The phenomenology of snowshoe hare cycles is addressed via
construction of a three-trophic-level population dynamics model in
which hare populations are limited by the availability of winter
browse from below and by predation from above. In the absence of
predators, the
model predicts annual oscillations, the magnitude of which depends
on habitat quality. With predators in the system, a wide range of
additional dynamics are possible: multi-annual cycles of various
periods, quasiperiodicity, and chaos. Parameterizing the model from
the literature leads to the conclusion that the model is compatible
with the principal features of the cycle in nature: its regularity,
mean period, and the observed range of peak-to-trough amplitudes.
The model also points to circumstances that can result in the
cycle's abolition as observed, for example, at the southern edge of
the hare's range. The model predicts that the increase phase of the
cycle is brought to a halt by food limitation, while the decline
from peak numbers is a consequence of predation. This is consistent
with factorial field experiments in which hare populations were
given supplemental food and partial surcease from predators. The
results of the experiments themselves are also reproducible by the
model.
Analysis of the model was carried out using a recently developed
methodology in which the original dynamical system is reformulated
as a perturbation of a Hamiltonian limit wherein exist infinite
numbers of periodic, quasiperiodic, and chaotic motions. The
periodic orbits are continued numerically into non-Hamiltonian
regions of parameter space corresponding to the situation in
nature. This procedure allows one to obtain an overall
understanding of the geometry of parametric dependencies. The
present study represents the first formulation of a full
three-trophic-level snowshoe hare model and the first time
any model of the cycle has been parameterized entirely using
independently-measured quantities.