Summary: Chronic activation of the stress axis caused by long‐term uncontrollable and unpredictable factors in the environment has been regarded as causing maladaptive and/or pathological effects, both by those studying animals in the laboratory and in nature. While pathology may apply to the former, I argue that it does not apply to the latter. Our thinking on the role of chronic stress in animals in nature has been heavily influenced by biomedical research, but much less so by the ecological and evolutionary context within which animals actually function. I argue that when such stressors occur (e.g. periods of high predation risk, food limitation, prolonged severe weather, social conflict, etc.), although the animal may be chronically stressed, its responses are adaptive and continue to promote fitness. Chronic stressors in nature can be subdivided into whether they are reactive (direct physiological challenges threatening homeostasis and not requiring cognitive processing – for example, food limitation) or anticipatory (perceived to be threatening and requiring cognitive processing – for example, high predation risk). For anticipatory stressors, their impact on the animal should not be based on their absolute duration (they may be acute), but rather by the duration of their physiological consequences. The anticipatory stressor of persistent high predation risk does not elicit chronic stress in all prey classes. Cyclic snowshoe hare and arctic ground squirrels exhibit evidence of chronic stress when predator numbers are high, but cyclic vole and noncyclic elk populations do not. I suggest that chronic stress has evolved to benefit the fitness of the former and not the later, with the key factors being lifespan and life history. I propose that chronic stress evolves in a species only if it is adaptive.

Habitat constrains and shapes successful ecological and physiological strategies and thus provides the context for the evolution of life-history traits. The stress axis plays a vital role in the endocrine system and is a critical component adapting mammals to particular habitat pressures. It is subject to both individual activational and organizational plasticity as well as to evolutionary modification. To illustrate, I examine the suite of traits of the stress axis associated with breeding frequency in male mammals, which varies in a continuum from semelparity to iteroparity. During the breeding season, males in species at the semelparous end of the continuum exhibit high concentrations of free corticosteroids, low concentrations of glucocorticoid-binding protein, a failure of the negative feedback system, a gonadal axis that is not inhibited by high corticosteroid concentrations, and immunosuppression. Iteroparous species exhibit the opposite traits. The evolutionary constraints selecting for the former may partially be related to phylogeny (in marsupials) as well as to an interaction of the restrictions imposed by the environment on female reproduction, the mating system, the high costs of reproduction, and the low adult survival during the nonbreeding season.

Northern environments present ecological and physiological problems for homeotherms that require adaptations to cope with severe and less predictable physical factors while at the same time continuing to have to cope with the biological ones, such as competition and predation. The stress axis plays a central role in these adaptations and I discuss the range of solutions that birds and mammals have evolved. The stress response in these animals is not static when a challenge occurs, but may be modulated depending on the biological function during the annual cycle (breeding versus nonbreeding), either under-responding to permit reproduction (some song birds) or responding vigorously, yet not having this compromise reproduction (Arctic ground squirrels). Both may trade off survival for reproduction. In contrast, the snowshoe hare shows the expected stress response to chronic high predation risk over 2–3 years: body resources are geared to survival and reproduction is inhibited. Two long term, persistent, and pervasive changes will confront northern birds and mammals in the 21^st century: global change and persistent organochlorine pollutants (POPs). These may result in either adaptations or shifts in distribution and abundance. For the former, latitudinal variation in the stress axis may help song birds respond rapidly; population variation in the stress axis response is unknown in northern mammals and relatively sedentary mammals may be unable to shift their distribution rapidly to adjust major climate shifts. For the latter, the few POPs studies that have examined the stress axis indicate marked negative effects.