Protandry, the earlier arrival of males to breeding areas than females, is a common pattern of sex-biased timing in many animal taxa (e.g. some insects, ®sh, amphibians, reptiles, birds and mammals). The adaptive signi®cance of protandry is not fully understood and, since the 1970s, at least seven hypotheses for protandry have been proposed. We describe each of these hypotheses and summarize what is known about each. In three of these hypotheses, the relative arrival timing of males and females has no direct ®tness consequences for males or females, but selection for different timing in each sex indirectly produces protandry. In the other four hypotheses, the difference between male and female timing has ®tness consequences for males or females and selection directly maintains the ®tness-maximizing degree of sex-biased timing. The hypotheses are not mutually exclusive, and the degree of multiple mating by males and the occurrence of male territoriality seem to determine the relative importance of each hypothesis. In order to understand the adaptive signi®cance of sex-biased timing, future studies need to consider all the alternatives and to assess the costs and bene®ts to males of early arrival relative to calendar date, to other males and to females.

The antagonistic pleiotropy theory of senescence postulates genes or traits that have opposite effects on early-life and late-life performances. Because selection is generally weaker late in life, genes or traits that improve early-life performance but impair late-life performance should come to predominate. Variation in the strength of age-specific selection should then generate adaptive variation in senescence. We demonstrate this mechanism by comparing early and late breeders within a population of semelparous capital-breeding sockeye salmon (Oncorhynchus nerka). We show that early breeders (but not late breeders) are under strong selection for a long reproductive lifespan (RLS), which facilitates defence of their nests against disturbance by later females. Accordingly, early females invest less energy in egg production while reserving more for nest defence. Variation along this reproductive trade-off causes delayed or slower senescence in early females (average RLS of 26 days) than in late females (reproductive lifespan of 12 days). We use microsatellites to confirm that gene flow is sufficiently limited between early and late breeders to allow adaptive divergence in response to selection. Because reproductive trade-offs should be almost universal and selection acting on them should typically vary in time and space, the mechanism described herein may explain much of the natural variation in senescence.

In many migratory birds, males precede females during migration and arrival at the breeding sites. Three proximate mechanisms are proposed to explain this phenomenon of protandry: males 1) winter closer to breeding sites, 2) start spring migration earlier, and/or 3) migrate faster than females. So far, the relative contribution of these mechanisms to protandry is unknown. The present study investigated the importance of each of the 3 proximate mechanisms of protandry for a songbird migrant wintering in Africa, the northern wheatear (Oenanthe oenanthe). Two subspecies co-occur in Europe on migration, of which the leucorhoa northern wheatears breeding from Iceland to Canada have to cross the North Atlantic, whereas the nominate form breeding in Europe does not face any significant sea barrier. We show that the leucorhoa subspecies had a significantly higher degree of protandry at stopover sites across Europe than the oenanthe subspecies (−6 vs. −2 days). Leucorhoa northern wheatear's higher degree of protandry was associated with a larger age effect, in which old males preceded young males, and greater sex-specific differences in wing shape and refueling yielding higher migration speeds in males than females. In oenanthe northern wheatears, light-level geolocators revealed that males did not winter closer to the breeding sites or migrate faster than females, but initiated spring migration earlier. Our results demonstrate that the significance of the mechanisms causing protandry can differ between related taxa and highlight the importance of the advancement in male arrival date with age as a potential factor shaping the degree of protandry.

We use a game-theoretic framework to investigate the reproductive phenology of female kokanee (Oncorhynchus nerka). As in the other semelparous species of Pacific salmon, females construct nests in gravel, spawn with males, bury their fertilized eggs, and defend their nest sites until they die several days later. Later-breeding females may reuse previous nest sites, and their digging behavior is thought to subject previously buried eggs to mortality. Using gametheoretic models, we show that females can reduce this risk by allocating resources to longevity (the period between arrival and death) as opposed to eggs. Waiting before territory settlement is also expected if it allows females to conserve energy and delay senescence. The models demonstrate how these costs and benefits interact to select for a seasonal decline in longevity, a well-known phenomenon in the salmonid literature, and a seasonal decline in wait duration. Both of these predictions were supported in a field study of kokanee. Female state of reproductive maturity was the most important proximate factor causing variation in longevity and wait duration. With more than 30% of territories being reused, dig-up is likely an important selective force in this population.

To learn more about the degree of individual variation in resource use by lake trout Salvelinus namaycush, ultrasonic telemetry was used to study their habitat use in a lake without pelagic schooling fish prey. Individuals spent most of their time within the metalimnion in favourable water temperatures. They also made frequent excursions, however, into lake temperatures exceeding their optimum for physiological performance at all temporal scales considered. Their frequent use of nearshore habitats suggested that feeding in littoral areas may be common. Habitat use was highly variable among individuals, but spatial habitat use by individuals showed remarkable consistency between years. In particular, some lake trout exhibited high site fidelity to shallow, nearshore areas, whereas others used deep areas extensively. This level of between-individual variation indicated niche partitioning by depth and the possibility of alternative foraging strategies.