Marco Festa-Bianchet

Population dynamics of large herbivores: variable recruitment with constant adult survival

Recent long-term studies of population ecology of large herbivorous mammals suggest that survival of prime-aged females varies little from year to year and across populations. Juvenile survival, on the other hand, varies considerably from year to year. The pattern of high and stable adult survival and variable juvenile survival is observed in contrasting environments, independently of the main proximal sources of mortality and regardless of whether mortality is stochastic or density-dependent. High yearly variability in juvenile survival may play a predominant role in population dynamics.

Undesirable evolutionary consequences of trophy hunting.

Phenotype-based selective harvests, including trophy hunting, can have important implications for sustainable wildlife management if they target heritable traits. Here we show that in an evolutionary response to sport hunting of bighorn trophy rams (Ovis canadensis) body weight and horn size have declined significantly over time. We used quantitative genetic analyses, based on a partly genetically reconstructed pedigree from a 30-year study of a wild population in which trophy hunting targeted rams with rapidly growing horns, to explore the evolutionary response to hunter selection on ram weight and horn size. Both traits were highly heritable, and trophy-harvested rams were of significantly higher genetic ‘breeding value’ for weight and horn size than rams that were not harvested. Rams of high breeding value were also shot at an early age, and thus did not achieve high reproductive success. Declines in mean breeding values for weight and horn size therefore occurred in response to unrestricted trophy hunting, resulting in the production of smaller-horned, lighter rams, and fewer trophies.

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.

Mass- and Density-Dependent Reproductive Success and Reproductive Costs in a Capital Breeder

For capital breeders, mass may affect reproductive potential. Reproductive expenditure may reduce future reproductive potential, particularly when resources are scarce. To test the hypothesis that reproductive success and the costs of reproduction vary according to mass and population density, we analyzed 25 yr of data on bighorn ewes (Ovis canadensis). The number of adult females was first limited by yearly removals, then allowed to triple. We found no survival costs of reproduction for ewes aged 4–7 yr. For ewes aged 8–14 yr, survival was density dependent for barren ewes but not for ewes that weaned lambs. Failure to lamb was rare and negatively correlated with fertility the following year. At low population density, lactation had a negative effect on mass gain but had a limited reproductive cost. At high density, heavy ewes had higher reproductive success than light ewes, and the reproductive cost and somatic costs of reproduction increased. The cost of reproduction was greater for light than for heavy ewes. Survival of weaned lambs to 1 yr was affected by population density but not by maternal mass or previous reproductive success. In large mammals, manipulations of reproductive effort are problematic, but long‐term monitoring of individual mass and reproductive success under varying conditions of resource availability can provide insights into the evolution of life histories.

Seasonal range selection in bighorn sheep: conflicts between forage quality, forage quantity, and predator avoidance

SummaryThe migratory and foraging behavior of individually marked bighorn ewes (Ovis canadensis) was studied to test the hypothesis that forage quality determined seasonal range selection. Forage quality was monitored through analysis of fecal crude protein. Ewes in the study population utilized two distinct ranges differing in elevation and possibly predation risk. Pregnant ewes migrated in May from the low-elevation winter range to lambing areas at higher elevation, before plant growth had started there. In so doing, they moved from a range of high-quality forage to one of low-quality forage, apparently to avoid predation on newborn lambs. Non-pregnant adult ewes migrated later. Most yearling ewes (which are not pregnant) migrated with the adult ewes to the lambing areas, but returned to the winter range within a few days, then migrated again to high-elevation areas in June. Forage quality was higher at high elevation from mid-June at least through July, but forage availability appeared to be lower than in the winter range. Seasonal range selection is likely determined by a combination of nutritional and antipredator constraints. The antipredator strategy of bighorn ewes does not always allow them to utilize the range with the best forage.

Predator-induced natural selection on temperament in bighorn ewes

Temperament traits in animals may have important fitness consequences, but have received little attention from ecologists or evolutionary biologists. A few studies have linked variation in temperament with fitness, but none has measured selection on temperament traits. We estimated the strength of selection on female boldness and docility on bighorn sheep ewes, Ovis canadensis. The Ram Mountain population experienced little predation pressure during the first 25 years of study, then 2 years (1997 and 1998) of frequent predation by cougars, Puma concolor, during which adult ewe mortality almost tripled over the long-term average, to 27% a year. During years of high predation, we found moderate selection favouring bold ewes, and age-specific selection on docility. Old ewes appeared more vulnerable to predation than young ewes. In contrast, no evidence of selection on temperament traits was observed during 2 years of low predation (1996 and 1999). These results suggest predator-induced selection favouring bold and nondocile ewes. Leadership was highly correlated with age and may increase the risk of predator encounter. Leadership alone, however, could not explain the higher vulnerability of old ewes to predation. Cougar predation on bighorn sheep occurs sporadically and unpredictably, probably because individual cougars often are prey specialists. Cougar predation may have limited microevolutionary effects on temperament in bighorn sheep, because it mostly affects ewes near the end of their reproductive life span and because of potential countervailing selection on boldness and docility. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.

Age-dependent sexual selection in bighorn rams

Although mating systems and sexual selection have been intensively studied in ungulate model systems, very few studies have combined genetic paternity analysis with individual phenotypic data over several breeding seasons. We used microsatellite paternity analysis to determine the parentage of 83 bighorn sheep (Ovis canadensis) born between 1995 and 2000 at Ram Mountain, Alberta, Canada. We could assign the paternity of 64 lambs at a high level of statistical confidence (95%). Within each season, the most successful ram sired an average of 35.5% of the lambs with assigned paternity, and a single ram sired 26.1% of all lambs over the six mating seasons. Although a few large-horned, mature (age 8+ years) rams had very high reproductive success, younger rams sired ca. 50% of the lambs. Mixed–effects models indicated that mating success increases as a nonlinear function of age, with horn length increasingly positive in correlation with mating success in older rams. These results indicate that young or small rams possibly achieve mating success through alternative mating tactics that are less dependent on body and weapon size, such as coursing and blocking. Sexual selection is therefore likely to have age–dependent effects on traits such as agility, body and horn size.

Senescence rates are determined by ranking on the fast-slow life-history continuum

Comparative analyses of survival senescence by using life tables have identified generalizations including the observation that mammals senesce faster than similar-sized birds. These generalizations have been challenged because of limitations of life-table approaches and the growing appreciation that senescence is more than an increasing probability of death. Without using life tables, we examine senescence rates in annual individual fitness using 20 individual-based data sets of terrestrial vertebrates with contrasting life histories and body size. We find that senescence is widespread in the wild and equally likely to occur in survival and reproduction. Additionally, mammals senesce faster than birds because they have a faster life history for a given body size. By allowing us to disentangle the effects of two major fitness components our methods allow an assessment of the robustness of the prevalent life-table approach. Focusing on one aspect of life history - survival or recruitment - can provide reliable information on overall senescence.

EFFECTS OF AGE, SEX, DISEASE, AND DENSITY ON SURVIVAL OF BIGHORN SHEEP

Longitudinal studies of survival are valuable because age-specific survival affects population dynamics and the evolution of several life history traits. We used capture–mark–recapture models to assess the relationship between survival and sex, age, population, year of study, disease, winter weather, and population density in two populations of bighorn sheep (Ovis canadensis) in Alberta, Canada. The Ram Mountain population, monitored for 20 yr, more than doubled in density; the Sheep River population, monitored for 13 yr, experienced a pneumonia epizootic. Yearling survival varied among years and was lower than that of older sheep of the same sex, except for yearling males at Ram Mountain. Yearling females at Ram Mountain were the only sex-age class exhibiting density dependence in survival. Senescence was evident for both sexes in both populations. Female survival from age 2 to age 7 was very high in both populations, but males aged 2 and 3 yr enjoyed better survival than males aged 4–6 yr. Our data support the suggestion that where hunters remove many males older than 5 yr of age, the natural mortality of males increases at 3–5 yr, possibly because young males suffer a mortality cost of participating in rutting activity. The decline in survival for sheep older than 7 yr was greater for males than for females. Survival was lower for males than for females, both among prime-aged sheep (0.896 vs. 0.939 at Sheep River; 0.837 vs. 0.945 at Ram Mountain) and among older sheep (0.777 vs. 0.859 at Sheep River; 0.624 vs. 0.850 at Ram Mountain), but not among yearlings. Survival of sheep aged 2–7 yr was not significantly different between the two populations. Winter weather did not affect survival. Survival of sheep 2 yr of age and older did not vary significantly between years, except at Sheep River where survival of prime-aged sheep of both sexes was lower in the year of the pneumonia epizootic. Studies of survival of mountain sheep based upon skull collections may have overestimated survival of young rams. Our results underline the need for accurate information on age-specific survival.

INDIVIDUAL DIFFERENCES, LONGEVITY, AND REPRODUCTIVE SENESCENCE IN BIGHORN EWES

Within individual iteroparous mammals, a high rate of reproduction in early life may occur at the cost of decreased reproduction near the end of life, leading to reproductive senescence. Using long-term data on marked individuals from two populations of bighorn sheep (Ovis canadensis), we tested for the existence of reproductive senescence and of trade-offs between longevity and early reproductive success in ewes, which have an observed maximum life-span of 19 yr. Lamb production decreased in older ewes, while weaning success for parous ewes was independent of age in one population and decreased with age in the other. The age-related decrease in lamb production followed a slight decrease in body mass, which began at ∼11 yr of age. Reproductive senescence in this species appears to begin 6–7 yr after the onset of survival senescence. Longevity was positively related to late-summer body mass at 6 yr. Contrary to reproductive cost theories, there was no negative relationship between early and late reproducti...