Online enhancements: supplementary tables and figure. Dryad data: http://dx.doi.org/10.5061/dryad.rg8d4.abstract: Age is a major factor explaining variation in life-history traits among individuals with typical patterns of increasing trait values early in life, maximum trait expression, and senescence. However, age-dependent variation in the expressions of sexually selected traits has received less attention, although such variation underpins differences in male competitive abilities and female preference, which are central to sexual selection. In contrast to previous studies focusing on single traits, we used repeated measures of seven sexually selected morphological and behavioral traits in male black grouse (Tetrao tetrix) to quantify the effects of age and life span on their expressions and quantified this variation in relation to male reproductive effort. Trait expression increased with age, but long-lived males had a slower increase and delayed maxima in trait values compared with shortlived males. There was evidence of terminal investment (increasing trait values during the last breeding season) in some traits and senescence in all traits. These trait dynamics were largely explained by the timing of male peak lekking effort. This study shows that fully understanding the variation in sexually selected traits and fitness benefits associated with sexual selection requires accounting for the complex interaction among individual age, life span, and the timing of individuals' investment in reproduction.

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Summary1. Individuals' reproductive success is often strongly associated with their age, with typical patterns of early-life reproductive improvement and late-life senescence. These age-related patterns are due to the inherent trade-offs between life-history traits competing for a limited amount of resources available to the organisms. In males, such trade-offs are exacerbated by the resource requirements associated with the expression of costly sexual traits, leading to dynamic changes in trait expression throughout their life span. 2. Due to the age dependency of male phenotypes, the relationship between the expression of male traits and mating success can also vary with male age. Hence, using longitudinal data in a lekking species with strong sexual selection -the black grouse Lyrurus tetrix -we quantified the effects of age, life span and age of first lek attendance (AFL) on male annual mating success (AMS) to separate the effects of within-individual improvement and senescence on AMS from selective (dis)appearance of certain phenotypes. Then, we used male AMS to quantify univariate and multivariate sexual selection gradients on male morphological and behavioural traits with and without accounting for age and age-related effects of other traits. 3. Male AMS increased with age, and there was no significant reproductive senescence. Most males never copulated, and of the ones that did, the majority had only one successful year. Life span was unrelated to AMS, but early AFL tended to lead to higher AMS at ages 1-3. AMS was related to most morphological and behavioural traits when male age was ignored. Accounting for age and age-specific trait effects (i.e. the interaction between a trait and age) reduced the magnitude of the selection gradients and revealed that behavioural traits are under consistent sexual selection, while sexual selection on morphological traits is stronger in old males. 4. Therefore, sexual selection in black grouse operates primarily on male behaviour and morphological traits may act as additional cues to supplement female choice. These results demonstrate the multifaceted influence of age on both fitness and sexual traits and highlight the importance of accounting for such effects when quantifying sexual selection.

Carotenoid-based traits commonly act as condition-dependent signals of quality to both males and females. Such colors are typically quantified using summary metrics (e.g., redness) derived by partitioning measured reflectance spectra into blocks. However, perceived coloration is a product of the whole spectrum. Recently, new methods have quantified a range of environmental factors and their impact on reflection data at narrow wavebands across the whole spectrum. Using this approach, we modeled the reflectance of red integumentary eye combs displayed by male black grouse (Lyrurus tetrix) as a function of ornament size and variables related to male quality. We investigated the strength and direction of effect sizes of variables at each waveband. The strongest effect on the spectra came from eye comb size, with a negative effect in the red part of the spectrum and a positive effect in ultraviolet reflectance. Plasma carotenoid concentration and body mass were also related to reflectance variance in differing directions across the entire spectra. Comparisons of yearlings and adults showed that the effects were similar but stronger on adult reflectance spectra. These findings suggest that reflectance in different parts of the spectrum is indicative of differing components of quality. This method also allows a more accurate understanding of how biologically relevant variables may interact to produce perceived coloration and multicomponent signals and where the strongest biological effects are found.

Lyrurus tetrix lek behaviours, we tested whether there were fine-scale differences in 21 reproductive effort (lek attendance, fighting rates) and whether these were related to age

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Male ornaments function as honest cues of male quality in many species and are subject to intra‐ and intersexual selection. These ornaments are generally studied during peak expression, however their size outside the breeding season may determine ultimate ornament size and costliness, and as such reproductive success. We investigated whether male black grouse Lyrurus tetrix eye comb size was related to age, condition and measures of male dominance before and during the breeding season. Total combined eye comb size began to increase ~70 d before the start of the breeding season. Adult males (aged ≥ 2 yr old) had consistently larger eye combs than younger males (1 yr old) both before and during the breeding season. Heavier and more dominant adult males (attending the lek more frequently and successfully reproducing) had larger eye combs. For younger males, those that were heavier had larger eye combs. Additionally, males that spent more time on the lek showed increased eye comb size as the breeding season approached. Overall we find that ornament size is positively related to dominance and condition before and during the breeding season. Since dominance is accrued through year‐round interactions in many species, the ability to maintain larger signals over prolonged periods, including outside of the breeding season, is likely to be beneficial for adults. For younger males, it is likely that they cannot sustain or are constrained from producing larger eye combs over long periods of time. They therefore prioritise growth of their ornaments later, and according to the amount of time they spend on the lek.