The population dynamics of the larch budmoth (LBM), Zeiraphera diniana, in the Swiss Alps are perhaps the best example of periodic oscillations in ecology (figure 7.1). These oscillations are characterized by a remarkably regular periodicity, and by an enormous range of densities experienced during a typical cycle (about 100,000-fold difference between peak and trough numbers). Furthermore, nonlinear time series analysis of LBM data (e.g., Turchin 1990, Turchin and Taylor 1992) indicates that LBM oscillations are definitely generated by a second-order dynamical process (in other words, there is a strong delayed density dependence—see also chapter 1). Analysis of time series data on LBM dynamics from five valleys in the Alps suggests that around 90% of variance in Rt is explained by the phenomenological time series model employing lagged LBM densities, R, =f(Ni-1,Ni-2,) (Turchin 2002). As discussed in the influential review by Baltensweiler and Fischlin (1988) about a decade ago, ecological theory suggests a number of candidate mechanisms that can produce the type of dynamics observed in the LBM (see also chapter 1). Baltensweiler and Fischlin concluded that changes in food quality induced by previous budmoth feeding was the most plausible explanation for the population cycles. During the last decade, the issue of larch budmoth oscillations was periodically revisited by various population ecologists looking for general insights about insect population cycles (e.g., Royama 1977, Bowers et al. 1993, Ginzburg and Taneyhill 1994, Den Boer and Reddingius 1996, Hunter and Dwyer 1998, Berryman 1999). These authors generally concurred with the view that budmoth cycles are driven by the interaction with food quality. A recent reanalysis of the rich data set on budmoth population ecology collected by Swiss researchers over a period of several decades, however, suggested that the role of parasitism is underappreciated (Turchin et al. 2002). Before focusing on the roles of food quality and parasitism in LBM dynamics, we briefly review the status of other hypotheses that were discussed in the literature on LBM cycles. First, the natural history of the LBM-larch system is such that food quantity is an unlikely factor to explain LBM oscillations.
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