Inbreeding depression in red deer calves

BACKGROUND Understanding the fitness consequences of inbreeding is of major importance for evolutionary and conservation biology. However, there are few studies using pedigree-based estimates of inbreeding or investigating the influence of environment and age variation on inbreeding depression in natural populations. Here we investigated the consequences of variation in inbreeding coefficient for three juvenile traits, birth date, birth weight and first year survival, in a wild population of red deer, considering both calf and mother's inbreeding coefficient. We also tested whether inbreeding depression varied with environmental conditions and maternal age. RESULTS We detected non-zero inbreeding coefficients for 22% of individuals with both parents and at least one grandparent known (increasing to 42% if the dataset was restricted to those with four known grandparents). Inbreeding depression was evident for birth weight and first year survival but not for birth date: the first year survival of offspring with an inbreeding coefficient of 0.25 was reduced by 77% compared to offspring with an inbreeding coefficient of zero. However, it was independent of measures of environmental variation and maternal age. The effect of inbreeding on birth weight appeared to be driven by highly inbred individuals (F = 0.25). On the other hand first year survival showed strong inbreeding depression that was not solely driven by individuals with the highest inbreeding coefficients, corresponding to an estimate of 4.35 lethal equivalents. CONCLUSIONS These results represent a rare demonstration of inbreeding depression using pedigree-based estimates in a wild mammal population and highlight the potential strength of effects on key components of fitness.This research was supported by a NERC grant to LEBK, JMP and THCB, NERC and BBSRC fellowships to DHN and a Royal Society fellowship to LEBK

Numerical assessment in the wild: insights from social carnivores

Playback experiments have proven to be a useful tool to investigate the extent to which wild animals understand numerical concepts and the factors that play into their decisions to respond to different numbers of vocalizing conspecifics. In particular, playback experiments have broadened our understanding of the cognitive abilities of historically understudied species that are challenging to test in the traditional laboratory, such as members of the Order Carnivora. Additionally, playback experiments allow us to assess the importance of numerical information versus other ecologically important variables when animals are making adaptive decisions in their natural habitats. Here, we begin by reviewing what we know about quantity discrimination in carnivores from studies conducted in captivity. We then review a series of playback experiments conducted with wild social carnivores, including African lions, spotted hyenas, and wolves, which demonstrate that these animals can assess the number of conspecifics calling and respond based on numerical advantage. We discuss how the wild studies compliment those conducted in captivity and allow us to gain insights into why wild animals may not always respond based solely on differences in quantity. We then consider the key role that individual discrimination and cross-modal recognition play in the ability of animals to assess the number of conspecifics vocalizing nearby. Finally, we explore new directions for future research in this area, highlighting in particular the need for further work on the cognitive basis of numerical assessment skills and experimental paradigms that can be effective in both captive and wild settings

Endocranial volume is heritable and is associated with longevity and fitness in a wild mammal

Research on relative brain size in mammals suggests that increases in brain size may generate benefits to survival and costs to fecundity: comparative studies of mammals have shown that interspecific differences in relative brain size are positively correlated with longevity and negatively with fecundity. However, as yet, no studies of mammals have investigated whether similar relationships exist within species, nor whether individual differences in brain size within a wild population are heritable. Here we show that, in a wild population of red deer ($\textit{Cervus elaphus}$), relative endocranial volume was heritable (h² = 63%; 95% credible intervals (CI) = 50-76%). In females, it was positively correlated with longevity and lifetime reproductive success, though there was no evidence that it was associated with fecundity. In males, endocranial volume was not related to longevity, lifetime breeding success or fecundity.Leverhulme Trust; Isaac Newton Trust; Natural Environmental Research Council (NE/L00688X/1); European Research Council (grant nos. 250098 and 294494); Australian Research Counci

Relative costs of offspring sex and offspring survival in a polygynous mammal.

Costs of reproduction are expected to be ubiquitous in wild animal populations and understanding the drivers of variation in these costs is an important aspect of life-history evolution theory. We use a 43 year dataset from a wild population of red deer to examine the relative importance of two factors that influence the costs of reproduction to mothers, and to test whether these costs vary with changing ecological conditions. Like previous studies, our analyses indicate fitness costs of lactation: mothers whose calves survived the summer subsequently showed lower survival and fecundity than those whose calves died soon after birth, accounting for 5% and 14% of the variation in mothers' survival and fecundity, respectively. The production of a male calf depressed maternal survival and fecundity more than production of a female, but accounted for less than 1% of the variation in either fitness component. There was no evidence for any change in the effect of calf survival or sex with increasing population density

Environmental coupling of selection and heritability limits evolution

There has recently been great interest in applying theoretical quantitative genetic models to empirical studies of evolution in wild populations. However, while classical models assume environmental constancy, most natural populations exist in variable environments. Here, we applied a novel analytical technique to a long-term study of birthweight in wild sheep and examined, for the first time, how variation in environmental quality simultaneously influences the strength of natural selection and the genetic basis of trait variability. In addition to demonstrating that selection and genetic variance vary dramatically across environments, our results show that environmental heterogeneity induces a negative correlation between these two parameters. Harsh environmental conditions were associated with strong selection for increased birthweight but low genetic variance, and vice versa. Consequently, the potential for microevolution in this population is constrained by either a lack of heritable variation ( in poor environments) or by a reduced strength of selection ( in good environments). More generally, environmental dependence of this nature may act to limit rates of evolution, maintain genetic variance, and favour phenotypic stasis in many natural systems. Assumptions of environmental constancy are likely to be violated in natural systems, and failure to acknowledge this may generate highly misleading expectations for phenotypic microevolution

Mating system of the Eurasian badger, Meles meles, in a high density population

Badgers are facultatively social, forming large groups at high density. Group-living appears to have high reproductive costs for females, and may lead to increased levels of inbreeding. The extent of female competition for reproduction has been estimated from field data, but knowledge of male reproductive success and the extent of extra-group paternity remains limited. Combining field data with genetic data (16 microsatellite loci), we studied the mating system of 10 badger social groups across 14 years in a high-density population. From 923 badgers, including 425 cubs, we were able to assign maternity to 307 cubs, with both parents assigned to 199 cubs (47%) with 80% confidence, and 14% with 95% confidence. Age had a significant effect on the probability of reproduction, seemingly as a result of a deficit of individuals aged two years and greater than eight years attaining parentage. We estimate that approximately 30% of the female population successfully reproduced in any given year, with a similar proportion of the male population gaining paternity across the same area. While it was known there was a cost to female reproduction in high density populations, it appears that males suffer similar, but not greater, costs. Roughly half of assigned paternity was attributed to extra-group males, the majority of which were from neighbouring social groups. Few successful matings occurred between individuals born in the same social group (22%). The high rate of extra-group mating, previously unquantified, may help reduce inbreeding, potentially making philopatry a less costly strategy

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. Read More: http://www.esajournals.org/doi/abs/10.1890/13-0967.

Parasite infections in a social carnivore: Evidence of their fitness consequences and factors modulating infection load

There are substantial individual differences in parasite composition and infection load in wildlife populations. Few studies have investigated the factors shaping this heterogeneity in large wild mammals or the impact of parasite infections on Darwinian fitness, particularly in juveniles. A host's parasite composition and infection load can be shaped by factors that determine contact with infective parasite stages and those that determine the host's resistance to infection, such as abiotic and social environmental factors, and age. Host–parasite interactions and synergies between coinfecting parasites may also be important. We test predictions derived from these different processes to investigate factors shaping infection loads (fecal egg/oocyte load) of two energetically costly gastrointestinal parasites: the hookworm Ancylostoma and the intracellular Cystoisospora, in juvenile spotted hyenas (Crocuta crocuta) in the Serengeti National Park, in Tanzania. We also assess whether parasite infections curtail survival to adulthood and longevity. Ancylostoma and Cystoisospora infection loads declined as the number of adult clan members increased, a result consistent with an encounter‐reduction effect whereby adults reduced encounters between juveniles and infective larvae, but were not affected by the number of juveniles in a clan. Infection loads decreased with age, possibly because active immune responses to infection improved with age. Differences in parasite load between clans possibly indicate variation in abiotic environmental factors between clan den sites. The survival of juveniles (<365 days old) to adulthood decreased with Ancylostoma load, increased with age, and was modulated by maternal social status. High‐ranking individuals with low Ancylostoma loads had a higher survivorship during the first 4 years of life than high‐ranking individuals with high Ancylostoma loads. These findings suggest that high infection loads with energetically costly parasites such as hookworms during early life can have negative fitness consequences

Low genetic variability, female-biased dispersal and high movement rates in an urban population of Eurasian badgersMeles meles

1. Urban and rural populations of animals can differ in their behaviour, both in order to meet their ecological requirements and due to the constraints imposed by different environments. The study of urban populations can therefore offer useful insights into the behavioural flexibility of a species as a whole, as well as indicating how the species in question adapts to a specifically urban environment. 2. The genetic structure of a population can provide information about social structure and movement patterns that is difficult to obtain by other means. Using non-invasively collected hair samples, we estimated the population size of Eurasian badgers Meles meles in the city of Brighton, England, and calculated population-specific parameters of genetic variability and sex-specific rates of outbreeding and dispersal. 3. Population density was high in the context of badger densities reported throughout their range. This was due to a high density of social groups rather than large numbers of individuals per group. 4. The allelic richness of the population was low compared with other British populations. However, the rate of extra-group paternity and the relatively frequent (mainly temporary) intergroup movements suggest that, on a local scale, the population was outbred. Although members of both sexes visited other groups, there was a trend for more females to make intergroup movements. 5. The results reveal that urban badgers can achieve high densities and suggest that while some population parameters are similar between urban and rural populations, the frequency of intergroup movements is higher among urban badgers. In a wider context, these results demonstrate the ability of non-invasive genetic sampling to provide information about the population density, social structure and behaviour of urban wildlife