Trophic cascades have been a central paradigm in explaining the structure of ecological communities but have been demonstrated mainly through comparative studies or experimental manipulations. In contrast, evidence for shifts in trophic cascades caused by intrinsically driven population dynamics is meager. By using empirical data of a cannibalistic fish population covering a 10-year period and a size-structured population model, we show the occurrence of a dynamic trophic cascade in a lake ecosystem, in which the community over time alternates between two different configurations. The intrinsically driven change in the size structure of the fish population from a dominance of stunted individuals to a dominance of gigantic cannibals among adult individuals is the driving force behind distinct abundance switches observed in zooplankton and phytoplankton. The presence of the phase with gigantic cannibals depends critically on the energy they extract from their victims, allowing strong reproduction for a number of years. C ommunity-wide trophic cascades, the propagation of indirect mutualism between nonadjacent trophic levels in food webs, have been suggested to occur more frequently in aquatic than in terrestrial systems (1-4). This suggestion is based on the arguments that terrestrial systems have a higher heterogeneity, a higher overall species diversity, and more chemical defenses among primary producers (higher plants vs. algae; ref. 1). Although the validity of all of these arguments has been questioned (2, 4), undoubtedly the empirical evidence for community-wide trophic cascades is, at present, substantially stronger for aquatic than for terrestrial systems. The empirical evidence largely stems from two sources: comparative studies of different systems in which the trophic structure, such as food chain length, differs (5-6) and experimental manipulations of top predators, either intentional or unintentional (species invasions; refs. 2, 3, and 7-9). In contrast, there is hardly any evidence for dynamic trophic cascades, in which major shifts in overall food-web structure are intrinsically driven by population dynamics. Only a few studies on recruitment variation have considered this aspect (10-12).Cannibalism has been shown to have a number of diverse effects on population dynamics and persistence (13-17). These effects, among others, include a potential for alternative stable states (18,19) and chaotic dynamics (20). Although many cannibalistic models ignore the energy that cannibals gain from cannibalism and, thus, are essentially ''infanticide'' models (13,15,20), some theoretical studies have shown that the energy extracted by the cannibal may have substantial impact on population persistence and individual life history (14, 16). Empirical evidence also suggests such an effect of energy extraction on population dynamics because of increased growth and thereby increased per-capita fecundity of cannibals (17).Here we present strong evidence for a whole-lake trophic cascade that is dynamic and intrinsically ...

Abstract. Recent size-structured cannibalistic models point to the importance of the energy gain by cannibals and also show that this gain may result in the emergence of giant individuals. We use a combination of a 10-year field study of a perch (Perca fluviatilis) population and quantitative within-season modeling of individual and population-level dynamics to investigate which mechanisms are most likely to drive the dynamics of the studied perch population. We focused on three main aspects to explain observed discrepancies between earlier model predictions and data: (1) introduction of more than one shared resource between cannibals and victims, (2) whether or not several victim age cohorts are necessary to allow giant growth, and (3) the intensity of inter-cohort competition between young-of-the-year (YOY) perch and 1-yr-old perch.At the start of the study period, the perch population was dominated by ''stunted'' perch individuals, and recruitment of perch to an age of 1-yr-old was negligible. Following a major death in adult perch, strong recruitments of perch to 1-yr-old were thereafter observed for a number of years. As 1-yr-olds these successful recruiters subsequently starved to death due to competition with the new YOY. The few surviving adult perch accelerated substantially in growth and became ''giants.'' At the end of the study period, the perch population moved back to the situation with stunted individuals. There was a high agreement between observed diets of cannibalistic perch and those predicted by the model for both the stunted and the giant phases. Analyses of growth rates showed that cannibalistic perch could become giants on a diet of YOY perch only, but that a supplement with the second shared resource (macroinvertebrates) was needed to reach the observed sizes. Modeling of growth and diet in the giant phase showed an exploitative competitive effect of YOY perch on 1-yr-old perch, but a restriction in habitat use of 1-yr-old perch had to be assumed to yield the observed growth rate and diet. The resource dynamics of zooplankton and macroinvertebrates were both accurately predicted by the model. Also, YOY perch mortality was accurately predicted and, furthermore, suggested that one of the trawling methods used may underestimate the number of YOY perch when they increase in size.We conclude that the presence of a second shared resource and the restricted habitat use and absence of cannibalistic consumption by 1-yr-old perch individuals are two important mechanisms to explain the discrepancy between model predictions and data. Our results also point to the fact that that the dynamics observed may be explained by complex dynamics not involving the presence of a giant and dwarf cycle.

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