Beyond Bergmann's rule: Global variability in human body composition is associated with annual average precipitation...

Human populations exhibit substantial geographical variability in body size and shape. However, the ecological stresses underlying this morphological variability remain poorly understood. The prevailing evolutionary explanation, "Bergmann's rule" assumes that morphological variability represents an adaptive response to average thermal conditions. We hypothesized that other climate factors-annual average precipitation, a marker of ecological productivity and inter-annual temperature volatility, a marker of infectious disease spikes-may also contribute to variability in body composition.

We explored this hypothesis by examining associations between these climate factors and geographic variability in body composition across 133 male and 105 female populations from nonindustrialized settings. We used monthly climate data over 113 years (1901-2013) to develop new climate indices for all worldwide land areas. We stratified our analyses by hot/cold setting (>/<20°C).

In hot environments, lean mass increased as predicted in association with ecological productivity, and decreased in association with ecological volatility. Conversely, levels of body fat increased in association with temperature volatility and precipitation. However, in cold settings, equivalent associations were only partially consistent with our hypotheses, and there was suggestive evidence of sex differences in these associations.

Beyond associations with mean annual temperature predicted by Bergmann's rule, variability in human body composition is also associated with mean annual temperature and inter-annual temperature volatility, with these associations further differing between hot and cold settings. Collectively, our results suggest that associations of human body composition with climate are complex for both physique (fat-free mass) and energy stores (adiposity).