Sea-Age Variation in Maiden Atlantic Salmon Spawners: Phenotypic Plasticity or Genetic Polymorphism?

Atlantic salmon exhibit a partially heritable polymorphism in which the morphs are distinguished by the duration and location of the sea-phase of their life-cycle. These morphs co-occur, albeit in characteristically different proportions, in most Scottish rivers and in both the spring and autumn spawner runs; early running fish being generally associated with upland spawning locations while late running fish are associated with lowland spawning. Thus, differences in riverine and marine environment appear to be linked to differences in the relative abundance of the morphs, rather than to the specific morph which is optimally adapted. In this paper, we report a model-based synthetic study aimed at understanding the key dynamic elements which determine the long-term stability of this polymorphism, and thus determine the relative abundance of the various sea-age morphs. Given the recent accumulation of evidence for a genetic basis for the polymorphism, we argue that the key dynamic mechanism which equalises the realized fitness of the sea-age morphs must be one or more morph-specific density dependencies in the riverine phase of the life-history. We explore a number of specific mechanisms, firmly based in known salmon biology, by which such morph-specific density dependence could occur and investigate the robustness of the co-existence which they imply. We conclude that the co-occurrence of multiple sea-age morphs of Atlantic salmon in Scottish rivers is a stable genetic polymorphism, maintained by some combination of physical separation and asymmetric competition between spawners of different morphs or the riverine stages of their offspring or both.