Anne Loison

AGE‐SPECIFIC SURVIVAL IN FIVE POPULATIONS OF UNGULATES: EVIDENCE OF SENESCENCE

Methodological problems in describing patterns of senescence in wild pop- ulations have until recently impeded progress in understanding the evolution of a process that decreases individual fitness. We investigated age- and sex-specific survival in five populations of three species of ungulates (roe deer, Capreolus capreolus; bighorn sheep, Ovis canadensis; and isard, Rupicapra pyrenaica), using recent statistical developments of capture-mark-recapture models and long-term (12 to 22 yr) data on marked individuals. The yearly survival of females aged 2-7 yr was remarkably similar and very high (92- 95%) in all five populations. Survival of adult males varied among species and populations. Survival decreased from 8 yr onward for both sexes in all populations, suggesting that senescence was a common phenomenon. Male survival was lower than female survival, and the gender difference increased with age. The extent of sex differences in survival was related neither to sexual dimorphism in mass nor to the level of polygyny, suggesting that species differences in social behavior, particularly mating system and the level of male- male aggression, may be more important than simply the level of polygyny in explaining sexual differences in survival. Our results underline the advantages of long-term monitoring of marked individuals for the study of evolutionary ecology.

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.

Habitat–performance relationships: finding the right metric at a given spatial scale

The field of habitat ecology has been muddled by imprecise terminology regarding what constitutes habitat, and how importance is measured through use, selection, avoidance and other bio-statistical terminology. Added to the confusion is the idea that habitat is scale-specific. Despite these conceptual difficulties, ecologists have made advances in understanding ‘how habitats are important to animals’, and data from animal-borne global positioning system (GPS) units have the potential to help this clarification. Here, we propose a new conceptual framework to connect habitats with measures of animal performance itself—towards assessing habitat–performance relationship (HPR). Long-term studies will be needed to estimate consequences of habitat selection for animal performance. GPS data from wildlife can provide new approaches for studying useful correlates of performance that we review. Recent examples include merging traditional resource selection studies with information about resources used at different critical life-history events (e.g. nesting, calving, migration), uncovering habitats that facilitate movement or foraging and, ultimately, comparing resources used through different life-history strategies with those resulting in death. By integrating data from GPS receivers with other animal-borne technologies and combining those data with additional life-history information, we believe understanding the drivers of HPRs will inform animal ecology and improve conservation.

Consequences of Harvesting on Age Structure, Sex Ratio and Population Dynamics of Red Deer Cervus elaphus in Central Norway

We explore some of the consequences of harvest on population growth rate, age and sex structure in a Norwegian population of red deer Cervus elaphus, using age-structured demographic models. Survival rates were estimated from individuals marked and monitored annually during 1977–1995, and information about reproduction patterns were obtained from hunting material in the same region. The population had an actual doubling time of 14 years, corresponding to a multiplication rate of 1.051. Harvesting led to a reduction of about 10% of the potential multiplication rate that equalled 1.166. Including stochasticity had only a small effect on the population multiplication rate. Due to a high hunting pressure, males had less than a 10% chance of reaching 4.5 years of age and the male-biased harvest strongly biased the sex ratio. Assuming that when the number of females per male increases above a given threshold some females would not manage to mate, we investigated at which level male harvesting could be maintained without having demographic consequences on the population growth rate. We concluded that the hunting pressure on males could probably be increased further but indirect consequences of a strongly biased sex ratio (e.g. on population genetic structure) remain to be studied. Variation in the multiplication rate mainly resulted from the variation in winter calf survival. In its present form the harvesting regime reduces the growth rate and biases the sex and age structure, but does not seem to threaten the populations viability and productivity.

Generation Time: A Reliable Metric to Measure Life-History Variation among Mammalian Populations

Oli and Dobson proposed that the ratio between the magnitude and the onset of reproduction (F/α ratio) allows one to predict the relative importance of vital rates on population growth rate in mammalian populations and provides a reliable measure of the ranking of mammalian species on the slow‐fast continuum of life‐history tactics. We show that the choice of the ratio F/α is arbitrary and is not grounded in demographic theory. We estimate the position on the slow‐fast continuum using the first axis of a principal components analysis of all life‐history variables studied by Oli and Dobson and show that most individual vital rates perform as well as the F/α ratio. Finally, we find, in agreement with previous studies, that the age of first reproduction is a reliable predictor of the ranking of mammalian populations along the slow‐fast continuum and that both body mass and phylogeny markedly influence the generation time of mammalian species. We conclude that arbitrary ratios such as F/α correlate with life‐history types in mammals simply because life‐history variables are highly correlated in response to allometric, phylogenetic, and environmental influences. We suggest that generation time is a reliable metric to measure life‐history variation among mammalian populations and should be preferred to any arbitrary combination between vital rates.

Short- and long-term effects of winter and spring weather on growth and survival of red deer in Norway

Abstract Populations of red deer (Cervus elaphus) in Norway have increased continuously over the last decades. We tested the possible effects of climate and increase in population size on the survival rates and body condition of individuals in one of the northernmost populations of red deer in Europe. Based on 678 individuals of known age marked between 1977 and 1995, we estimated annual survival rates, the probabilities of being harvested and the recapture probability according to sex, age, year, winter and spring weather, population size, and, body weight and body condition, using capture-mark-recapture models. Winter harshness negatively influenced the body weight of yearlings and the survival of calves of both sexes. Spring weather influenced the survival of males in all age classes. A negative trend during the study period was detected in body weight and condition of calves and yearlings, but not in any age- or sex- specific survival rates. No significant gender differences in mean survival were shown in any age class. Moreover, there was little (male) or no (female) detectable between-year variation in survival rates for yearlings and adults. Winter weather acts as a limiting factor on population growth through a short-term effect on first-year survival and a long-term effect on body weight. We discuss the surprising low sex differences in natural survival rates and the differential effects of winter harshness on body weight, body condition and survival in relation to life history characteristics of red deer.

Cohort effects and deer population dynamics

Abstract Among-individual variation of life history traits in a given population of deer has most often been accounted for by differences among sex and age classes and by current environmental variation (mostly changes in density and climatic conditions). However, among-individual differences in fitness can also be generated by differences in environmental conditions during the year of birth. Such cohort effects can be divided into two different components. First, among-year differences in environmental conditions at birth may produce large yearly variation in recruitment that generates a direct numerical effect (i.e., a high proportion of newborns are recruited in good years, whereas very low proportions are recruited when environmental conditions encountered by newborns during their first weeks of life are poor). Second, when recruited into the population, individuals born in a good year may reproduce earlier, reach a larger body mass, and have a higher reproductive success than individuals born in a poor year. We call the long-lasting influence on individual fitness of environmental conditions during the year of birth a delayed quality effect. Here, we first review briefly evidence of numerical and quality effects recently accumulated in deer populations. Then, by using long-term monitoring (> 20 y) of two contrasted roe deer (Capreolus capreolus) populations, we assess the influence of the among-cohort differences in reproductive traits, age- and sex-specific survival rates, and population growth rate. Results show that cohort effects (1) are widespread in deer populations and (2) may have a major influence on population dynamics, especially in low-performance populations. Cohort effects should therefore be accounted for in management plans of deer populations.

Fitness costs of reproduction depend on life speed: empirical evidence from mammalian populations

Fitness costs of reproduction play a key role in understanding the evolution of reproductive tactics. Nevertheless, the detection and the intensity of costs of reproduction vary according to which life-history traits and species are studied. We propose an evolutionary model demonstrating that the chance of detecting a cost of reproduction should be lower when the fitness component studied has a low rather than high variance. Consequently, the fitness component that is affected the most by costs of reproduction should vary with life speed. Since long-lived species have developed a strategy that avoids jeopardizing their survival and short-lived species favour current reproduction, variance in survival is smaller and variance in reproduction higher in long-lived vs. short-lived species. We review empirical studies of costs of reproduction in free-ranging mammals, comparing evidence of costs reported among species and focal traits. In support of our model, more studies reported evidence of reproductive costs of reproduction in ungulates than in rodents, whereas survival costs of reproduction were more frequent in rodents than in ungulates. The life-history model we propose is expected to apply to any species, and hence provides a better understanding of life-history variation, which should be relevant to all evolutionary ecologists.

Good reindeer mothers live longer and become better in raising offspring.

Longevity is the main factor influencing individual fitness of long-lived, iteroparous species. Theories of life history evolution suggest this is because increased longevity allows individuals to (i) have more breeding attempts (time component), (ii) accumulate experience so as to become better able to rear offspring (experience component) or (iii) because individuals reaching old age have above-average quality (quality component). We assess empirically the relative influences of time, experience and quality on the relationship between longevity and individual fitness among female reindeer. Fitness increased with longevity due to all three processes. All females increased in success with age up to their penultimate year of life (experience component), the success of the terminal-breeding occasion was strongly dependent on longevity. Long-lived females had more successful breeding attempts during their life (time component), and had higher reproductive success at all ages, especially during the last year of life (individual quality component) than short-lived females. Our study reveals a more complex relationship between longevity and fitness in large mammals than the simple increase of the number of reproductive attempts when living longer.

THE FACTORIAL DECOMPOSITION OF THE MAHALANOBIS DISTANCES IN HABITAT SELECTION STUDIES

The Mahalanobis distances have been introduced in habitat selection studies for the estimation of environmental suitability maps (ESMs). The pixels of raster maps of a given area correspond to points in the multidimensional space defined by the mapped environmental variables (ecological space). The Mahalanobis distances measure the distances in this space between these points and the mean of the ecological niche (i.e., the hypothesized optimum for the species) regarding the structure of the niche. The map of these distances over the area of interest is an estimated ESM. Several authors recently noted that the use of a single optimum for the niche of a species may lead to biased predictions of animal occurrence. They proposed to use instead a minimum set of basic habitat requirements, found by partitioning the Mahalanobis distances into a restricted set of biologically meaningful axes. However, the statistical approach they proposed does not take into account the environmental conditions on the area where the niche was sampled (i.e., the environmental availability), and we show that including this availability is necessary. We used their approach as a basis to develop a new exploratory tool, the Mahalanobis distance factor analysis (MADIFA), which performs an additive partitioning of the Mahalanobis distances taking into account this availability. The basic habitat requirements of a species can be derived from the axes of the MADIFA. This method can also be used to compute ESMs using only this small number of basic requirements, therefore including only the biologically relevant information. We also prove that the MADIFA is complementary to the commonly used ecological-niche factor analysis (ENFA). We used the MADIFA method to analyze the niche of the chamois Rupicapra rupicapra in a mountainous area. This method adds to the existing set of tools for the description of the niche.