Costs of mating competition limit male lifetime breeding success in polygynous mammals.

This is the accepted version of the paper. The final published version is available from the Royal Society at http://rspb.royalsocietypublishing.org/content/281/1786/20140418.abstract.Although differences in breeding lifespan are an important source of variation in male fitness, the factors affecting the breeding tenure of males have seldom been explored. Here, we use cross-species comparisons to investigate the correlates of breeding lifespan in male mammals. Our results show that male breeding lifespan depends on the extent of polygyny, which reflects the relative intensity of competition for access to females. Males have relatively short breeding tenure in species where individuals have the potential to monopolize mating with multiple females, and longer ones where individuals defend one female at a time. Male breeding tenure is also shorter in species in which females breed frequently than in those where females breed less frequently, suggesting that the costs of guarding females may contribute to limiting tenure length. As a consequence of these relationships, estimates of skew in male breeding success within seasons overestimate skew calculated across the lifetime and, in several polygynous species, variance in lifetime breeding success is not substantially higher in males than in females.The Leverhulme Trust, the Isaac Newton Trust, and the European Research Council provided the funding for this study

Validating methods for estimating endocranial volume in individual red deer (Cervus elaphus).

Comparing brain sizes is a key method in comparative cognition and evolution. Brain sizes are commonly validated by interspecific comparisons involving animals of varying size, which does not provide a realistic index of their accuracy for intraspecific comparisons. Intraspecific validation of methods for measuring brain size should include animals of the same age and sex to ensure that individual differences can be detected in animals of similar size. In this study we compare three methods of measuring the endocranial volume of 33 red deer skulls to investigate the accuracy of each method. Methods for estimating endocranial volume included scanning each skull using computerised tomography (CT) and quantifying the volume with OsiriX software, filling the cranium with glass beads and measuring the bead volume, and linear measurements (length, width, and height) of the cranium using callipers. CT scan volumes were highly correlated with results from the bead method, but only moderately correlated with the linear method. This study illustrates the importance of validating intraspecies measurement methods, which allows for the accurate interpretation of results.This is the author accepted manuscript. The final version is available from Elsevier via http://dx.doi.org/10.1016/j.beproc.2012.10.01

Ontogenetic changes in alarm-call production and usage in meerkats ( Suricata suricatta ): adaptations or constraints?

In many species, individuals suffer major mortality in their first year because of predation. Behaviours that facilitate successful escape are therefore under strong selection, but anti-predator skills often emerge gradually during an individual's early development. Using long-term data and acoustic recordings of alarm calls collected during natural predator encounters, we aimed to elucidate two largely unsolved issues in anti-predator ontogeny: (1) whether incorrect predator assignment is adaptively age-appropriate, given that vulnerability often changes during development, or whether age-related differences reflect true mistakes made by immature individuals; and (2) the extent to which the development of adult-like competence in alarm-call production and usage is simply a function of maturational processes or dependent upon experience. We found that young meerkats (Suricata suricatta) were less likely to give alarm calls than adults, but alarmed more in response to non-threatening species compared to adults. However, stimuli that pose a greater threat to young than adults did not elicit more calling from young; this argues against age-related changes in vulnerability as the sole explanation for developmental changes in calling. Young in small groups, who were more likely to watch out for predators, alarmed more than less vigilant young in larger groups. Moreover, despite similarities in acoustic structure between alarm call types, calls appeared in the repertoire at different rates, and those that were associated with frequently encountered predators were produced relatively early on. These results indicate that experience is a more plausible explanation for such developmental trajectories than maturatio

A single trapped ion in a finite range trap

This paper presents a method to describe dynamics of an ion confined in a realistic finite range trap. We model this realistic potential with a solvable one and we obtain dynamical variables (raising and lowering operators) of this potential. We consider coherent interaction of this confined ion in a finite range trap and we show that its center-of-mass motion steady state is a special kind of nonlinear coherent states. Physical properties of this state and their dependence on the finite range of potential are studied

No apparent benefits of allonursing for recipient offspring and mothers in the cooperatively breeding meerkat.

Cooperative behaviours by definition are those that provide some benefit to another individual. Allonursing, the nursing of non-descendent young, is often considered a cooperative behaviour and is assumed to provide benefits to recipient offspring in terms of growth and survival, and to their mothers, by enabling them to share the lactation load. However, these proposed benefits are not well understood, in part because maternal and litter traits and other ecological and social variables are not independent of one another, making patterns hard to discern using standard univariate analyses. Here, we investigate the potential benefits of allonursing in the cooperatively breeding Kalahari meerkat, where socially subordinate females allonurse the young of a dominant pair without having young of their own. We use structural equation modelling to allow us to account for the interdependence of maternal traits, litter traits and environmental factors. We find no evidence that allonursing provides benefits to pups or mothers. Pups that received allonursing were not heavier at emergence and did not have a higher survival rate than pups that did not receive allonursing. Mothers whose litters were allonursed were not in better physical condition, did not reconceive faster and did not reduce their own nursing investment compared to mothers who nursed their litters alone. These patterns were not significantly influenced by whether mothers were in relatively good, or poor, condition. We suggest that allonursing may persist in this species because the costs to allonurses may be low. Alternatively, allonursing may confer other, more cryptic, benefits to pups or allonurses, such as immunological or social benefits.KJM was supported by a research grant from the Cambridge Philosophical Society. KEM was supported by a US National Science Foundation grant to Alison Bell and KEM (NSF IOS 1121980).This is the accepted manuscript. The final version is available at http://onlinelibrary.wiley.com/doi/10.1111/1365-2656.12343/abstract

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

Social and environmental factors affect tuberculosis related mortality in wild meerkats.

Tuberculosis (TB) is an important and widespread disease of wildlife, livestock and humans world-wide, but long-term empirical datasets describing this condition are rare. A population of meerkats (Suricata suricatta) in South Africa's Kalahari Desert have been diagnosed with Mycobacterium suricattae, a novel strain of TB, causing fatal disease in this group-living species. This study aimed to find characteristics associated with clinical TB in meerkats. These characteristics could subsequently be used to identify 'at-risk' animals within a population, and target these individuals for control measures. We conducted a retrospective study based on a unique, long-term life-history dataset of over 2000 individually identified animals covering a 14-year period after the first confirmatory diagnosis of TB in this population in 2001. Individual- and group-level risk factors were analysed using time-dependent Cox regression to examine their potential influence on the time to development of end-stage TB. Cases of disease involved 144 individuals in 27 of 73 social groups, across 12 of 14 years (an incidence rate of 3·78 cases/100 study years). At the individual level, increasing age had the greatest effect on risk of disease with a hazard ratio of 4·70 (95% CI: 1·92-11·53, P < 0·01) for meerkats aged 24-48 months, and a hazard ratio of 9·36 (3·34-26·25, P < 0·001) for animals aged over 48 months (both age categories compared with animals aged below 24 months). Previous group history of TB increased the hazard by a factor of 4·29 (2·00-9·17, P < 0·01), and an interaction was found between this variable and age. At a group level, immigrations of new group members in the previous year increased hazard by a factor of 3·00 (1·23-7·34, P = 0·016). There was weaker evidence of an environmental effect with a hazard ratio for a low rainfall (<200 mm) year of 2·28 (0·91-5·72, P = 0·079). Our findings identify potential individual characteristics on which to base targeted control measures such as vaccination. Additional data on the dynamics of the infection status of individuals and how this changes over time would complement these findings by enhancing understanding of disease progression and transmission, and thus the implications of potential management measures

Effects of cortisol administration on cooperative behavior in meerkat helpers

Experimental administration of stress hormones to meerkat helpers does not affect their contributions to cooperative activities. Previous work on cooperative breeders suggested that levels of stress hormones are related to helping behavior. We tested this experimentally by injecting meerkat helpers with cortisol, whilst matched controls received a saline injection. This did not lead to changes in cooperative behaviour, but induced females, and not males, to spend more time near pups and less time foragin

Flexible alarm calling in meerkats: the role of the social environment and predation urgency

Flexible vocal production has been demonstrated in several vertebrate species, with much work focusing on the role of the social "audience” in explaining variation in call production. It is, however, likely that the decision to call is an emergent property of both external and internal factors, and the extent to which these factors are integrated has been little investigated. We addressed this question by examining the production of alarm calls in wild male meerkats (Suricata suricatta) in different social environments and different predator-encounter contexts. Males searching for reproductive opportunities (rovers) were followed 1) in their home group and when prospecting either 2) solitarily or 3) in a coalition with other males. Results showed that conspecific presence influenced the production of flee-alarm and recruitment calls. Solitary rovers were less likely to produce flee-alarm calls compared with when they are with conspecifics, whether coalitionary rovers or the rover's home group. Experimentally elicited recruitment calls were also produced less when males were solitary than when in their home group. Bark vocalizations, emitted when meerkats were safe, were always produced irrespective of conspecific presence, indicating that these calls function to address predators. The probability of producing flee alarms also increased with the urgency of the predation event. Our results indicate that variation in alarm call production depends on whom the call is addressed to and also on the motivational state of the caller. We argue that neglecting to integrate internal and external factors when elucidating mechanisms underlying vocal production can potentially lead to misguided, parsimonious conclusions regarding vocal flexibility in animal

Differences in cooperative behavior among Damaraland mole rats are consequences of an age-related polyethism.

In many cooperative breeders, the contributions of helpers to cooperative activities change with age, resulting in age-related polyethisms. In contrast, some studies of social mole rats (including naked mole rats, Heterocephalus glaber, and Damaraland mole rats, Fukomys damarensis) suggest that individual differences in cooperative behavior are the result of divergent developmental pathways, leading to discrete and permanent functional categories of helpers that resemble the caste systems found in eusocial insects. Here we show that, in Damaraland mole rats, individual contributions to cooperative behavior increase with age and are higher in fast-growing individuals. Individual contributions to different cooperative tasks are intercorrelated and repeatability of cooperative behavior is similar to that found in other cooperatively breeding vertebrates. Our data provide no evidence that nonreproductive individuals show divergent developmental pathways or specialize in particular tasks. Instead of representing a caste system, variation in the behavior of nonreproductive individuals in Damaraland mole rats closely resembles that found in other cooperatively breeding mammals and appears to be a consequence of age-related polyethism.This study was funded by an European Research Council grant to THCB (294494).This is the author accepted manuscript. The final version is available from the National Academy of Sciences via http://dx.doi.org/10.1073/pnas.160788511