We tested hypotheses on how animals should respond to heterospecifics encountered in the environment. Hypotheses were formulated from models parameterized to emphasize four factors that are expected to influence species discrimination: mating and territorial interactions; sex differences in resource value; environments in which heterospecifics were common or rare; and the type of identity cues available for species recognition. We also considered the role of phylogeny on contemporary responses to heterospecifics. We tested the extent these factors explained variation among taxa in species discrimination using a meta-analysis of three decades of species recognition research. A surprising outcome was the absence of a general predictor of when species discrimination would most likely occur. Instead, species discrimination is dictated by the benefits and costs of responding to a conspecific or heterospecific that are governed by the specific circumstances of a given species. The phylogeny of species recognition provided another unexpected finding: the evolutionary relationships among species predicted whether courting males within species – but not females – would discriminate against heterospecifcs. This implies that species recognition has evolved quite differently in the sexes. Finally, we identify common pitfalls in experimental design that seem to have affected some studies (e.g., poor statistical power) and provide recommendations for future research.