Indirect and direct models of sexual selection make different predictions regarding the quantitative genetic relationships between sexual ornaments and fitness. Indirect models predict that ornaments should have a high heritability and that strong positive genetic covariance should exist between fitness and the ornament. Direct models, on the other hand, make no such assumptions about the level of genetic variance in fitness and the ornament, and are therefore likely to be more important when environmental sources of variation are large. Here we test these predictions in a wild population of the blue tit (Parus caeruleus), a species in which plumage coloration has been shown to be under sexual selection. Using 3 years of cross-fostering data from over 250 breeding attempts, we partition the covariance between parental coloration and aspects of nestling fitness into a genetic and environmental component. Contrary to indirect models of sexual selection, but in agreement with direct models, we show that variation in coloration is only weakly heritable ðh 2 ! 0:11Þ, and that two components of offspring fitness-nestling size and fledgling recruitment-are strongly dependent on parental effects, rather than genetic effects. Furthermore, there was no evidence of significant positive genetic covariation between parental colour and offspring traits. Contrary to direct benefit models, however, we find little evidence that variation in colour reliably indicates the level of parental care provided by either males or females. Taken together, these results indicate that the assumptions of indirect models of sexual selection are not supported by the genetic basis of the traits reported on here.
In the plant–insect–insectivorous bird food chain, directional changes in climate can result in mismatched phenology, potentially affecting selection pressures. Phenotypic plasticity in the timing of breeding, characterized by reaction norm slopes, can help maximize fitness when faced with earlier prey emergence. In temperate passerines, the timing of tree budburst influences food availability for chicks through caterpillar phenology and the resulting food abundance patterns. Thus, the timing of tree budburst might serve as a more direct proxy for the cue to time egg‐laying. The evolutionary potential of breeding plasticity relies on heritable variation, which is based upon individual variation, yet studies on individual variation in plasticity are few. Here, we tested for the laying date—budburst date and the clutch size—laying date reaction norms, and examined 1) the among‐individual variance in reaction norm intercepts and slopes; and 2) the selection differentials and gradients on these intercepts and slopes. Using long‐term data of oak (genus Quercus) budburst and blue tit (Cyanistes caeruleus) reproduction, we applied within‐subject centering to detect reaction norms, followed by bivariate random regression to quantify among‐individual variance in reaction norm properties and their covariance with fitness. Individuals significantly differed in intercepts and slopes of both laying date—budburst date and clutch size—laying date reaction norms, and directional selection was present for an earlier laying date and a larger clutch size (intercepts), but not on plasticity (slopes). We found that individuals have their own regimes for adjusting egg‐laying and clutch size. This study provides further support of individual variation of phenotypic plasticity in birds.
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