Contrasts in kinship structure in mammalian societies

Comparative studies of mammals confirm Hamilton’s prediction that differences in cooperative and competitive behavior across species will be related to contrasts in kinship between group members. Although theoretical models have explored the factors affecting kinship within social groups, few have analyzed the causes of contrasts in kinship among related species. Here, we describe interspecific differences in average kinship between group members among social mammals and show that a simple mathematical model that includes the number of breeding females, male reproductive skew, and litter size successfully predicts ~95% of observed variation in average kinship between group members across a sample of mammals. Our model shows that a wide range of conditions can generate groups with low average relatedness but only a small and rather specific set of conditions are likely to generate high average levels of relatedness between their members, providing insight into the relative rarity of advanced forms of cooperation in mammalian societies

Maternal longevity and offspring sex in wild ungulates

In species with sexual size dimorphism, the offspring of the larger sex usually have greater energy requirements and may lead to greater fitness costs for parents. The effects of offspring sex on maternal longevity, however, have only been tested in humans. Human studies produced mixed results and considerable debate mainly owing to the difficulty of distinguishing the effects of sexual dimorphism from sociocultural factors. To advance this debate, we examined how the relative number of sons influenced maternal longevity in four species of free-living ungulates (Soay sheep Ovis aries ; bighorn sheep, Ovis canadensis ; red deer, Cervus elaphus ; mountain goat, Oreamnos americanus ), with high male-biased sexual size dimorphism but without complicating sociocultural variables. We found no evidence for a higher cumulative cost of sons than of daughters on maternal longevity. For a given number of offspring, most females with many sons in all four populations lived longer than females with few sons. The higher cost of sons over daughters on maternal lifespan reported by some human studies may be the exception rather than the rule in long-lived iteroparous species

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

Pregnancy is detected via odour in a wild cooperative breeder

Among mammals, scent has long been known to encode oestrus, however in many species detecting pregnancy may also be important in terms of both competition and mate-choice. Here we show, through odour presentation experiments, that pregnancy is discernible via scent by both sexes in the cooperatively breeding banded mongoose. Males spent more time investigating and were more likely to scent mark the odours of non-pregnant females, compared to pregnant females. Females showed increased levels of scent marking when odours were of the same reproductive state as themselves. These results present the first direct demonstration that pregnancy is detectable via scent in wild cooperative breeders. Detecting pregnancy may be particularly important in cooperative breeders as, in addition to the competition between males for receptive mates, there is also intense competition between females for access to alloparental care. Consequently, dominant females benefit from targeting reproductive suppression towards subordinates that represent direct threats, such as pregnant females

The effects of recruitment to direct predator cues on predator responses in meerkats

Behavioral responses of animals to direct predator cues (DPCs; e.g., urine) are common and may improve their survival. We investigated wild meerkat (Suricata suricatta) responses to DPCs by taking an experimental approach. When meerkats encounter a DPC they often recruit group members by emitting a call type, which causes the group members to interrupt foraging and approach the caller. The aim of this study was to identify the qualities of olfactory predator cues, which affect the strength of response by meerkats, and determine the benefits of responses to such cues. Experimental exposure to dog (Canis lupus) urine as a DPC revealed that the recruited individuals increased vigilance to fresh urine in comparison to older urine, whereas a higher quantity of urine did not induce such an effect. Both freshness and higher quantities increased the proportion of group members recruited. These results indicate that recruitment might play a crucial role in correctly assessing the current level of danger and that recruiting might facilitate group decision making. To test the prediction that the reaction to a DPC enhances early predator response, we presented a DPC of a predator and a control cue of a herbivore, and each time simultaneously moved a full-mounted caracal (Caracal caracal) in the vicinity of the group. Meerkats responded earlier to the caracal when the DPC was presented, indicating that the response to a DPC facilitates predator response and that they use information from the cue that reliably reflects the risk in the current momen

On the exactness of the Semi-Classical Approximation for Non-Relativistic One Dimensional Propagators

For one dimensional non-relativistic quantum mechanical problems, we investigate the conditions for all the position dependence of the propagator to be in its phase, that is, the semi-classical approximation to be exact. For velocity independent potentials we find that: (i) the potential must be quadratic in space, but can have arbitrary time dependence. (ii) the phase may be made proportional to the classical action, and the magnitude (``fluctuation factor'') can also be found from the classical solution. (iii) for the driven harmonic oscillator the fluctuation factor is independent of the driving term.Comment: 7 pages, latex, no figures, published in journal of physics

Maternal effects and early-life performance are associated with parasite resistance across life in free-living Soay sheep

Maternal effects occur when the maternal phenotype influences that of the offspring in addition to the effects of maternal genes, and may have a considerable influence on offspring parasite resistance. These effects, and the effects of early levels of reproduction and parasite resistance, may persist into later life and even influence ageing rates. Here we analyse a 20-year longitudinal data set collected on a free-living population of Soay sheep, to investigate the associations between a suite of maternal phenotypic traits and early-life performance on measures of parasite resistance across life. Our results show that maternal effects are important in determining offspring parasite resistance, since lambs born as twins and those born to the youngest and oldest mothers show higher parasite burdens. We show that the association between parasite resistance and natal litter size persists into adulthood. We also show that age-specific changes in parasite resistance in males are associated with natal litter size, and that age-specific changes in females are influenced by early-life levels of reproduction and parasite infection. These results add to the growing evidence that conditions experienced by individuals during development can have a profound influence on immediate and late-life performance and may even influence ageing

The population growth consequences of variation in individual heterozygosity.

Heterozygosity has been associated with components of fitness in numerous studies across a wide range of taxa. Because heterozygosity is associated with individual performance it is also expected to be associated with population dynamics. However, investigations into the association between heterozygosity and population dynamics have been rare because of difficulties in linking evolutionary and ecological processes. The choice of heterozygosity measure is a further issue confounding such studies as it can be biased by individual differences in the frequencies of the alleles studied, the number of alleles at each locus as well as the total number of loci typed. In this study, we first examine the differences between the principal metrics used to calculate heterozygosity using long-term data from a marked population of Soay sheep (Ovis aries). Next, by means of statistical transformation of the homozygosity weighted by loci index, we determine how heterozygosity contributes to population growth in Soay sheep by modelling individual contributions to population growth (p(t(i))) as a function of several covariates, including sex, weight and faecal egg count--a surrogate of parasitic nematode burden in the gut. We demonstrate that although heterozygosity is associated with some components of fitness, most notably adult male reproductive success, in general it is only weakly associated with population growth