PATTERNS OF PREDATION ON NONCYCLIC LEMMINGS

Noncyclic populations of microtine rodents may be limited within a relatively constant range of densities by generalist predators with a prey base sufficiently diverse to sustain them when rodents are scarce (generalist predator hypothesis). Collared lemmings (Dicrostonyx kilangmiutak) at Pearce Point, Northwest Territories, Canada, are noncyclic and limited to fairly constant low densities in summer by predation, principally by red fox and Rough-legged Hawks. We tested four predictions of the generalist predator hypothesis as a possible explanation for relatively constant lemming densities: (1) predators do not show strong numerical responses to lemming density; (2) the proportion of lemming biomass in predator diets declines with declining lemming abundance, compensated for by increased consumption of alternative prey; (3) predators show a type-III functional response to lem- ming density; and (4) at low densities, predation on lemmings ceases. The first prediction was not satisfied by the principal predators: at very low lemming densities, Rough-legged Hawks did not settle, and breeding success of red foxes and hawks was limited by lemming abundance. However, a number of generalist predators (Golden Eagle, grizzly bear, arctic ground squirrel, Peregrine Falcon, and Gyrfalcon) did not respond numerically to the lowest lemming densities. The second prediction was partly supported: all predators consumed lemmings at a lower rate as lemming densities declined. However, Rough-legged Hawks were not able to compensate fully for the declining consumption by increasing their use of alternative prey, and red foxes were able to do so in one of three years. Regarding the third prediction, foxes showed some evidence of a type-III functional response but hawks did not. As for the fourth prediction, most predators still consumed lemmings at very low densities; lemmings lacked a secure refuge. The Pearce Point system differs from those where microtine dynamics are relatively constant and nonirruptive because of persistent predation by generalists. In terms of biomass, lemmings are the principal prey for their dominant predators. These predators (the semi- generalist red fox and the specialist Rough-legged Hawk) rely on lemmings to breed, but drive them to densities too low to sustain breeding by these same predators in the subsequent spring. In this regard, the system is similar to one driven by specialists. In some winters, however, populations recover because lemmings breed under the snow and most summer predators are absent. As a result, lemming densities in spring are often high enough for specialists and semigeneralists to initiate breeding. When winter breeding and survival fail to allow population growth, hawks and foxes may fail to breed and then leave the system. Even so, summer generalists still persist and continue to consume lemmings, curtailing potential irruptive growth. In this regard, the system is similar to one where prey are relatively constant because of generalists. Community dynamics at Pearce Point can best be understood as a combination of three dominant processes. Summer predation by specialists and semigeneralists results in desta- bilizing declines. Winter breeding, coupled with good survival, can lead to destabilizing growth. However, this growth is curtailed in the following summer by either destabilizing specialist predation or the stabilizing influence of generalist predation. When lemmings are scarce, the semigeneralist red fox and some generalist predators rely on arctic ground squirrels as their primary prey or their principal alternative prey. The ground squirrel appears to be the critical species maintaining this relatively diverse arctic tundra predator community and the relatively constant lemming densities.