Multiple pathways mediate the effects of climate change on maternal reproductive traits in a red deer population

Temporal changes in phenological traits arising as a consequence of recent rapid environmental change have been widely demonstrated in animal populations. Increasingly, studies are seeking to understand the impact of changes in such traits on individual fitness and population dynamics, with the ultimate aim of predicting population persistence or extinction under different climate scenarios. Here, we examined the effects of environmental change on maternal reproductive traits in a wild population of red deer (Cervus elaphus) and sought to explain why, despite a rapid advance in offspring birth dates, we observed no apparent consequences for offspring fitness. By using path analysis, we identified both direct and indirect paths along which changes in environmental conditions affected birth date, birth mass, juvenile survival, and female fecundity. In general, warmer temperatures were associated with earlier birth dates and greater birth mass, and higher rainfall was associated with reduced juvenile survival and reduced female fecundity. We also examined concurrent effects of population density, maternal age, and reproductive history, and found that temporal stasis in average trait values, at least in part, could be explained by antagonistic roles of direct and indirect effects of changing climate and increasing population density. Identification of the many mechanisms that contribute to the dynamics of phenotypic traits is challenging; this study demonstrates the need to consider both climatic and demographic variation in order to understand the fitness consequences of changes in phenological traits

Re-mating across years and intralineage polygyny are associated with greater than expected levels of inbreeding in wild red deer.

The interaction between philopatry and nonrandom mating has important consequences for the genetic structure of populations, influencing co-ancestry within social groups but also inbreeding. Here, using genetic paternity data, we describe mating patterns in a wild population of red deer (Cervus elaphus) which are associated with marked consequences for co-ancestry and inbreeding in the population. Around a fifth of females mate with a male with whom they have mated previously, and further, females frequently mate with a male with whom a female relative has also mated (intralineage polygyny). Both of these phenomena occur more than expected under random mating. Using simulations, we demonstrate that temporal and spatial factors, as well as skew in male breeding success, are important in promoting both re-mating behaviours and intralineage polygyny. However, the information modelled was not sufficient to explain the extent to which these behaviours occurred. We show that re-mating and intralineage polygyny are associated with increased pairwise relatedness in the population and a rise in average inbreeding coefficients. In particular, the latter resulted from a correlation between male relatedness and rutting location, with related males being more likely to rut in proximity to one another. These patterns, alongside their consequences for the genetic structure of the population, have rarely been documented in wild polygynous mammals, yet they have important implications for our understanding of genetic structure, inbreeding avoidance and dispersal in such systems

Consistent within-individual plasticity is sufficient to explain temperature responses in red deer reproductive traits

Warming global temperatures are affecting a range of aspects of wild populations, but the exact mechanisms driving associations between temperature and phenotypic traits may be difficult to identify. Here, we use a 36-year data-set on a wild population of red deer to investigate the causes of associations between temperature and two important components of female reproduction: timing of breeding and offspring size. By separating within- versus between-individual associations with temperature for each trait, we show that within-individual phenotypic plasticity (changes within a female’s lifetime) was entirely sufficient to generate the observed population-level association with temperature at key times of year. However, despite apparently adequate statistical power, we found no evidence of any variation between females in their responses (i.e. no ‘IxE’ interactions). Our results suggest that female deer show plasticity in reproductive traits in response to temperatures in the year leading up to calving, and that this response is consistent across individuals, implying no potential for either selection or heritability of plasticity. We estimate that the plastic response to rising temperatures explained 24% of the observed advance in mean calving date over the study period. We highlight the need for comparable analyses of other systems to determine the contribution of within-individual plasticity to population-level responses to climate change

Individual differences, density dependence and offspring birth traits in a population of red deer

Variation between individuals is an essential component of natural selection and evolutionary change, but it is only recently that the consequences of persistent differences between individuals on population dynamics have been considered. In particular, few authors have addressed whether interactions exist between individual quality and environmental variation. In part, this is due to the difficulties of collecting sufficient data, but also the challenge of defining individual quality. Using a long-established study population of red deer, Cervus elaphus, inhabiting the North Block of the Isle of Rum, and three quality measures, this paper investigates how differences in maternal quality affect variation in birth body mass and date, as population density varies, and how this differs depending on the sex of the offspring and the maternal quality measure used. Significant interactions between maternal quality, measured as a hind's total contribution to population growth, and population density are reported for birth mass, but only for male calves. Analyses using dominance or age at primiparity to define maternal quality showed no significant interactions with population density, highlighting the difficulties of defining a consistent measure of individual quality

The red deer rut revisited: female excursions but no evidence females move to mate with preferred males. Behav. Ecol

In polygynous mammals, most variance in male reproductive success is expected to result from competition between males, and the role of female behavior remains poorly understood. Contests between red deer males during the annual rut are one of the most famous examples of male-male competition. However, anecdotal evidence suggests females in estrus make substantial movements, changing harems, and potentially disrupting the outcome of male contests. In other polygynous mammals, such movements have been interpreted as evidence of female mating preferences. Here, we use 34 years of detailed observational data on wild red deer to show that 43% of estrous females are found in different harems between successive observations and that 64% of such females make substantial movements (up to 4 km) when this occurs. Approximately 45% of these movements result in the male into whose harem a female moved fathering her offspring. We then test whether females move nonrandomly with respect to male phenotype, consistent with the hypothesis that females move to mate with preferred males. Although in general, females were more likely to be found in larger harems and the harems of younger males after harem changes, these effects were not specific to estrous females. Further, estrous females were not more likely to be found in the harems of less related males. We therefore find little support for the idea that estrous females move between harems to mate with a preferred male; as a result, the reasons females make such extraordinary movements when in estrus remain unclear

Male Rut Data

Male rut data, from observations. Please see ReadMe file

Data from: Re-mating across years and intra-lineage polygyny are associated with greater than expected levels of inbreeding in wild red deer

The interaction between philopatry and non-random mating has important consequences for the genetic structure of populations, influencing co-ancestry within social groups but also inbreeding. Here, using genetic paternity data, we describe mating patterns in a wild population of red deer (Cervus elaphus) which are associated with marked consequences for co-ancestry and inbreeding in the population. Around a fifth of females mate with a male with whom they have mated previously, and further, females frequently mate with a male with whom a female relative has also mated (intra-lineage polygyny). Both of these phenomena occur more than expected under random mating. Using simulations, we demonstrate that temporal and spatial factors, as well as skew in male breeding success, are important in promoting both re-mating behaviours and intra-lineage polygyny. However, the information modelled was not sufficent to explain the extent to which these behaviours occured. We show that re-mating and intra-lineage polygyny are associated with increased pairwise relatedness in the population and a rise in average inbreeding coefficients. The latter in particular resulted from a non-random spatial distribution of the rutting male population with respect to relatedness, such that related males were more likely to rut in proximity to one another. These patterns, alongside their consequences for the genetic structure of the population, have rarely been documented in wild polygynous mammals, yet they have important implications for our understanding of genetic structure, inbreeding avoidance and dispersal in such systems

Pedigree

Pedigree, calculated as described in the paper

Sex differences in the consequences of maternal loss in a long-lived mammal, the red deer (Cervus elaphus)

In several primates, the presence of mothers affects the growth, survival and reproduction of their offspring, but similar effects have not yet been demonstrated in ungulates. Here, we investigate the effects of the mother's presence in a population of r