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Hamel, Sandra; Gaillard, Jean-Michel; Douhard, Mathieu; Festa-Bianchet, Marco; Pelletier, Fanie; Yoccoz, Nigel G. 2017-09-21 Heterogeneity among individuals influences the life-history trajectories we observe at the population level because viability selection, selective immigration and emigration processes, and ontogeny change the proportion of individuals with specific trait values with increasing age. Here, we review the two main approaches that have been proposed to account for these processes in life-history trajectories, contrasting how they quantify ontogeny and selection, and proposing ways to overcome some of their limitations. Nearly all existing approaches to model individual heterogeneity assume either a single normal distribution or a priori known groups of individuals. Ontogenetic processes, however, can vary across individuals through variation in life-history tactics. We show the usefulness of describing ontogenetic processes by modelling trajectories with a mixture model that focuses on heterogeneity in life-history tactics. Additionally, most methods examine individual heterogeneity in a single trait, ignoring potential correlations among multiple traits caused by latent common sources of individual heterogeneity. We illustrate the value of using a joint modelling approach to assess the presence of a shared latent correlation and its influence on life-history trajectories. We contrast the strengths and limitations of different methods for different research questions, and we exemplify the differences among methods using empirical data from long-term studies of ungulates.
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Douhard, Mathieu; Festa-Bianchet, Marco; Coltman, Dave W.; Pelletier, Fanie; Coltman, David W. 2016-01-13 Parents should bias sex allocation toward offspring of the sex most likely to provide higher fitness returns. Trivers and Willard proposed that for polygynous mammals, females should adjust sex-ratio at conception or bias allocation of resources toward the most profitable sex, according to their own body condition. However, the possibility that mammalian fathers may influence sex allocation has seldom been considered. Here, we show that the probability of having a son increased from 0.31 to 0.60 with sire reproductive success in wild bighorn sheep (Ovis canadensis). Furthermore, our results suggest that females fertilized by relatively unsuccessful sires allocated more energy during lactation to daughters than to sons, while the opposite occurred for females fertilized by successful sires. The pattern of sex-biased offspring production appears adaptive because paternal reproductive success reduced the fitness of daughters and increased the average annual weaning success of sons, independently of maternal allocation to the offspring. Our results illustrate that sex allocation can be driven by paternal phenotype, with profound influences on the strength of sexual selection and on conflicts of interest between parents.
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Douhard, Mathieu; Pigeon, Gabriel; Festa-Bianchet, Marco; Coltmann, Dave W.; Guillemette, Simon; Pelletier, Fanie; Coltman, David W. 2016-11-30 The development of male secondary sexual characters such as antlers or horns has substantial biological and socio-economic importance because in many species these traits affect male fitness positively through sexual selection and negatively through trophy hunting. Both environmental conditions and selective hunting can affect horn growth but their relative importance remains unexplored. We first examined how a large-scale climate index, the Pacific Decadal Oscillation (PDO), local weather and population density influenced both absolute and relative annual horn growth from birth to 3 years of male bighorn sheep (Ovis canadensis) over 42 years. We then examined the relative influence of environmental conditions and evolution mainly driven by trophy hunting on male horn length at 3 years of age. Horn growth was positively influenced by low population density and warm spring temperature, suggesting that ongoing climate change should lead to larger horns. Seasonal values of PDO were highly correlated. Horn growth increased with PDO in spring or summer at low density, but was weak at high density regardless of PDO. The interaction between population density and PDO in spring or summer accounted for a similar proportion of the observed annual variation in horn growth (32% or 37%) as did the additive effects of spring temperature and density (34%). When environmental conditions deteriorated, males allocated relatively more resources to summer mass gain than to horn growth, suggesting a conservative strategy favoring maintenance of condition over allocation to secondary sexual characters. Population density explained 27% of the variation in horn length, while evolutionary effects explained 9% of the variance. Thus, our study underlines the importance of both evolution and phenotypic plasticity on the development of a secondary sexual trait.
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Douhard, Mathieu; Festa-Bianchet, Marco; Landes, Julie; Pelletier, Fanie 2019-03-07 1. Environmental conditions during early development, from conception to sexual maturity, can have lasting consequences on fitness components. Although adult lifespan often accounts for much of the variation in fitness in long-lived animals, we know little about how early environment affects adult lifespan in the wild, and even less about whether these effects differ between the sexes. 2. Using data collected over 45 years from wild bighorn sheep (Ovis canadensis), we investigated the effects of early environment on adult mortality in both sexes, distinguishing between natural and anthropogenic sources of mortality. 3. We used the average body mass of yearlings (at about 15 months of age) as a yearly index of environmental quality. We first examined sex differences in natural mortality responses to early environment by censoring harvested males in the year they were shot. We then investigated sex differences in the effects of early environment on overall mortality (natural and hunting mortality combined). Finally, we used path analysis to separate the direct influence of early environment from indirect influences, mediated by age at first reproduction, adult mass and horn length. 4. As early environmental conditions improved, natural adult mortality decreased in both sexes, although for males the effect was not statistically supported. Sex differences in the effects of early environment on adult mortality were detected only when natural and hunting mortality were pooled. Males that experienced favorable early environment had longer horns as adults and died earlier because of trophy hunting, which does not mimic natural mortality. Females that experienced favorable early environment started to reproduce earlier and early primiparity was associated with reduced mortality, suggesting a silver-spoon effect. 5. Our results show that early conditions affect males and females differently because of trophy hunting. These findings highlight the importance of considering natural and anthropogenic environmental factors across different life stages to understand sex differences in mortality.

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