A nonlinear demographic model was used to predict the population dynamics of the flour beetle Tribolium under laboratory conditions and to establish the experimental protocol that would reveal chaotic behavior. With the adult mortality rate experimentally set high, the dynamics of animal abundance changed from equilibrium to quasiperiodic cycles to chaos as adult-stage recruitment rates were experimentally manipulated. These transitions in dynamics corresponded to those predicted by the mathematical model. Phase-space graphs of the data together with the deterministic model attractors provide convincing evidence of transitions to chaos.

Our approach to testing nonlinear population theory is to connect rigorously mathematical models with data by means of statistical methods for nonlinear time series. We begin by deriving a biologically based demographic model. The mathematical analysis identifies boundaries in parameter space where stable equilibria bifurcate to periodic 2—cycles and aperiodic motion on invariant loops. The statistical analysis, based on a stochastic version of the demographic model, provides procedures for parameter estimation, hypothesis testing, and model evaluation. Experiments using the flour beetle Tribolium yield the time series data. A three—dimensional map of larval, pupal, and adult numbers forecasts four possible population behaviors: extinction, equilibria, periodicities, and aperiodic motion including chaos. This study documents the nonlinear prediction of periodic 2—cycles in laboratory cultures of Tribolium and represents a new interdisciplinary approach to understanding nonlinear ecological dynamics.

A defining hypothesis of theoretical ecology during the past century has been that population fluctuations might largely be explained by relatively low-dimensional, nonlinear ecological interactions, provided such interactions could be correctly identified and modeled. The realization in recent decades that such nonlinear interactions might result in chaos and other exotic dynamic behaviors has been exciting but tantalizing, in that attributing the fluctuations of a particular real population to the complex dynamics of a particular mathematical model has proved to be an elusive goal. We experimentally tested a modelpredicted sequence of transitions (bifurcations) in the dynamic behavior of a population from stable equilibria to quasiperiodic and periodic cycles to chaos to three-cycles using cultures of the flour beetle Tribolium. The predictions arose from a system of difference equations (the LPA model) describing the nonlinear life-stage interactions, predominantly cannibalism. We built a stochastic version of the model incorporating demographic variability and obtained conditional least-squares estimates for the model parameters. We generated 2000 ''bootstrapped data sets'' with a time-series bootstrap technique, and for each set we reestimated the model parameters. The resulting 2000 bootstrapped parameter vectors were used to obtain confidence intervals for the model parameters and estimated distributions of the Liapunov exponents for the deterministic portion (the skeleton) of the model as well as for the full stochastic model. Frequency distributions of estimated dynamic behaviors of the skeleton at each experimental treatment were produced. For one treatment, over 83% of the bootstrapped parameter estimates corresponded to chaotic attractors, and the remainder of the estimates yielded high-period cycles. The low-dimensional skeleton accounted for at least 90% of the variability in the population abundances and accurately described the responses of populations to experimental demographic manipulations, including treatments for which the predicted dynamic behavior was chaos. Demographic stochasticity described the remaining noise quite well. We conclude that the fluctuations of experimental flour beetle populations are explained largely by known nonlinear forces involving cannibalistic-stage interactions. Claims of dynamic behavior such as periodic cycles or chaos must be accompanied by a consideration of the reliability of the estimated parameters and a realization that the population fluctuations are a blend of deterministic forces and stochastic events.

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