Body mass is estimated for extinct species of Sigmodon. These data are then used in appropriate equations derived among Recent mammals to estimate a suite of physiological and ecological variables which are followed through almost 4 ma of cotton rat history. A statistical trend towards large size is documented. Despite large swings in population size and other parameters, hypothetical values of population metabolism remain virtually constant, suggesting negligible population energetic benefit to size change in either direction. Studies of extant cotton rats and unrelated taxa sharing the same adaptive zone suggest that there is now, and has been in the past, negligible thermoregulatory advantage to modification of cotton rat body mass. Large size appears to be associated in Sigmodon with heightened aggression to the point that two species, particularly if of dichotomous size, cannot coexist in the same microhabitat. At least with regard to cotton rats, this conclusion represents a challenge to the comfortable hypotheses of coevolution and character displacement. The overall trend toward large size during Pleistocene time is considered then as the interplay of selection acting to favor large size in areas of sympatry and stochastic processes originating cotton rat populations of different body size. Because morphology and ecological strategies are stable within the cotton rat adaptive zone for millions of years, it is suggested that mammalian speciation events that result in exploitation of a new adaptive zone are uncommon and occasionally cross higher taxonomic categories. These events are defined as first-order speciation events, in contrast to second-order events that occur in clades within the same adaptive zone.