Comparing fitness measures and the influence of age of first reproduction in Columbian ground squirrels

The timing of life-history traits may have strong influences on the evolution of life cycles and on population demography. This is especially true of the age at which females first reproduce (Cole’s principle). We examined whether the age at which females first reproduce influences fitness in Columbian ground squirrels (Urocitellus columbianus), for which females varied in the age at which they initially produce weaned offspring, from ages 1 through 5 years. With 148 females with complete known life spans in a 28-year data set, we examined four fitness measures: individual fitness (λ ind), individual fitness relative to the pattern of growth of the population (λ rel), lifetime reproductive success (LRS), and LRS relative to the total LRS for each female’s cohort (LRSrel). These metrics were calculated for offspring produced at the time of weaning and offspring that survived to emerge after their first hibernation period. Individual fitness (λ ind) was significantly associated with population growth during a female’s lifetime (λ Leslie; R2 = 0.523, P < 0.0001), indicating the need to adjust individual fitness for demonstrated changes in population growth and thus producing a relative individual fitness index (λ rel). We regressed λ rel on age at first reproduction, and found significant selection favoring earlier reproductive success (β ± SE = −0.20 ± 0.06; R2 = 0.306, P < 0.0001). When using an earlier (offspring at weaning) versus later (those that survived their first hibernation) measure of fecundity, we found that the latter introduced considerable variation, likely environmental, into the estimate of selection. This greatly weakened the regression of relative fitness on the age at first successful reproduction. LRS and LRSrel exhibited nonsignificant changes with age at first reproduction. Finally, those females that reproduced successfully at younger ages had similar litter sizes but significantly shorter life spans than females that matured when older, perhaps reflecting costs to early reproduction.

Ectoparasites and fitness of female Columbian ground squirrels

Parasites play an important role in the evolution of host traits via natural selection, coevolution and sexually selected ornaments used in mate choice. These evolutionary scenarios assume fitness costs for hosts. To test this assumption, we conducted an ectoparasite removal experiment in free-living Columbian ground squirrels ( Urocittelus columbianus ) in four populations over three years. Adult females were randomly chosen to be either experimentally treated with anti-parasite treatments (spot-on solution and flea powder, N = 61) or a sham treatment (control, N = 44). We expected that experimental females would show better body condition, increased reproductive success and enhanced survival. Contrary to our expectations, body mass was not significantly different between treatments at mating, birth of litter or weaning of young. Further, neither number nor size of young at weaning differed significantly between the two treatments. Survival to the next spring for adult females and juveniles was not significantly different between experimental and control treatments. Finally, annual fitness was not affected by the treatments. We concluded that females and their offspring were able compensate for the presence of ectoparasites, suggesting little or no fitness costs of parasites for females in the different colonies and during the years of our experiments.