Patterns in food-web structure have frequently been examined in static food webs, but few studies have attempted to delineate patterns that materialize in food webs under nonequilibrium conditions. Here, using one of nature's classical nonequilibrium systems as the food-web database, we test the major assumptions of recent advances in food-web theory. We show that a complex web of interactions between insect herbivores and their natural enemies displays significant architectural flexibility over a large fluctuation in the natural abundance of the major herbivore, the spruce budworm ( Choristoneura fumiferana ). Importantly, this flexibility operates precisely in the manner predicted by recent foraging-based food-web theories: higher-order mobile generalists respond rapidly in time and space by converging on areas of increasing prey abundance. This “birdfeeder effect” operates such that increasing budworm densities correspond to a cascade of increasing diversity and food-web complexity. Thus, by integrating foraging theory with food-web ecology and analyzing a long-term, natural data set coupled with manipulative field experiments, we are able to show that food-web structure varies in a predictable manner. Furthermore, both recent food-web theory and longstanding foraging theory suggest that this very same food-web flexibility ought to be a potent stabilizing mechanism. Interestingly, we find that this food-web flexibility tends to be greater in heterogeneous than in homogeneous forest plots. Because our results provide a plausible mechanism for boreal forest effects on populations of forest insect pests, they have implications for forest and pest management practices.

We conducted a 14‐yr intensive study of spruce budworm (Choristoneura fumiferana (Clem.)) survivorship at three study plots in largely balsam fir (Abies balsamea (L.) Mill.) stands in New Brunswick, Canada, to elucidate certain key mechanisms underlying spruce budworm outbreak cycles. The study covered a peak‐to‐declining phase (from 1981 and 1994) of the budworm outbreak cycle that had started in the early 1960s. Frequent sampling was carried out in each plot‐year to construct a practically continuous survivorship curve, and the annual variation in population density was estimated. We found a high level of correlation between the studied phase of the outbreak cycle and annual variations in the survivorship over the postdiapause period, suggesting that postdiapause survivorship was the chief determinant of the cycle. We found the annual changes in population density in the present study to be closely similar in pattern to those from the provincial budworm surveys conducted in much larger areas. This implies that the mechanism underlying the population process found in the few study plots in largely balsam fir stands also applies to the process in much larger areas of diverse stand types. The main source of postdiapause mortality is found to be natural enemies. The impacts of parasitoids and disease are evaluated by rearing budworm samples in the laboratory. Hymenopteran and dipteran parasitoids are by far the major sources of mortality, and microsporidians are the most prevalent pathogen. Occurrences of other entomopathogenic fungi and viruses were insignificant throughout the study. Seasonal changes in laboratory survivorship are compared with the corresponding field survivorship to estimate the effect of predation. No major mortality factor is found to singly play a predominant role in determining the outbreak cycle. Conversely, some minor factors are shown to have played significant roles. Thus, the importance of recognizing the action of natural enemies as a complex is emphasized for understanding the budworm outbreak cycle. Finally, centered on the roles played by the chronological succession of natural enemies in the present study, the results of budworm research in New Brunswick since the mid‐1940s are synthesized to outline basic mechanisms underlying the outbreak processes as a guide for further studies.

A grid of 25 traps baited with virgin female moths was used to monitor the abundance of male spruce budworm in relatively sparse populations for 12 successive years. Concurrent sampling of third-instar larvae showed that the male moth counts of one generation had a predictive relationship to third-instar larval counts of the next generation and therefore the sex-attractant trap is a feasible means of measuring budworm abundance at low population densities. But three sampling constraints must be noted: (a) a moth invasion will generally invalidate the relationship;(b)extrapolation of a moth–larvae relationship observed in one forest biotype to biotypes with different host characteristics is risky; (c) the probability of a female attracting a particular male decreases sharply with increasing population density.Trapping success is discussed with respect to location and height within the stand and to weather conditions.