Evolutionary biologists have developed several indices, such as selection gradients (β) and the opportunity for sexual selection (Is), to quantify the actual and/or potential strength of sexual selection acting in natural or experimental populations. In a recent paper, Klug et al. (J. Evol. Biol.23, 2010, 447) contend that selection gradients are the only legitimate metric for quantifying sexual selection. They argue that Is and similar mating‐system‐based metrics provide unpredictable results, which may be uncorrelated with selection acting on a trait, and should therefore be abandoned. We find this view short‐sighted and argue that the choice of metric should be governed by the research question at hand. We describe insights that measures such as the opportunity for selection can provide and also argue that Klug et al. have overstated the problems with this approach while glossing over similar issues with the interpretation of selection gradients. While no metric perfectly characterizes sexual selection in all circumstances, thoughtful application of existing measures has been and continues to be informative in evolutionary studies.

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Leks are classic models for studies of sexual selection due to extreme variance in male reproductive success, but the relative influence of intrasexual competition and female mate choice in creating this skew is debatable. In the lekking lance-tailed manakin (Chiroxiphia lanceolata), these selective episodes are temporally separated into intrasexual competition for alpha status and female mate choice among alpha males that rarely interact. Variance in reproductive success between status classes of adult males (alpha versus non-alpha) can therefore be attributed to male-male competition whereas that within status largely reflects female mate choice. This provides an excellent opportunity for quantifying the relative contribution of each of these mechanisms of sexual selection to the overall opportunity for sexual selection on males (I males ). To calculate variance in actual reproductive success, we assigned genetic paternity to 92.3% of 447 chicks sampled in seven years. Reproduction by non-alphas was rare and apparently reflected status misclassifications or opportunistic copulations en route to attaining alpha status rather than alternative mating strategies. On average 31% (range 7-44%, nZ6 years) of the total I males was due to variance in reproductive success between alphas and non-alphas. Similarly, in a cohort of same-aged males followed for six years, 44 -58% of the total I males was attributed to variance between males of different status. Thus, both intrasexual competition for status and female mate choice among lekking alpha males contribute substantially to the potential for sexual selection in this species.

Male lance-tailed manakins (Chiroxiphia lanceolata) cooperate in complex courtship displays, but the dominant (alpha) partner monopolizes mating opportunities. This raises the question of why subordinates (betas) cooperate. Three nonexclusive hypotheses explain the adaptive basis of helping behavior by subordinate males: cooperation may increase (1) subordinates' immediate reproductive success, (2) the reproductive success of close relatives, or (3) subordinates' chances of future reproduction. I demonstrated that beta males rarely sired chicks and were unrelated to their alpha partners but received delayed direct benefits from cooperation; betas had an increased probability of becoming an alpha when compared to males that had not been betas. To investigate the mechanism by which betas attain these adaptive benefits, I examined betas' success in replacing their alpha partners both in natural turnover events and when alphas were experimentally removed. Beta males did not consistently inherit alpha roles in the same territories where they served their beta tenure, arguing that queuing for status does not fully explain the benefits of cooperation for betas. Instead, betas may be apprenticing to develop effective and appropriate displays that enhance their subsequent success as alphas. Complex social affiliations appear to mediate selective pressure for cooperation in this species.Keywords: lance-tailed man akin, Chiroxiphia, cooperative courtship, relatedness, delayed direct benefits.Cooperative reproduction is a classic paradox in evolutionary biology because cooperating individuals apparently sacrifice their own reproductive potential to assist the reproductive efforts of others. Much theory about cooperation is informed by studies of cooperative breeding, in * Present address: