhabit the southernmost extent of their range and already rear during the coolest part of the year, suggesting that a warming climate will truncate rather than shift their annual presence. Our findings are concerning for coldwater diadromous species in general because warming climates may constrain watershed use and diminish viability of life histories (e.g., late springtime rearing) and associated portfolio benefits over the long term. Lower watershed nurseries for coldwater fish in warming climates may be enhanced through allocating coldwater reservoir releases to prolong juvenile rearing periods downstream or restorations that facilitate colder conditions.

Project description:Size-selective mortality owing to lack of energy reserves during the first marine winter has been suggested to be a result of juvenile salmon failing to reach a critical size or condition by the end of their first marine summer and not surviving the following winter due to this presumed energy deficit. This hypothesis implies strong size dependency of mortality, and is subject to empirical data support for acceptance. Scale circulus spacing has been interpreted as an index for body size, and we reviewed the effect of size-selective mortality with a knife-edge mortality function on descriptive statistics for a scale circulus spacing index (SCSI). In order to invoke size selection as an important driver of mortality during the first year of ocean rearing, it is necessary to demonstrate not only that size-selective mortality is directed towards the smaller members of the population, but that the selective nature of the mortality can account for a substantial portion of the observed mortality. If the assumption is made that a random sample of a single juvenile population has been obtained, then studies that employ a SCSI to infer size-selective mortality coupled with a critical size limit must demonstrate a shift toward larger values of the SCSI, but also a concomitant reduction in the variance and range of the SCSI and an increase in the skewness and kurtosis of the SCSI values. Through simulation we found that the percentage of adults that displayed a SCSI value greater than the maximum observed in the juvenile sample was highly dependent on the initial juvenile sample size and size-selective mortality rate. Geographical distributions of juvenile Pacific salmon can be stratified by size, with larger individuals migrating earlier from local ocean entry locations than smaller individuals, and thus differential timing migration of juveniles based upon body size prior to the collection of the marine juvenile sample may be a more plausible explanation of published trends in the SCSI, rather than invoking substantial size-selective mortality and a critical size limit.

Dataset Information

Direction and magnitude of natural selection on body size differ among age-classes of seaward-migrating Pacific salmon.

Publications

Direction and magnitude of natural selection on body size differ among age-classes of seaward-migrating Pacific salmon.

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