Like most heavily preyed-upon animals, snowshoe hares (Lepus americanus) have to balance conflicting demands of obtaining food at a high rate and avoiding predators. Adopting foraging behaviours to minimise predation risk may also lead to a decline in condition, and hence fecundity. Predictions of three hypotheses (condition constraint hypothesis, predator-avoidance constraint hypothesis, predation-sensitive foraging (PSF) hypothesis) were tested by comparing changes in the survival and condition of snowshoe hares on four experimental areas in winter during a cyclic peak and decline (1989)(1990)(1991)(1992)(1993) near Kluane Lake, Yukon, Canada, where (i) predation risk was reduced by excluding terrestrial predators (FENCE), (ii) food supply was supplemented with rabbit chow a d libitum (FOOD), (iii) these two treatments were combined (FENCE+FOOD), and (iv) an unmanipulated CONTROL was used.Different pattems of survival and changes in body mass were observed in the presence and absence of terrestrial predators. On the CONTROL area, female body mass and fecundity declined, even though sufficient winter forage was apparently available in all years. A similar decrease in body mass was observed on the FOOD treatment, but only during the third year of the population decline. In contrast, female body mass remained high throughout the decline in the absence of terrestrial predators in the FENCE+FOOD and FENCE treatments. Winter survival declined on CONTROL and FENCE areas during the first year of the population decline (1991), but remained higher on FOOD until 1992 and FENCE+FOOD until 1993. These results generally supported the PSF hypothesis where terrestrial predators were present (CONTROL and FOOD grids). Where terrestrial predators were absent (FENCE and FENCE+FOOD), the results supported the alternative condition constraint hypothesis. The evidence suggests that a cascade of sublethal behavioural and physiological effects associated with increased predation risk contribute to the population decline and delayed recovery of cyclic low-phase populations of snowshoe hares.
IntroductionWhere predation is implicated in the regulation of prey populations, anti-predator behaviour will probably play a role in the process (Abrams 1989(Abrams , 1990(Abrams , 1992a(Abrams , 1992b(Abrams , 1992c(Abrams , 1993Matsuda and Abrams 1994). The theoretical basis for exploring this problem of a trade-off between gaining energy and avoiding predation has developed rapidly in recent years (Gilliam and Fraser 1987; Houston 1987, 1990;Ludwig and Rowe 1990;Brown 1992;Houston et al. 1993;Clark 1993Clark , 1994 Oksanen and Lundberg 1994). Numerous studies have investigated the effects of increased predation risk on foraging behaviour of small mammals (Holmes 1984;Kotler 1984;Anderson 1986; Desy et al. 1990;Lima and Dill 1990;Cassini and Galante 1992;Dickman 1992;Kotler et al. 1992;Saarikko 1992;Hughes et al. 1994), and in most of these studies increased risk of predation resulted in a shift in patterns of habitat use, diet and tim...