Sexual Selection in Dung Beetles .2. Female Fecundity as an Estimate of Male Reproductive Success in Relation to Horn Size, and Alternative Behavioral Strategies in Onthophagus-Binodis Thunberg (Scarabaeidae, Onthophagini)

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Abstract
One component of the reproductive success (fitness) of Onthophagus binodis Thunberg males was estimated by the number of offspring their mate produced relative to male horn and body size. O. binodis males consist of large horned and small hornless morphs. Female fecundity was significantly increased when reproducing with: (1) large horned males compared with small hornless males; (2) the horned morph compared with the hornless morph of males with similar body size. Horned males cooperating with females invest a considerable effort in providing each egg with dung. Hornless males do not appear to assist females after mating. Alternative male mating strategies are predicted under intense intrasexual competition. As hornless O. binodis males persist in large numbers alongside cooperative, mate guarding horned males, selection should favour alternative tactics by hornless males to gain reproductive opportunities.