Allen's rule is an ecogeographical rule formulated by Joel Asaph Allen in 1877, broadly stating that animals adapted to cold climates have shorter and thicker limbs and bodily appendages than animals adapted to warm climates. More specifically, it states that the body surface-area-to-volume ratio for homeothermic animals varies with the average temperature of the habitat to which they are adapted (i.e. the ratio is low in cold climates and high in hot climates). Explanation Allen's rule predicts that endothermic animals with the same body volume should have different surface areas that will either aid or impede their heat dissipation. Because animals living in cold climates need to conserve as much heat as possible, Allen's rule predicts that they should have evolved comparatively low surface area-to-volume ratios to minimize the surface area by which they dissipate heat, allowing them to retain more heat. For animals living in warm climates, Allen's rule predicts the opposite: that they should have comparatively high ratios of surface area to volume. Because animals with low surface area-to-volume ratios would overheat quickly, animals in warm climates should, according to the rule, have high surface area-to-volume ratios to maximize the surface area through which they dissipate heat. In animals Though there are numerous exceptions, many animal populations appear to conform to the predictions of Allen's rule. The polar bear has stocky limbs and very short ears that are in accordance with the predictions of Allen's rule. In 2007, R.L. Nudds and S.A. Oswald studied the exposed lengths of seabirds' legs and found that the exposed leg lengths were negatively correlated with Tmaxdiff (body temperature minus minimum ambient temperature), supporting the predictions of Allen's rule. J.S. Alho and colleagues argued that tibia and femur lengths are highest in populations of the common frog that are indigenous to the middle latitudes, consistent with the predictions of Allen's rule for ectothermic organisms. Populations of the same species from different latitudes may also follow Allen's rule. R.L. Nudds and S.A. Oswald argued in 2007 that there is poor empirical support for Allen's rule, even if it is an "established ecological tenet". They said that the support for Allen's rule mainly draws from studies of single species, since studies of multiple species are "confounded" by the scaling effects of Bergmann's rule and alternative adaptations that counter the predictions of Allen's rule. J.S. Alho and colleagues argued in 2011 that, although Allen's rule was