Tim Coulson

Noise and determinism in synchronized sheep dynamics

A major debate in ecology concerns the relative importance of intrinsic factors and extrinsic environmental variations in determining population size fluctuations. Spatial correlation of fluctuations in different populations caused by synchronous environmental shocks,, is a powerful tool for quantifying the impact of environmental variations on population dynamics,. However, interpretation of synchrony is often complicated by migration between populations,. Here we address this issue by using time series from sheep populations on two islands in the St Kilda archipelago. Fluctuations in the sizes of the two populations are remarkably synchronized over a 40-year period. A nonlinear time-series model shows that a high and frequent degree of environmental correlation is required to achieve this level of synchrony. The model indicates that if there were less environmental correlation, population dynamics would be much less synchronous than is observed. This is because of a threshold effect that is dependent on population size; the threshold magnifies random differences between populations. A refined model showsthat part of the required environmental synchronicity can be accounted for by large-scale weather variations. These results underline the importance of understanding the interaction between intrinsic and extrinsic influences on population dynamics.

Microsatellites reveal heterosis in red deer

The fitness consequences of inbreeding and outbreeding are poorly understood in natural populations. We explore two microsatellite–based variables, individual heterozygosity (likely to correlate with recent inbreeding) and a new individual–specific internal distance measure, meand2 (focusing on events deeper in the pedigree), in relation to two measures of fitness expressed early in life, birth weight and neonatal survival, in 670 red deer calves (Cervus elaphus) born on the Isle of Rum between 1982 and 1996. For comparison, we also analyse inbreeding coefficients derived from pedigrees in which paternity was inferred by molecular methods.Only 14 out of 231 calves (6.1%) had non–zero inbreeding coefficients, and neither inbreeding coefficient nor individual heterozygosity was consistently related to birth weight or neonatal survival. However, meand2 was consistently related to both fitness measures. Low meand2 was associated with low birth weight, especially following cold Aprils, in which foetal growth is reduced. Low meand2 was also associated with low neonatal survival, but this effect was probably mediated by birth weight because fitting birth weight to the neonatal survival model displaced meand2 as an explanatory variable. We conclude that in the deer population fitness measures expressed early in life do not show evidence of inbreeding depression, but they do show evidence of heterosis, possibly as a result of population mixing. We also demonstrate the practical problems of estimating inbreeding via pedigrees compared with a direct marker–based estimate of individual heterozygosity. We suggest that, together, individual heterozygosity and meand2, estimated using microsatellites, are useful tools for exploring inbreeding and outbreeding in natural populations.

The use of photographic rates to estimate densities of tigers and other cryptic mammals

The monitoring and management of species depends on reliable population estimates, and this can be both difficult and very costly for cryptic large vertebrates that live in forested habitats. Recently developed camera trapping techniques have already been shown to be an effective means of making mark-recapture estimates of individually identifiable animals (e.g. tigers). Camera traps also provide a new method for surveying animal abundance. Through computer simulations, and an analysis of the rates of camera trap capture from 19 studies of tigers across the species range, we show that the number of camera days/tiger photograph correlates with independent estimates of tiger density. This statistic does not rely on individual identity and is particularly useful for estimating the population density of species that are not individually identifiable. Finally, we used the comparison between observed trapping rates and the computer simulations to estimate the minimum effort required to determine that tigers,n or other species, do not exist in an area, a measure that is critical for conservation planning.

Sexually antagonistic genetic variation for fitness in red deer.

Evolutionary theory predicts the depletion of genetic variation in natural populations as a result of the effects of selection, but genetic variation is nevertheless abundant for many traits that are under directional or stabilizing selection. Evolutionary geneticists commonly try to explain this paradox with mechanisms that lead to a balance between mutation and selection. However, theoretical predictions of equilibrium genetic variance under mutation–selection balance are usually lower than the observed values, and the reason for this is unknown. The potential role of sexually antagonistic selection in maintaining genetic variation has received little attention in this debate, surprisingly given its potential ubiquity in dioecious organisms. At fitness-related loci, a given genotype may be selected in opposite directions in the two sexes. Such sexually antagonistic selection will reduce the otherwise-expected positive genetic correlation between male and female fitness. Both theory and experimental data suggest that males and females of the same species may have divergent genetic optima, but supporting data from wild populations are still scarce. Here we present evidence for sexually antagonistic fitness variation in a natural population, using data from a long-term study of red deer (Cervus elaphus). We show that male red deer with relatively high fitness fathered, on average, daughters with relatively low fitness. This was due to a negative genetic correlation between estimates of fitness in males and females. In particular, we show that selection favours males that carry low breeding values for female fitness. Our results demonstrate that sexually antagonistic selection can lead to a trade-off between the optimal genotypes for males and females; this mechanism will have profound effects on the operation of selection and the maintenance of genetic variation in natural populations.

Empirical evidence of density-dependence in populations of large herbivores

Density‐dependence is a key concept in population dynamics. Here, we review how body mass and demographic parameters vary with population density in large herbivores. The demographic parameters we consider are age‐ and sex‐specific reproduction, survival and dispersal. As population density increases, the body mass of large herbivores typically declines, affecting individual performance traits such as age of first reproduction and juvenile survival. We documented density‐dependent variations in reproductive rates for many species from the Arctic to subtropical zones, both with and without predation. At high density, a trade‐off between growth and reproduction delays the age of primiparity and often increases the costs of reproduction, decreasing both survival and future reproductive success of adult females. Density‐dependent preweaning juvenile survival occurs more often in polytocous than monotocous species, while the effects of density on post‐weaning juvenile survival are independent of litter size. Responses of adult survival to density are much less marked than for juvenile survival, and may be exaggerated by density‐dependent changes in age structure. The role of density‐dependent dispersal in population dynamics remains uncertain, because very few studies have examined it. For sexually dimorphic species, we found little support for higher sensitivity to increasing density in the life history traits of males compared to females, except for young age classes. It remains unclear whether males of dimorphic species are sensitive to male density, female density or a combination of both. Eberhardts model predicting a sequential effect of density on demographic parameters (from juvenile survival to adult survival) was supported by 9 of 10 case studies. In addition, population density at birth can also lead to cohort effects, including a direct effect on juvenile survival and longterm effects on average cohort performance as adults. Density effects typically interact with weather, increasing in strength in years of harsh weather. For some species, the synchronization between plant phenology and reproductive cycle is a key process in population dynamics. The timing of late gestation as a function of plant phenology determines whether density‐dependence influences juvenile survival or adult female reproduction. The detection of density‐dependence can be made difficult by nonlinear relationships with density, high sampling variability, lagged responses to density changes, changes in population age structure, and temporal variation in the main factors limiting population growth. The negative feedbacks of population size on individual performance, and hence on life history traits, are thus only expected in particular ecological contexts and are most often restricted to certain age‐specific demographic traits.

Conservation: Reproductive collapse in saiga antelope harems

A common assumption is that breeding in polygnous systems is not limited by the number of males because one male can inseminate many females. But here we show that reproductive collapse in the critically endangered saiga antelope (Saiga tatarica tatarica) is likely to have been caused by a catastrophic drop in the number of adult males in this harem-breeding ungulate, probably due to selective poaching for their horns. Fecundity and calf survival are known to be affected by markedly skewed sex ratios, but in the saiga antelope the sex ratio has become so distorted as to lead to a drastic decline in the number of pregnancies — a finding that has implications both for the conservation of the species and for understanding the reproductive ecology of polygynous ungulates.

The Evolutionary Demography of Ecological Change: Linking Trait Variation and Population Growth

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.

The relative roles of density and climatic variation on population dynamics and fecundity rates in three contrasting ungulate species

The relative influences of density–dependent and–independent processes on vital rates and population dynamics have been debated in ecology for over half a century, yet it is only recently that both processes have been shown to operate within the same population. However, generalizations on the role of each process across species are rare. Using a process–orientated generalized linear modelling approach we show that variations in fecundity rates in populations of three species of ungulates with contrasting life histories are associated with density and winter weather in a remarkably similar manner. However, there are differences and we speculate that they are a result of differences in size between the species. Much previous research exploring the association between vital rates, population dynamics and densitydependent and–independent processes has used pattern–orientated approaches to decompose time–series into contributions from density–dependent and–independent processes. Results from these analyses are sometimes used to infer associations between vital rates, density and climatic variables. We compare results from pattern–orientated analyses of time–series with process–orientated analyses and report that the two approaches give different results. The approach of analysing relationships between vital rates, density and climatic variables may detect important processes influencing population dynamics that timeseries methodologies may overlook.

POPULATION SUBSTRUCTURE, LOCAL DENSITY, AND CALF WINTER SURVIVAL IN RED DEER (CERVUS ELAPHUS)

Population substructure and the effects of scale have recently received much theoretical attention, but few studies have examined these factors in free-living populations of vertebrates. We used > 200 000 sightings of recognized females recorded over a con- tinuous 20-yr period to explore population substructure and spatial heterogeneity in red deer on the Isle of Rum, Scotland. We used hierarchical cluster analysis to group individuals together by their proximities in space, and we explored the influence of scale, considering scales ranging between the whole population and groups of one or two individuals. Inter- mediate scales were isolated as being the best at describing calf winter survival, the key factor in determining future population density. The most statistically explanatory scale isolated a population substructure related to vegetation, with higher local densities occurring around herb-rich Festuca-Agrostis grassland. Calves at high local density were most likely to die. Patterns of local population density varied between seasons in relation to food availability. High-resolution scales were the best descriptors of calf winter survival in summer; coarser scales were better in winter. In both summer and winter, local population density was more important than total population density in influencing calf winter survival. The effects on calf survival of local population density during the summer interacted significantly with calf sex and the mothers reproductive status. In this study, the technique of grouping animals by their proximity in space was more realistic and informative than discrete spatial divisions of the study area.

THE DEMOGRAPHIC CONSEQUENCES OF RELEASING A POPULATION OF RED DEER FROM CULLING

A change in population density can generate spatial and demographic effects that can have an impact on fluctuations in population size for many years. Although the demographic effects of time lags have been incorporated into analyses of time series data, there are few detailed descriptions of the long-term demographic consequences of a change in density. We use detailed, individual-based data from a population of red deer ( Cervus elaphus) from the North Block of the Isle of Rum, Scotland, to describe long-term de- mographic and spatial effects of a change in density. The population was released from hunting pressure in 1972. Over the following 10 years population density doubled and, since the early 1980s, has fluctuated around ecological carrying capacity. The cessation of culling led to long-term transient spatial and demographic effects that have persisted for 30 years. Different vital rates responded to the increase in density at different rates, causing long-term changes to the demographic and spatial structure of the population. These changes altered the impact of different age- and sex-specific vital rates on annual changes in pop- ulation size. These changes are still ongoing, 30 years after cessation of the cull, suggesting that a change in density may generate transient dynamics that persist for several generations.