No abstract

Farm Atlantic salmon escape and invade rivers throughout the North Atlantic annually, which has generated growing concern about their impacts on native salmon populations. A large-scale experiment was therefore undertaken in order to quantify the lifetime success and interactions of farm salmon invading a Norwegian river. Sexually mature farm and native salmon were genetically screened, radio tagged and released into the River Imsa where no other salmon had been allowed to ascend. The farm ¢shes were competitively and reproductively inferior, achieving less than one-third the breeding success of the native ¢shes. Moreover, this inferiority was sex biased, being more pronounced in farm males than females, resulting in the principal route of gene £ow involving native males mating with farm females. There were also indications of selection against farm genotypes during early survival but not thereafter. However, evidence of resource competition and competitive displacement existed as the productivity of the native population was depressed by more than 30%. Ultimately, the lifetime reproductive success (adult to adult) of the farm ¢shes was 16% that of the native salmon. Our results indicate that such annual invasions have the potential for impacting on population productivity, disrupting local adaptations and reducing the genetic diversity of wild salmon populations.

The magnitude of ¢tness variation caused by maternal e¡ects and, thus, the adaptive signi¢cance of maternal traits may depend on environmental quality, generating crossing reaction norms among o¡spring phenotypes that shape life-history evolution. By manipulating intraclutch variation in egg size and comparing siblings we examined the maternal e¡ects of egg size on o¡spring performance and tested for the existence of reaction norms to environmental quality using the brown trout Salmo trutta. When sibling groups of small and large eggs were reared separately in a hatchery environment initial size di¡er-ences disappeared rapidly. However, in semi-natural environments and under direct competition, juveniles from large eggs experienced growth and survival advantages over siblings from small eggs. Moreover, distinct reaction norms existed, with the di¡erences in performance of juveniles from small and large eggs being most pronounced in the poorer growth environments. Our results provide the ¢rst direct evidence, to our knowledge, for a causal relationship between egg size and ¢tness-related traits in ¢shes, independent of potentially confounding genetic e¡ects. Moreover, they indicate that previous studies have been biased by experimental conditions that excluded competitive asymmetries and environmental variability. The existence of reaction norms indicates a shift in optimal egg size across gradients of environmental quality that probably shapes the evolution of this trait.

Here we critically review the scale and extent of adaptive genetic variation in Atlantic salmon (Salmo salar L.), an important model system in evolutionary and conservation biology that provides fundamental insights into population persistence, adaptive response and the effects of anthropogenic change. We consider the process of adaptation as the end product of natural selection, one that can best be viewed as the degree of matching between phenotype and environment. We recognise three potential sources of adaptive variation: heritable variation in phenotypic traits related to fitness, variation at the molecular level in genes influenced by selection, and variation in the way genes interact with the environment to produce phenotypes of varying plasticity. Of all phenotypic traits examined, variation in body size (or in correlated characters such as growth rates, age of seaward migration or age at sexual maturity) generally shows the highest heritability, as well as a strong effect on fitness. Thus, body size in Atlantic salmon tends to be positively correlated with freshwater and marine survival, as well as with fecundity, egg size, reproductive success, and offspring survival. By contrast, the fitness implications of variation in behavioural traits such as aggression, sheltering behaviour, or timing of migration are largely unknown. The adaptive significance of molecular variation in salmonids is also scant and largely circumstantial, despite extensive molecular screening on these species. Adaptive variation can result in local adaptations (LA) when, among other necessary conditions, populations live in patchy environments, exchange few or no migrants, and are subjected to differential selective pressures. Evidence for LA in Atlantic salmon is indirect and comes mostly from ecological correlates in fitness-related traits, the failure of many translocations, the poor performance of domesticated stocks, results of a few common-garden experiments (where different populations were raised in a common environment in an attempt to dissociate heritable from environmentally induced phenotypic variation), and the pattern of inherited resistance to some parasites and diseases. Genotype x environment interactions occurr for many fitness traits, suggesting that LA might be important. However, the scale and extent of adaptive variation remains poorly understood and probably varies, depending on habitat heterogeneity, environmental stability and the relative roles of selection and drift. As maladaptation often results from phenotype-environment mismatch, we argue that acting as if populations are not locally adapted carries a much greater risk of mismanagement than acting under the assumption for local adaptations when there are none. As such, an evolutionary approach to salmon conservation is required, aimed at maintaining the conditions necessary for natural selection to operate most efficiently and unhindered. This may require minimising alterations to native genotypes and habitats to which populations have likely become ad...

The breeding system of Atlantic salmon (Salmo salar) is shaped both by natural selection for offspring production and by sexual selection for access to mating opportunities. These evolutionary forces operate with differing intensities in the two sexes to shape their breeding behaviour and tactics. Female breeding success is largely dependent on egg production, access to breeding territories, and nest quality and survival. By contrast, male breeding success is largely determined by access to ovipositing females. As such, the breeding system of Atlantic salmon is similar to that of other members of the subfamily Salmoninae. However, early male maturity, a common pattern within the Salmoninae, reaches its greatest expression in both terms of frequency and magnitude of the mature male size difference in Atlantic salmon. Despite generalities, spawning populations of Atlantic salmon are not static, as they exhibit spatial and temporal variability in demography (e.g., spawner density, sex ratio, age at maturity, and body size). Events, both natural and anthropogenic (e.g., exploitation, habitat alteration, and climatic changes), affect this variability and ultimately shape the breeding system.

Why do highly fecund organisms apparently sacrifice offspring size for increased numbers when offspring survival generally increases with size? The theoretical tools for understanding this evolutionary trade-off between number and size of offspring have developed over the past 25 years; however, the absence of data on the relation between offspring size and fitness in highly fecund species, which would control for potentially confounding variables, has caused such models to remain largely hypothetical. Here we manipulate egg size, controlling for maternal trait interactions, and determine the causal consequences of offspring size in a wild population of Atlantic salmon. The joint effect of egg size on egg number and offspring survival resulted in stabilizing phenotypic selection for an optimal size. The optimal egg size differed only marginally from the mean value observed in the population, suggesting that it had evolved mainly in response to selection on maternal rather than offspring fitness. We conclude that maximization of maternal fitness by sacrificing offspring survival may well be a general phenomenon among highly fecund organisms.

The latitudinal variation in clutch size found in many animal species, including Pacific salmon, has been an enigmatic problem in ecology. We analyze egg number and egg size of 17 populations of coho salmon (Oncorhynchus kisutch) distributed over a latitudinal gradient in North America. These populations have a significant latitudinal increase in their egg number. But this increase is accompanied by a significant latitudinal decrease in their egg size. The total biomass of eggs produced also declines with latitude. Thus, the positive latitudinal trend in egg number cannot be explained by latitudinal variation in the total investment in eggs. This suggests that local optima in egg size may result in latitudinal clines in egg number. This reasoning, that egg number evolves around selection for egg size, is in fact predicted by life history theory and may explain the clutch size patterns observed in many organisms.

In the breeding system of Pacific salmon, females compete for oviposition territories, and males compete to fertilize eggs. The natural selection in females and sexual selection in males likely has been responsible for their elaborate breeding morphologies and the dimorphism between the sexes. We quantified direct-selection intensities during breeding on mature coho salmon (Oncorhynchus kisutch), measured for seven phenotypic characters, including three secondary sexual characters. Wild and sea-ranched hatchery coho were used to enhance the range of phenotypes over which selection could be examined. The fish were allowed to breed in experimental arenas where we could quantify components of breeding success as well as estimate overall breeding success. We found that without competition, natural selection acts only on female body size for increased egg production; there is no detectable selection on males for the phenotypic distribution we used. Under competition, the opportunity for selection increased sixfold among females. Natural selection favored female body size and caudal-peduncle (tail) depth. Increased body size meant increased egg production and access to nesting territories. The caudal peduncle, used in burst swimming and nest digging, influenced both successful egg deposition and nest survival. Increasing density increased competition among females, though it did not significantly intensify natural selection on their characters. In males, competition increased the opportunity for selection 52-fold, which was nine times greater than for females. Sexual selection favored male body size and hooked snout length, both characters directly influencing male access to spawning opportunities. Selection on male body size was also affected significantly by breeding density. The ability of large males to control access to spawning females decreased at higher densities reflecting an increase in the operational sex ratio. Further, the relative success of small males, which could sneak access to spawning females, appeared to increase as that of intermediate-sized males decreased. Such disruptive selection may be responsible for the evolution of alternative reproductive tactics in salmon.