Food and space: resources defended by territorial parous female Columbian ground squirrels

The importance of food and space, as resources defended by parous female Columbian ground squirrels (Spermophilus columbianus), was studied by manipulating one of these resources, that is increasing the quality of forage (through urine fertilization) on small plots within territories. Use of these fertilized plots by ground squirrels increased more than 100-fold when averaged over 2 years after manipulation. This increase, however, was not equal for each sex and age class: parous females used the fertilized plots relatively more, and nonparous females less, than either yearling or adult males. The number of parous females with territories overlapping the experimental plots also increased after fertilization, but the size of their territories declined only slightly, by less than 10%. Parous females with access to the fertilized plots, relative to those without such access, had greater body mass and larger litters that both weighed more at birth and gained body mass subsequently more rapidly. Parous females on territories with fertilized plots showed higher levels of agonism than those on territories lacking such plots. Most of the agonism was centered on the experimental plots and more of it was directed at young of other females than at their own young. Such differential treatment of kin, however, did not extend to their offspring of the previous year. We suggest that for parous females of this ground squirrel, both food and space (at least that normally needed to supply sufficient forage) are important resources to defend, and both may have played a significant role in the evolution of territoriality in females of this species.