Bordered tug-of-war models are neither general nor predictive of reproductive skew

Models of reproductive skew assume reproductive shares are either conceded, competed over, or both. Previous mathematical evaluations found that simultaneous concessions and contests are evolutionarily unstable. Recently, Shen and Reeve (2010) challenged these conclusions and developed a series of sub-models they argued to be a unified approach to reproductive skew: the general bordered tug-of-war (BTOW). However, BTOW fails as a general model for two reasons: (1) the BTOW strategy cannot invade populations where individuals either only compete for or only concede reproductive shares and (2) contrary to Shen and Reeve's assertion, BTOW populations are easily invaded by strategies with fewer or no concessions, but competing at lower levels. The failure of BTOW as a general model has major implications for interpreting experiments on reproductive skew. A large number of studies have measured the effects of genetic relatedness and competitive ability on reproductive skew, with a great majority finding no significant correlation between variation in within-group relatedness or competitive ability and across-group differences in skew. No model of reproductive skew except one variant of the BTOW predicts such results. With the rejection of BTOW as a valid general model, it is clear that these results are contradictory to reproductive skew theory rather than supportive of it.