Chaos theory discloses triggers and drivers of plankton dynamics in stable environment.

Despite the enticing discoveries of chaos in nature, triggers and drivers of this phenomenon remain a classical enigma which needs irrefutable empirical evidence. Here we analyze results of the yearlong replicated mesocosm experiment with multi-species plankton community that allowed revealing signs of chaos at different trophic levels in strictly controlled abiotic environment. In mesocosms without external stressors, we observed the "paradox of chaos" when biotic interactions (internal drivers) were acting as generators of internal abiotic triggers of complex plankton dynamics. Chaos was registered as episodes that vanished unpredictably or were substituted by complex behaviour of other candidates when longer time series were considered. Remarkably, episodes of chaos were detected even in the most abiotically stable conditions. We developed the Integral Chaos Indicator to validate the results of the Lyapunov exponent analysis. These findings are essential for modelling and forecasting behaviour of a variety of natural and other global systems.

Coupled predator-prey oscillations in a chaotic food web.

Coupling of several predator-prey oscillations can generate intriguing patterns of synchronization and chaos. Theory predicts that prey species will fluctuate in phase if predator-prey cycles are coupled through generalist predators, whereas they will fluctuate in anti-phase if predator-prey cycles are coupled through competition between prey species. Here, we investigate predator-prey oscillations in a long-term experiment with a marine plankton community. Wavelet analysis of the species fluctuations reveals two predator-prey cycles that fluctuate largely in anti-phase. The phase angles point at strong competition between the phytoplankton species, but relatively little prey overlap among the zooplankton species. This food web architecture is consistent with the size structure of the plankton community, and generates highly dynamic food webs. Continued alternations in species dominance enable coexistence of the prey species through a non-equilibrium 'killing-the-winner' mechanism, as the system shifts back and forth between the two predator-prey cycles in a chaotic fashion.

Chaos in a long-term experiment with a plankton community.

Mathematical models predict that species interactions such as competition and predation can generate chaos. However, experimental demonstrations of chaos in ecology are scarce, and have been limited to simple laboratory systems with a short duration and artificial species combinations. Here, we present the first experimental demonstration of chaos in a long-term experiment with a complex food web. Our food web was isolated from the Baltic Sea, and consisted of bacteria, several phytoplankton species, herbivorous and predatory zooplankton species, and detritivores. The food web was cultured in a laboratory mesocosm, and sampled twice a week for more than 2,300 days. Despite constant external conditions, the species abundances showed striking fluctuations over several orders of magnitude. These fluctuations displayed a variety of different periodicities, which could be attributed to different species interactions in the food web. The population dynamics were characterized by positive Lyapunov exponents of similar magnitude for each species. Predictability was limited to a time horizon of 15-30 days, only slightly longer than the local weather forecast. Hence, our results demonstrate that species interactions in food webs can generate chaos. This implies that stability is not required for the persistence of complex food webs, and that the long-term prediction of species abundances can be fundamentally impossible.