The biophysics of Bergmann's rule: a comparison of the effects of pelage and body size variation on metabolic rate
Should an animal extending its range into a cooler climate rely most on pelage or on body size change to minimize its mass-specific metabolic rate? The various examples of animals following Bergmann's rule support the latter. The fact that an increase in size will result in an increase in total metabolic rate (though coupled to the decrease in the mass-specific metabolic rate) suggests that increases in the insulative value of the pelage would be the preferred strategy. We used a thermal simulation model to compare the relative effects of increasing body mass versus increasing pelage insulative properties on the mass-specific metabolic rate. We found that even the fur of summer-adapted small mammals from temperate climates is extremely dense compared with that of larger mammals and is near the point at which increases in density increase, rather than decrease, heat loss as a result of the high conductivity of individual hairs compared with the layer of still air that it encloses. Small mammals also have lower fur depths, presumably as a result of biomechanical constraints. Seasonal changes in pelage observed in small mammals have very modest effects on mass-specific metabolism. Summer-adapted temperate large mammals, however, are less heavily insulated and, consequently, have substantial latitude for increasing insulation as a means of minimizing mass-specific metabolism. Thus, Bergmann's rule should be more relevant to small mammals than to large ones.