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Adaptive explanations for dormancy often invoke bet hedging, where reduced mean fitness can be adaptive if it associates with reduced fitness variance. Sex allocation theory typically ignores variance effects and focuses on mean fitness. For many cyclical parthenogens, these themes become linked, as only sexually produced eggs undergo the dormancy needed to survive harsh conditions. We ask how sex allocation and the timing of sex evolve when this constraint exists in the form of a trade-off between asexual reproduction and sexual production of dormant eggs-the former being crucial for within-season success and the latter for survival across seasons. We show that male production can be temporally separated from or co-occur with sex, depending on whether direct (time) or indirect (population density) cues of the season's end are available and whether population growth is density dependent. Sex generally occurs late in the season but is induced earlier in unpredictable environments. When only indirect cues are available, the temporal spread of sex, and with it the production of dormant stages, is even larger and, given sufficient mortality, leads to endogenous population cycles in which frequent sex coincides with high densities. In all scenarios, algorithms maximizing geometric mean fitness have reduced fitness variance compared with a hypothetical non-bet hedger, confirming that the timing of male production and sex in facultative seasonal settings can be bet-hedging traits.

Coevolution with avian brood parasites shapes a range of traits in their hosts, including morphology, behavior, and breeding systems. Here we explore whether brood parasitism is also associated with the evolution of host clutch size. Several studies have proposed that hosts of highly virulent parasites could decrease the costs of parasitism by evolving a smaller clutch size, because hosts with smaller clutches will lose fewer progeny when their clutch is parasitized. We describe a model of the evolution of clutch size, which challenges this logic and shows instead that an increase in clutch size (or no change) should evolve in hosts. We test this prediction using a broad-scale comparative analysis to ask whether there are differences in clutch size within hosts and between hosts and nonhosts. Consistent with our model, this analysis revealed that host species do not have smaller clutches and that hosts that incur larger costs from raising a parasite lay larger clutches. We suggest that brood parasitism might be an influential factor in clutch-size evolution and could potentially select for the evolution of larger clutches in host species.

Successful invasions by sexually reproducing species depend on the ability of individuals to mate. Finding mates can be particularly challenging at low densities (a mate-finding Allee effect), a factor that is only implicitly accounted for by most invasion models, which typically assume asexual populations. Existing theory on single-sex populations suggests that dispersal evolution in the presence of a mate-finding Allee effect slows invasions. Here we develop a two-sex model to determine how mating system, strength of an Allee effect, and dispersal evolution influence invasion speed. We show that mating system differences can dramatically alter the spread rate. We also find a broader spectrum of outcomes than earlier work suggests. Allowing dispersal to evolve in a spreading context can sometimes alleviate the mate-finding Allee effect and slow the rate of spread. However, we demonstrate the opposite when resource competition among females remains high: evolution then acts to speed up the spread rate, despite simultaneously exacerbating the Allee effect. Our results highlight the importance of the timing of mating relative to dispersal and the strength of resource competition for consideration in future empirical studies.

Within and across taxa, there is much variation in the mode of fertilization, that is, whether eggs and/or sperm are released or kept inside or on the surface of the parent's body. Although the evolutionary consequences of fertilization mode are far-reaching, transitions in the fertilization mode itself have largely escaped the- oretical attention. Here we develop the first evolutionary model of egg retention and release, which also considers transitions between hermaphroditism and dioecy as well as egg size evolution. We provide a unifying explanation for reported associations between small body size, hermaphroditism, and egg retention in marine invertebrates that have puzzled researchers for more than 3 decades. Our model, by including sperm limitation, shows that all these patterns can arise as an evolutionary response to local competition between eggs for fertilization. This can provide a general explanation for three em- pirical patterns: sperm casters tend to be smaller than related broad- cast spawners, hermaphroditism is disproportionately common in sperm casters, and offspring of sperm casters are larger. Local gamete competition also explains a universal sexual asymmetry: females of some species retain their gametes while males release theirs, but the opposite ("egg casting") lacks evolutionary stability and is apparently not found in nature.

Mate choice frequently operates differently for males and females as a consequence of male competition for mates. Competitive interactions can alter the fitness payoffs of choice and the realization of preferences under natural conditions, yet the majority of male choice studies still use binary trials that ignore social factors. Here we test the importance of contest dynamics in male choice using a framework in which females are considered analogous to foraging patches that are subject to competition. We track the mate choices and interactions of 640 spiders (Nephila plumipes) before and after manipulation of competition within enclosures, modeling the expected fitness payoffs of each male's actual choices and comparing these with all alternative choices. Many males choose new mates once social conditions change and achieve higher fitness than predicted under random movement. Males do not simply move to larger females but choose favorable competitive environments that balance competition and female fecundity, thereby increasing their fitness payoffs. Further, we show for the first time that prior-residence effects, which are known to influence male contests, also have a strong influence in male reproductive contests and can shape male mate choice. These results highlight the importance of situating male choice studies in the relevant social context, as intrasexual interactions can have profound effects on the realization and payoffs of male mate-choice strategies.

Explaining the "prior-residence effect" (automatic owner status of individuals who arrived first in an area) was one of the very first applications of game theory in animal behavior. These models, however, predict paradoxical solutions where intruders always win, with no satisfactory explanation for the absence of such cases in nature. We propose a solution based on new developments in evolutionary game theory. A self-consistent model with feedbacks between individual behavior and population dynamics produces qualitatively different frequency-dependent selection on intruders (floaters) than on territory owners. Starting with an ancestral population with no respect for ownership, the most likely evolutionary end point is complete or partial respect. Conventional rules of conflict resolution thus can rely on "uncorrelated asymmetries" without differences in resource-holding power or territory value, although they will be strengthened by such differences. We also review the empirical literature on animal contests, testing whether asymmetries in resource-holding power are required to explain the observations. Despite much empirical effort, results remain inconclusive, because experiments are often unable to distinguish between the motivation of individuals to fight and the behavioral outcome of a contest. To help arrive at conclusive answers, we suggest a standardized empirical approach to quantify prior-residence effects.

When should males begin guarding a resource when both resources and guarders vary in quality? This general problem applies, for example, to migrant birds occupying territories in the spring and to precopula in crustaceans where males grab females before they molt and become receptive. Previous work has produced conflicting predictions. Theory on migrant birds predicts that the strongest competitors should often arrive first, whereas some models of mate guarding have predicted that the strongest competitors wait and then simply usurp a female from a weaker competitor. We build a general model of resource guarding that allows varying the ease with which takeovers occur. The model is phrased in terms of mate-guarding crustaceans, but the same logic can be applied to other forms of resource acquisition where priority plays a role but takeovers might be possible too. The race to secure breeding positions can lead to strong competitors (large males) taking females earliest, even though this means accepting a lower-quality female. Paradoxically, this means that small males, which have fewer breeding opportunities, are more choosy than larger ones. Such solutions are found when takeovers are impossible. The easier the takeovers and the higher the rate of finding guarded resources, the more likely are solutions where guarding durations are short, where strong competitors begin guarding only just before breeding, and where they do this by usurping the resource. The relationship between an individual's competitive ability and its timing of resource acquisition can also be nonlinear if takeovers are moderately common; if this is the case, then males of intermediate size guard the longest.

Life-history theory concerns the optimal spread of reproduction over an organism's life span. In variable environments, there may be extrinsic differences between breeding periods within an organism's life, affecting both offspring and parent and giving rise to intergenerational trade-offs. Such trade-offs are often discussed in terms of reproductive value for parent and offspring. Here, we consider parental life-history optimization in response to varying offspring values of a population regulated by territoriality, where the quality of the environment varies periodically. Periods are interpreted as either within-year (seasonality) or between-years variation (cyclicity). The evolutionarily stable strategy in a general model with two-phased periodicity in the environment can generate either higher or lower effort in the more favorable of the two phases; hence knowing survival prospects of offspring does not suffice for predicting reproductive effort-the future of all descendants and the parent must be tracked. We also apply our method to data on the Ural owl Strix uralensis, a species preying on cyclically fluctuating voles. The observed dynamics are best predicted by assuming delayed reproductive costs and Type II functional response. Accounting for varying offspring values can lead to cases where both reproductive effort and recruitment of offspring are higher in the phase when voles are not maximally abundant, a pattern supported by our data.

Charles Darwin (1871) defined sexual selection as a difference inreproductive success due to competition over mates, a definition that is still valid today. Being able to quantify such differences is thus essential for explaining the consequences of sexual selection, say, the ornamentation rdisplay behavior of males. In the specific ase of lekking (aggregated, non-resource-based display of males), attempts to do so have proven difficult. Any measure that does not take into account the possibility of random mating to produce the observed pattern of mating success is unsatisfactory as a measure of the intensity of sexual selection (Sutherland 1987; Mackenzie et al. 1995b). That variation in male mating success on leks is systematically higher than expected from pure random mating has been verified by Bradbury et al. (1985) and Mackenzie et al. (1995a). However, these studies do not yet provide means to estimate the expected mating success of individuals displaying together. Since a lek is not itself a trait but an assemblage of several males, each trying to optimize their own fitness (Hoglund and Alatalo 1995), fitness considerations of single individuals are required in explaining the evolution of lekking. The skew index, defined as the ratio of observed to maximum skew (Keller and Vargo 1993; Reeve and Ratnieks 1993), has been gaining popularity as a measure of skewed reproductive success (e.g., Bourke and Heinze 1994; Keller and Reeve 1994; Reeve and Keller 1995), and it has also been used to estimate the expected fitness of lekking individuals (Widemo and Owens 1995). However, this measure has been shown to have undesirable discontinuities (Pamilo and Crozier 1996). Moreover, it interprets random variation as real differences in mating success. This is most clearly seen in models of pure random mating, which nevertheless lead to positive values of the skew measure (Mackenzie et al. 1995b). Recently, Keller and Krieger (1996) proposed a method for correcting the skew index for random variation. We shall show, however, that the corrected measure still is not satisfactory, and we suggest a novel, statistically rigorous method. We give two criteria for a proper measure of skewed mating success. First, it should allow determining the most probable underlying distribution of mating

Sexual selection is often quantified using Bateman gradients, which represent sex-specific regression slopes of reproductive success on mating success and thus describe the expected fitness returns from mating more often. Although the analytical framework for Bateman gradients aimed at covering all sexual systems, empirical studies are biased toward separate-sex organisms, probably because important characteristics of other systems remain incompletely treated. Our synthesis complements the existing Bateman gradient approach with three essential reproductive features of simultaneous hermaphrodites. First, mating in one sex may affect fitness via the opposite sex, for example, through energetic trade-offs. We integrate cross-sex selection effects and show how they help characterizing sexually mutualistic versus antagonistic selection. Second, male and female mating successes may be correlated, complicating the interpretation of Bateman gradients. We show how to quantify the impact of this correlation on sexual selection and propose a principal component analysis on male and female mating success to facilitate interpretation. Third, self-fertilization is accounted for by adding selfed progeny as a separate category of reproductive success to analyses of Bateman gradients. Finally, using a worked example from the snail Biomphalaria glabrata, we illustrate how the extended analytical framework can enhance our understanding of sexual selection in hermaphroditic animals and plants. © 2010 by The University of Chicago.

When several individuals simultaneously provide for offspring, as in families, the effort of any one individual will depend on the efforts of the other family members. This conflict of interest among family members is made more complicated by their relatedness because relatives share genetic interest to some degree. The conflict resolution will also be influenced by the differences in reproductive value between breeders and helpers. Here, we calculate evolutionarily stable provisioning efforts in families with up to two helpers. We explicitly consider that the behavioral choices are made in a life-history context, and we also consider how group sizes change dynamically; this affects, for example, average relatedness among group members. We assume two different scenarios: intact families in which the breeder is 100% monogamous and stepfamilies in which the breeder shifts mate between breeding events. The average relatedness among family members is allowed to evolve in concert with changes in provisioning effort. Our model shows that an individual's provisioning effort is not easy to predict from either its relatedness to the offspring or its reproductive value. Instead, it is necessary to consider the inclusive fitness effect of provisioning, which is determined by a combination of relatedness, reproductive value, and the reproductive value of the offspring.

We examine the evolutionary stability of year-round residency in territorial populations, where breeding sites are a limiting resource. The model links individual life histories to the population-wide competition for territories and includes spatial variation in habitat quality as well as a potential parent-offspring conflict over territory ownership. The general form of the model makes it applicable to the evolution of dispersal, migration, partial migration, and delayed dispersal (offspring retention). We show that migration can be evolutionarily stable only if year-round residency in a given area would produce a sink population, where mortality exceeds reproduction. If this applies to a fraction of the breeding habitat only, partial migration is expected to evolve. In the context of delayed dispersal, habitat saturation has been argued to form an ecological constraint on independent breeding, which favors offspring retention and cooperative breeding. We show that habitat saturation must be considered as a dynamic outcome of birth, death, and dispersal rates in the population, rather than an externally determined constraint. Although delayed dispersal often associates with intense competition for territories, life-history traits have direct effects on stable dispersal strategies, which can often override the effect of habitat saturation. As an example, high survival of floaters selects against delayed dispersal, even though it increases the number of competitors for each breeding vacancy (the "habitat saturation factor"). High survival of territory owners, by contrast, generally favors natal philopatry. We also conclude that spatial variation in habitat quality only rarely selects for delayed dispersal. Within a population, however, offspring retention is more likely in high-quality territories.