The responses of salmonid fishes to the problems posed by marginal hab
itats are genetic exercises in population insurance. The costs increas
e as the risks increase, but the risks are met by a wide repertoire of
biological capacities. The most general proximate response to adversi
ty is behavioral: ontogenetic niche shifts are an acknowledgment that
a series of environments becomes marginal for all salmonids during dev
elopment. Physiological tolerances and developmental flexibility gover
n the timing of these movements. Such shifts are the product of natura
l selection in relatively predictable environments, but less predictab
le or catastrophic events are accommodated at a different genetic leve
l. Low spatial flexibility is counteracted by temporal insurance, and
vice versa. Thus, high homing precision is coupled with complex multip
le-age structures, and simple age structuring is coupled with relative
ly high spatial straying