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1. Given energetic constraints, female reproductive strategy is expected to be shaped by a trade-off between offspring size and offspring number, the optimal resolution of which may vary with environmental conditions. 2. We tested the hypothesis that selection will favour the production of larger litters, even at some expense to offspring size, under good conditions (and vice versa in harsh environments) using data from a long-term study of an unmanaged population of Soay sheep on the islands of St Kilda, NW Scotland. 3. Both litter size (which is either 1 or 2) and offspring birth weight are under positive selection through female annual fitness, but the strength of selection varies systematically with environmental conditions. Age effects were also detected, with selection weakening as female age increases. 4. Consistent with theoretical predictions, the selective trade-off between litter and offspring size is shifted in favour of the latter under poor environmental conditions. Thus, direct selection on offspring birth weight increases under harsh environmental conditions, particularly for females producing twins. 5. However, singletons are only favoured when environmental conditions are very poor, and offspring weight is constrained to be low. Thus, the current low incidence of twinning (13·5% of litters produced since 1985) appears to be suboptimal with respect to the empirically estimated selection regime. Thus, litter size, a trait known to be heritable, may be expected to evolve upwards in this population. 6. Our study highlights the necessity of incorporating environmental heterogeneity and age structure into analyses of natural selection, and suggests that the common assumption of optimality used in models of life history may sometimes be problematic. [ABSTRACT FROM AUTHOR]

Van Tienderen recently published a method that links selection gradients between a phenotypic trait and multiple fitness components with the effects of these fitness components on the population growth rate (mean absolute fitness). The method allows selection to be simultaneously estimated across multiple fitness components in a population dynamic framework. In this paper we apply the method to a population of red deer living in the North Block of the Isle of Rhum, Scotland. We show that (1) selection on birth date and birth weight can operate through multiple fitness components simultaneously; (2) our estimates of the response to selection are consistent with the observed change in trait values that we cannot explain with environmental and phenotypic covariates; (3) selection on both traits has fluctuated over the course of the study; (4) selection operates through different fitness components in different years; and (5) no environmental covariates correlate with selection because different fitness components respond to density and climatic variation in contrasting ways. [ABSTRACT FROM AUTHOR]

Summary 1. We use structured demographic accounting to decompose the contribution to the variance in relative population growth between 1971 and 1997 in female red deer on Rum, Scotland, in terms of different fitness components and age categories. 2. During the first 10 years (1971–80) the variance in relative population growth was small (0·00246) as the population grew from 58 to 148 having been culled previously. After this the relative variance in population growth increased to 0·00575 between 1981 and 1989, and increased again to 0·00848 between 1990 and 1997. 3. In the first phase of population growth (1971–80) changes in the birth rate contributed most to the variance in relative population growth (30%), then adult winter survival (23%), with calf summer and calf winter survival contributing less than 10%. Birth rate tended to covary strongly with adult winter survival, and to a lesser extent with calf winter survival. 4. The contribution of birth rate to the variance in relative population growth diminished to less than 5% in both the periods 1981–89 and 1990–97. In contrast, the contribution of adult winter survival increased to 36% in 1980s and 40% in 1990s. At the same time, the covariation of adult winter survival with calf winter survival increased successively to reach 40% in 1990–97. 5. Changes in population age structure were small and contributed little to the variance in relative population growth (≈ 1% over the entire study). In the 1970s the 5–7-year-old group contributed most (23%) to the variance in relative population growth, though this covaried with 8–12-year-olds. In the 1980s the 8–12-year-olds contributed most (27%) and now the covariation was with older animals (> 12-year-olds). This pattern persisted in the 1990s, though there was covariation with 5–7-year-olds too. 6. Birth rate, calf winter survival and adult winter survival were all... [ABSTRACT FROM AUTHOR]

1. Long-term studies allow the outcomes of repeated selection events to be monitored. Here, we investigate phenotypic selection in successive winter mortality events in the Soay sheep of St Kilda, Scotland, between 1985 and 1996. Selection of three quantitative morphometric traits, body weight, hindleg length and incisor arcade breadth, was investigated in different sectors of the population. 2. Evidence from fitness differentials of positive directional selection for large size was repeatedly found in lambs and adult females. Selection in the opposing direction was only found in one year in lambs. 3. Selection gradients showed that in most years when significant selection occurred, body weight was the focus of direct selection, whereas selection of hindleg length and incisor breadth was indirect, arising from their correlation with body weight. 4. Selection was strongest in years of low over-winter survival and almost absent in years when survival was high. Intensity of selection was greatest in lambs, emphasizing the differences in selection pressure experienced by different sectors of the population, in addition to the temporal variation in selection pressure due to population density and environmental conditions. 5. Despite repeated positive selection of body weight, no evidence of a change in the population mean was found over the course of the study. [ABSTRACT FROM AUTHOR]

Inbreeding depression is of major concern for the conservation of threatened species, and inbreeding avoidance is thought to be a key driver in the evolution of mating systems. However, the estimation of individual inbreeding coefficients in natural populations has been challenging, and, consequently, the full effect of inbreeding on fitness remains unclear. Genomic inbreeding coefficients may resolve the long-standing paucity of data on inbreeding depression in adult traits and total fitness. Here we investigate inbreeding depression in a range of life history traits and fitness in a wild population of red deer (Cervus elaphus) in Scotland using individual inbreeding coefficients derived from dense Single-Nucleotide Polymorphism (SNP) data (Fgrm). We find associations between[Formula: see text]and annual breeding success in both sexes, and between maternal inbreeding coefficient and offspring survival. We also confirm previous findings of inbreeding depression in birth weight and juvenile survival. In contrast, inbreeding coefficients calculated from a deep and comparatively complete pedigree detected inbreeding depression in juvenile survival, but not in any adult fitness component. The total effect of inbreeding on lifetime breeding success (LBS) was substantial in both sexes: for Fgrm = 0.125, a value resulting from a half-sib mating, LBS declined by 72% for females and 95% for males. Our results demonstrate that SNP-based estimates of inbreeding provide a powerful tool for evaluating inbreeding depression in natural populations, and suggest that, to date, the prevalence of inbreeding depression in adult traits may have been underestimated.

1. Trade-offs among life-history traits are common because individuals have to partition limited resources between multiple traits. Reproductive costs are generally assumed to be high, resulting in reduced survival and fecundity in the following year. However, it is common to find positive rather than negative correlations between life-history traits. 2. Here, we use a data set from the individual-based study of red deer on the Isle of Rum to examine how these costs vary between individuals and at different ages, using multi-state mark-recapture methodology. 3. Females that had reproduced frequently in the past incurred lower costs of reproduction in terms of survival in the following year and were more likely to reproduce in two consecutive years. Older individuals and those that had not reproduced frequently exhibited higher costs. 4. These results highlight the importance of considering heterogeneity and individual quality when examining trade-offs and demonstrate the effectiveness of using detailed long-term data sets to explore life-history strategies using multi-state mark-recapture models., (© 2010 The Authors. Journal of Animal Ecology © 2010 British Ecological Society.)

We used long-term data on movements, survival, and reproduction of female red deer (Cervus elaphus Linnaeus) of the Isle of Rum, Scotland (1970-2001), to explain variation in life history (age at maturity) from a hind's access to habitat resources and exposure to local density, and cross-generational (maternal) effects on observed relationships. We described each hind's use of resources relative to availability in the study area from an individual-based resource selection function (RSF); we defined local density as the total number of hinds aged > or = 1 year within the subpopulation cluster to which an individual belonged. The likelihood of a hind producing her first calf in the period from birth to the end of the summer in which she turned age 3 was positively related to relative use of Agrostis/Festuca grasslands and other low-elevation communities, and inversely related to a hind's mean annual local density. However, when we considered both a daughter's RSF and exposure to local density and her mother's RSF and exposure to local density, maternal data alone most parsimoniously explained variation in age at maturity of daughters. Mothers were able to lower age at maturity in their daughters in two, non-mutually exclusive ways. First, birth mass of daughters was inversely related to age at maturity, and mothers that used relatively less uplands (Calluna-dominated heath and heather moorland) and occupied areas of lower density produced larger offspring. Second, mothers could establish a home range that enabled daughters to mature in areas with access to high quality Agrostis/Festuca grasslands at low density. Lifetime reproductive success was inversely associated with a hind's age at maturity via extension of the reproductive life span. Longevity did not change in association with age at maturity. Patterns in how animals use available habitat resources may depend on that of previous generations, especially at larger scales of resource selection.

Two contrasting approaches to the analysis of population dynamics are currently popular: demographic approaches where the associations between demographic rates and statistics summarizing the population dynamics are identified; and time series approaches where the associations between population dynamics, population density, and environmental covariates are investigated. In this paper, we develop an approach to combine these methods and apply it to detailed data from Soay sheep (Ovis aries). We examine how density dependence and climate contribute to fluctuations in population size via age- and sex-specific demographic rates, and how fluctuations in demographic structure influence population dynamics. Density dependence contributes most, followed by climatic variation, age structure fluctuations and interactions between density and climate. We then simplify the density-dependent, stochastic, age-structured demographic model and derive a new phenomenological time series which captures the dynamics better than previously selected functions. The simple method we develop has potential to provide substantial insight into the relative contributions of population and individual-level processes to the dynamics of populations in stochastic environments.

Population dynamics and evolutionary change are linked by the fundamental biological processes of birth and death. This means that population growth may correlate with the strength of selection, whereas evolutionary change can leave an ecological signature. We decompose population growth in an age-structured population into contributions from variation in a quantitative trait. We report that the distribution of body sizes within a population of Soay sheep can markedly influence population dynamics, accounting for up to one-fifth of observed population growth. Our results suggest that there is substantial opportunity for evolutionary dynamics to leave an ecological signature and visa versa.

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.

Individuals are predicted to maximize lifetime reproductive success (LRS) through selective use of resources; however, a wide range of ecological and social processes may prevent individuals from always using the highest-quality resources available. Resource selection functions (RSFs) estimate the relative amount of time an individual spends using a resource as a function of the proportional availability of that resource. We quantified the association between LRS and coefficients of individual-based RSFs describing lifetime resource selection for 267 female red deer (Cervus elaphus) of the Isle of Rum, Scotland, from 1970 to 2001. LRS was significantly related to first- and second-order effects of selection for Agrostis/Festuca grassland and proximity to the sea coast (quality of forage within Agrostis/Festuca grassland was highest nearest the coast (ratio of short:long grassland)). The benefits of selecting for quality in Agrostis/Festuca grassland, however, traded-off with increases in LRS gained by avoiding conspecific density. LRS was inversely associated with local density, which was highest along the coast, and reproductive benefits of selecting Agrostis/Festuca grassland diminished with increasing density. We discuss the relevance of these results to our understanding of the spatial distribution of red deer abundance, and potential applications of our approach to evolutionary and applied ecology.

A growing literature now documents the presence of fine-scale genetic structure in wild vertebrate populations. Breeding population size, levels of dispersal and polygyny--all hypothesized to affect population genetic structure--are known to be influenced by ecological conditions experienced by populations. However the possibility of temporal or spatial variation in fine-scale genetic structure as a result of ecological change is rarely considered or explored. Here we investigate temporal variation in fine-scale genetic structure in a red deer population on the Isle or Rum, Scotland. We document extremely fine-scale spatial genetic structure (< 100 m) amongst females but not males across a 24-year study period during which resource competition has intensified and the population has reached habitat carrying capacity. Based on census data, adult deer were allocated to one of three subpopulations in each year of the study. Global F(ST) estimates for females generated using these subpopulations decreased over the study period, indicating a rapid decline in fine-scale genetic structure of the population. Global F(ST) estimates for males were not different from zero across the study period. Using census and genetic data, we illustrate that, as a consequence of a release from culling early in the study period, the number of breeding females has increased while levels of polygyny have decreased in this population. We found little evidence for increasing dispersal between subpopulations over time in either sex. We argue that both increasing female population size and decreasing polygyny could explain the decline in female population genetic structure.

While vocal tract resonances or formants are key acoustic parameters that define differences between phonemes in human speech, little is known about their function in animal communication. Here, we used playback experiments to present red deer stags with re-synthesized vocalizations in which formant frequencies were systematically altered to simulate callers of different body sizes. In response to stimuli where lower formants indicated callers with longer vocal tracts, stags were more attentive, replied with more roars and extended their vocal tracts further in these replies. Our results indicate that mammals other than humans use formants in vital vocal exchanges and can adjust their own formant frequencies in relation to those that they hear.

Understanding how climate can interact with other factors in determining patterns of species abundance is a persistent challenge in ecology. Recent research has suggested that the dynamics exhibited by some populations may be a non-additive function of climate, with climate affecting population growth more strongly at high density than at low density. However, we lack methodologies to adequately explain patterns in population growth generated as a result of interactions between intrinsic factors and extrinsic climatic variation in non-linear systems. We present a novel method (the Functional Coefficient Threshold Auto-Regressive (FCTAR) method) that can identify interacting influences of climate and density on population dynamics from time-series data. We demonstrate its use on count data on the size of the Soay sheep population, which is known to exhibit dynamics generated by nonlinear and non-additive interactions between density and climate, living on Hirta in the St Kilda archipelago. The FCTAR method suggests that climate fluctuations can drive the Soay sheep population between different dynamical regimes--from stable population size through limit cycles and non-periodic fluctuations.

Grazing by hill sheep and red deer prevents the regeneration of woodland in many parts of the Scottish Highlands and has also led to extensive loss of heather cover. Conservation bodies claim that there has been a rapid rise in Highland deer numbers caused by inadequate management and that these need to be drastically reduced. Here we show that the recent increase in red deer stocks has probably been overestimated and suggest that the gradual rise in numbers since 1970 may be a consequence of a reduction in sheep stocks and of changes in winter weather, rather than of a reduction in culling rate. Although there would be environmental benefits in reducing deer numbers, there is an equal need to reduce the numbers of hill sheep in many parts of the Highlands.

It has been suggested that fluctuating asymmetry (FA) in secondary sexual traits may be a useful indicator of either individual quality or environmental stress. We tested this concept using a series of analyses of FA in male antler size in a wild red deer (Cervus elaphus) population, using four measures of size repeated across successive years on the same individuals. We found no consistent evidence of correlations between traits in levels of FA, nor of any associations between known environmental or developmental conditions. None of the four measures of FA showed a significant heritability (average h2 = 0.041), nor was there any evidence of inbreeding depression. For three of the four traits, fluctuating asymmetry did not predict either annual or lifetime breeding success. However there were significant associations between breeding success and FA in antler length. Given the series of null results in our other tests, it seems likely that this was a direct mechanistic effect rather than because measures of FA were indicative of individual quality or condition.

Populations of red deer that are limited by food, like those of many other ungulates, commonly include more females than males. We assessed the contribution of variation in sex- and age-specific rates of mortality and emigration to density-dependent changes in the adult sex ratio, using long-term observations and demographic experiments involving the red deer population on Rum, Scotland. We incorporated these effects in a stochastic model of local populations under different management regimes to show here that, when female numbers are allowed to increase to more than 60% of the ecological carrying capacity, the sustainable annual harvest of males from local deer populations will fall. Because males are typically culled by fee-paying hunters and generate more income than females, income will decrease as the male harvest falls. Because numbers of female deer throughout much of the Highlands probably exceed the threshold at which male density starts to be affected, many managers might be able to raise income from local deer populations by reducing female numbers, with potential benefits to the vegetation of Scottish Highland environments.

Evolutionary and conservation biologists have a long-standing interest in the consequences of inbreeding. It is generally recognized that inbred individuals may experience reduced fitness or inbreeding depression. By the same token, relatively outbred individuals can have greater than average fitness, i.e. heterosis. However, nearly all of the empirical evidence for inbreeding depression comes from laboratory or domestic species. Inbreeding depression and heterosis are difficult to detect in natural populations due to the difficulties in establishing pedigrees. An alternative method is to correlate heterozygosity, which is measured using genetic markers, with a trait related to fitness. The typically studied traits, such as juvenile survival and growth rates, either cover only early life or are weakly correlated with lifetime breeding success (LBS). In this paper we show that heterozygosity is positively associated with male and female adult LBS in a wild population of red deer (Cervus elaphus) on the Isle of Rum, Scotland. To the authors' knowledge, this is the first time that inbreeding depression and/or heterosis have been detected for a trait highly correlated with overall fitness in both sexes in a wild population.

Classical population genetics theory predicts that selection should deplete heritable genetic variance for fitness. We show here that, consistent with this prediction, there was a negative correlation between the heritability of a trait and its association with fitness in a wild population of red deer (Cervus elaphus) and there was no evidence of significant heritability of total fitness. However, the decline in heritability was caused, at least in part, by increased levels of residual variance in longevity and, hence, in total fitness: in this population, longevity is known to be heavily influenced by environmental factors. Other life history traits that were not associated with longevity, such as average annual breeding success, had higher heritabilities. Coefficients of additive genetic variance differed markedly between traits, but highly skewed measures, such as male breeding success, generally had greater coefficients of variance than morphometric traits. Finally, there were significant maternal effects in a range of traits, particularly for females.

In sexually dimorphic mammals, high population density is commonly associated with increased mortality of males relative to females and with female-biased adult sex ratios. This paper investigates the consequences of these changes on the distribution of male breeding success, the intensity of competition for females and the opportunity for sexual selection. After the red deer (Cervus elaphus L.) population of the North Block of Rum (Inner Hebrides) was released from culling, female numbers rose and male numbers declined, leading to an adult sex ratio of around one male to two females. This change was the result of increased mortality of males relative to females during the first two years of life; of increased emigration rates by young males; and of reduced immigration by males from outside the study area. The increasing bias in the adult sex ratio affected the timing of breeding as well as the distribution of mating success in males. As the adult sex ratio became increasingly biased towards females, the degree of skew in mating success (calculated across all harem-holders) increased, but mature males defended harems for shorter periods and a higher proportion of males held harems. In addition, a higher proportion of calves were fathered by immigrant males and the proportion fathered by males born in the study area declined. These results support the contention that, where high population density is associated with a female-biased adult sex ratio, competition for mates is likely to decline.

Theoretical models of the effect of population bottlenecks on genetic variation assume that individuals are removed at random from the population. We investigated this assumption in a naturally regulated, unstable population of Soay sheep (Ovis aries). During rapid population declines or 'crashes', individuals were not removed at random with respect to genotype: we found associations between individual survival and certain genotypes at five polymorphic protein or microsatellite DNA loci (Ada, Got, Tf, MAF18 and OPACAP). Some loci appeared to show simple associations with survival whereas others had more complex interactions with crash year or age: all displayed different patterns of association between the sexes. Simple overdominance was not a general feature of our data; it seems likely that fluctuating selecting, countervailing selection in different fitness components or frequency-dependent selection may explain the pattern and complexity of the associations shown at different loci. Our study cannot distinguish between selection acting at these loci or at other, closely linked loci. However, our empirical study implies that the molecular genetic outcome of population bottlenecks in natural populations does not always follow theoretical expectations based on the random removal of genotypes. Bottlenecks in which individuals are removed at random are distinct from bottlenecks in which there is scope for selection via non-random survival of individuals.

In a population of free-ranging red deer hinds on the Isle of Rhum (Inner Hebrides) we investigated relationships between four aspects of reproductive performance (fertility, calf birth weight, birth sex ratio and calving date) and four variables likely to affect the mother's condition: age, reproductive status, home range area and year of calving. Fertility was significantly related to mother's age, reproductive status and home range area as well as to year of calf's birth. Stag calves were heavier than hind calves and birth weight was significantly related to mother's age, home range area and year of (calf's) birth but not to mother's reproductive status. Birth sex ratio did not differ from parity, and was not significantly associated with any of the four variables examined. Birth date was significantly related to the mother's reproductive status, home range area and year of (calf's) birth but not to mother's age or the sex of the calf.