Reproductive allocation in pulsed-resource environments: a comparative study in two populations of wild boar.

Pulsed resources influence the demography and evolution of consumer populations and, by cascading effect, the dynamics of the entire community. Mast seeding provides a case study for exploring the evolution of life history traits of consumers in fluctuating environments. Wild boar (Sus scrofa) population dynamics is related to seed availability (acorns/beechnuts). From a long-term monitoring of two populations subjected to markedly different environmental contexts (i.e., both low vs. high frequency of pulsed resources and low vs. high hunting pressure in Italy and in France, respectively), we assessed how pulsed resources shape the reproductive output of females. Using path analyses, we showed that in both populations, abundant seed availability increases body mass and both the absolute and the relative (to body mass) allocation to reproduction through higher fertility. In the Italian population, females equally relied on past and current resources for reproduction and ranked at an intermediate position along the capital-income continuum of breeding tactics. In contrast, in the French population, females relied on current more than past resources and ranked closer to the income end of the continuum. In the French population, one-year old females born in acorn-mast years were heavier and had larger litter size than females born in beechnut-mast years. In addition to the quantity, the type of resources (acorns/beechnuts) has to be accounted for to assess reliably how females allocate resources to reproduction. Our findings highlight a high plasticity in breeding tactics in wild boar females and provide new insight on allocation strategies in fluctuating environments.

The relationship between phenotypic variation among offspring and mother body mass in wild boar: evidence of coin-flipping?

1. In highly variable environments, the optimal reproductive tactics of iteroparous organisms should minimize variance in yearly reproductive success to maximize the long-term average reproductive success. To minimize among-year variation in reproductive success, individuals can either minimize the variance in the number of offspring produced at each reproductive attempt (classical bet-hedging) or maximize the phenotypic diversity of offspring produced within or among reproductive attempts (coin-flipping). 2. From a long-term detailed study of an intensively exploited population facing a highly unpredictable environment, we identify a continuum of reproductive tactics in wild boar females depending on their body mass. 3. At one end, light females adjusted litter size to their body mass and produced highly similar-sized offspring within a litter. These females fitted the hypothesis of individual optimization commonly reported in warm-blooded species, which involves both an optimal mass and an optimal number of offspring for a given mother. At the other end of the continuum, heavy females produced litters of variable size including a mixture of heavy and light offspring within litters. 4. Prolific heavy wild boar females diversify the phenotype of their offspring, providing a first evidence for coin-flipping in a warm-blooded species.

Fluctuating food resources influence developmental plasticity in wild boar.

To maximize long-term average reproductive success, individuals can diversify the phenotypes of offspring produced within a reproductive event by displaying the 'coin-flipping' tactic. Wild boar (Sus scrofa scrofa) females have been reported to adopt this tactic. However, whether the magnitude of developmental plasticity within a litter depends on stochasticity in food resources has not been yet investigated. From long-term monitoring, we found that juvenile females produced similar-sized fetuses within a litter independent of food availability. By contrast, adult females adjusted their relative allocation to littermates to the amount of food resources, by providing a similar allocation to all littermates in years of poor food resources but producing highly diversified offspring phenotypes within a litter in years of abundant food resources. By minimizing sibling rivalry, such a plastic reproductive tactic allows adult wild boar females to maximize the number of littermates for a given breeding event.

Assessing whether mortality is additive using marked animals: a Bayesian state-space modeling approach.

Whether different sources of mortality are additive, compensatory, or depensatory is a key question in population biology. A way to test for additivity is to calculate the correlation between cause-specific mortality rates obtained from marked animals. However, existing methods to estimate this correlation raise several methodological issues. One difficulty is the existence of an intrinsic bias in the correlation parameter. Although this bias can be formally expressed, it requires knowledge about natural survival without any competing mortality source, which is difficult to assess in most cases. Another difficulty lies in estimating the true process correlation while properly accounting for sampling variation. Using a Bayesian approach, we developed a state-space model to assess the correlation between two competing sources of mortality. By distinguishing the mortality process from its observation through dead recoveries and live recaptures, we estimated the process correlation. To correct for the intrinsic bias, we incorporated experts' opinions on natural survival. We illustrated our approach using data on a hunted population of wild boars. Mortalities were not additive and natural mortality increased with hunting mortality likely as a consequence of non-controlled mortality by crippling loss. Our method opens perspectives for wildlife management and for the conservation of endangered species.

On the evolutionary consequences of increasing litter size with multiple paternity in wild boar (Sus scrofa scrofa).

Understanding how some species may be able to evolve quickly enough to deal with anthropogenic pressure is of prime interest in evolutionary biology, conservation, and management. Wild boar (Sus scrofa scrofa) populations keep growing all over Europe despite increasing hunting pressure. In wild boar populations subject to male-selective harvesting, the initially described polygynous mating system may switch to a promiscuous/polyandrous one. Such a change in the mating system, where potentially more males sire a litter at one reproductive event, may be associated with the retention of high genetic diversity and an increase of litter size. We tested these hypotheses by estimating the number of sires per litter based on a six-year long monitoring of a wild boar population subject to particularly high harvesting pressure. Our results show a high and stable genetic diversity and high rates of multiple paternity compared to other populations, thus depicting a promiscuous/polyandrous mating system in this population. We also show that litter size is positively linked to the number of sires, suggesting that multiple paternity increases fecundity. We finally discuss that multiple paternity may be one of the factors allowing rapid evolution of this population by maintaining both genetic and phenotypic diversity.

High hunting pressure selects for earlier birth date: wild boar as a case study.

Exploitation by humans affects the size and structure of populations. This has evolutionary and demographic consequences that have typically being studied independent of one another. We here applied a framework recently developed applying quantitative tools from population ecology and selection gradient analysis to quantify the selection on a quantitative trait-birth date-through its association with multiple fitness components. From the long-term monitoring (22 years) of a wild boar (Sus scrofa scrofa) population subject to markedly increasing hunting pressure, we found that birth dates have advanced by up to 12 days throughout the study period. During the period of low hunting pressure, there was no detectable selection. However, during the period of high hunting pressure, the selection gradient linking breeding probability in the first year of life to birth date was negative, supporting current life-history theory predicting selection for early births to reproduce within the first year of life with increasing adult mortality.