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.

The efficient and effective monitoring of individuals and populations is critically dependent on correct species identification. While this point may seem obvious, identifying the majority of the more than 100 natural enemies involved in the spruce budworm (Choristoneura fumiferana – SBW) food web remains a non-trivial endeavor. Insect parasitoids play a major role in the processes governing the population dynamics of SBW throughout eastern North America. However, these species are at the leading edge of the taxonomic impediment and integrating standardized identification capacity into existing field programs would provide clear benefits. We asked to what extent DNA barcoding the SBW food web would alter our understanding of the diversity and connectence of the food web and the frequency of generalists vs. specialists in different forest habitats. We DNA barcoded over 10% of the insects collected from the SBW food web in three New Brunswick forest plots from 1983 to 1993. For 30% of these specimens, we amplified at least one additional nuclear region. When the nodes of the food web were estimated based on barcode divergences (using molecular operational taxonomic units (MOTU) or phylogenetic diversity (PD) – the food web became much more diverse and connectence was reduced. We tested one measure of food web structure (the “bird feeder effect”) and found no difference compared to the morphologically based predictions. Many, but not all, of the presumably polyphagous parasitoids now appear to be morphologically-cryptic host-specialists. To our knowledge, this project is the first to barcode a food web in which interactions have already been well-documented and described in space, time and abundance. It is poised to be a system in which field-based methods permit the identification capacity required by forestry scientists. Food web barcoding provided an effective tool for the accurate identification of all species involved in the cascading effects of future budworm outbreaks. Integrating standardized barcodes within food webs may ultimately change the face of community ecology. This will be most poignantly felt in food webs that have not yet been quantified. Here, more accurate and precise connections will be within the grasp of any researcher for the first time.

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.