Jon T. Jorgenson

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.

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.

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...

Body mass and individual fitness in female ungulates: bigger is not always better

In female vertebrates, differences in fitness often correspond to differences in phenotypic quality, suggesting that larger females have greater fitness. Variation in individual fitness can result from variation in life span and/or variation in yearly reproductive success, but no study has yet assessed the relationships between the components of fitness and phenotypic quality while controlling for life span. We tried to fill this gap using data from long–term monitoring (23 years) of marked roe deer and bighorn sheep, two ungulates with very different life histories. In both species, we found a strong positive relationship between an adult females mass and her probability of reaching old age: over the long term, bigger is indeed better for ungulate females. On the other hand, we found no evidence in either species that heavier females had higher fitness when differences in life span were accounted for: over the short term, bigger is not necessarily better. Our results indicate that, while broad differences in phenotypic quality affect individual fitness, when differences in life span are accounted for phenotypic quality has no residual effect on fitness. Therefore, within a given range of phenotypic quality, bigger is not always better, for reasons which may differ between species.

Life History Consequences of Variation in Age of Primiparity in Bighorn Ewes

We investigated the life history consequences of age at primiparity in two marked populations of bighorn sheep (Ovis canadensis), studied for 19 and 12 yr. Some ewes first lambed at 2 yr of age, others at 3 or 4 yr. Lambs of 2-yr-old ewes were smaller and less viable than lambs of older ewes. Lactation at 2 yr did not affect survival to 3 yr, but in one population it was associated with reduced survival from 3 to 4 yr. Two yr olds that weaned lambs gained less mass during summer than other 2 yr olds. Overwinter mass loss from 2 to 3 yr was correlated with autumn body mass, and was lower for parous than for nonporous ewes. Reproductive success at 3 yr was independent of reproductive status at 2 yr. At 4 yr, ewes that lactated as 2 yr olds were smaller than ewes that had not lactated as 2 yr olds. In one population, negative effects of early maturation upon mass gain and subsequent reproductive success became evident at high density, and very few 2 yr olds lambed when population density was high. The lifetime reproductive success of early- maturing ewes was not lower than that of late-maturing ewes. To account for individual variation in initial reproductive potential, we controlled body mass as a yearling, a variable that affected reproductive success. Mass as a yearling, however, did not interact with age of primiparity to affect survival, growth, or reproductive success. Uncertainties about re- source availability and possible effects of genotype and body fat may explain why many ewes postponed their first reproduction despite apparently low life history costs of early maturation.

Heritability of body mass varies with age and season in wild bighorn sheep

Heritabilities (h2) of body mass at different ages and seasons were estimated using offspring–mother regression and restricted maximum likelihood (REML) methods for bighorn sheep on Ram Mountain, Alberta. Both methods resulted in similar estimates of h2 for adults, but for lambs and yearlings heritability was underestimated by offspring–mother regression relative to REML, possibly because of higher maternal-effects bias for offspring–mother regression. Heritabilities of body mass in bighorn were similar to published estimates for domestic sheep. Heritability estimated by offspring–mother regression increased after 2 years of age. The REML method suggested that heritability was moderate for lambs and yearlings, very low at 2 years of age, and increased afterwards. The increase in heritability with age was attributed to declining negative maternal effects. Very low h2 estimates at 2 years of age, obtained with both methods, appeared to be caused by a combination of high environmental variance and very low genetic variance. Body mass of bighorn sheep has a pronounced seasonal cycle, and h2 was lower in June than in September for 2-year-olds and older sheep, and associated with both lower VA and higher VE in spring.

Harvesting bighorn ewes: consequences for population size and trophy ram production

We wanted to test whether ewe hunting would cause a decline in population size or in trophy ram production, and whether a reduction in ewe density would increase the size of ram horns. Thus, we examined the consequences of a bighorn sheep (Ovis canadensis) ewe hunting season through an experimental manipulation of an isolated population in Alberta, 1972-91. The number of ewes remained stable during 9 years despite yearly removals of 12-24% of the total ewe population. The removals did not affect (P>0.5) ewe mortality due to other causes, lamb production by adult ewes, or lamb survival