Abrupt transitions of the top-down controlled Black Sea pelagic ecosystem during 1960–2000: Evidence for regime-shifts under strong fishery exploitation and nutrient enrichment modulated by climate-induced variations

Abstract

Functioning of the Black Sea ecosystem has profoundly changed since the early 1970s under cumulative effects of excessive nutrient enrichment, strong cooling/warming, over-exploitation of pelagic fish stocks, and population outbreak of gelatinous carnivores. Applying a set of criteria to the long-term (1960–2000) ecological time-series data, the present study demonstrates that the Black Sea ecosystem was reorganised during this transition phase in different forms of top-down controlled food web structure through successive regime-shifts of distinct ecological properties. The Secchi disc depth, oxic–anoxic interface zone, dissolved oxygen and hydrogen sulphide concentrations also exhibit abrupt transition between their alternate regimes, and indicate tight coupling between the lower trophic food web structure and the biogeochemical pump in terms of regime-shift events. The first shift, in 1973–1974, marks a switch from large predatory fish to small planktivore fish-controlled system, which persisted until 1989 in the form of increasing small pelagic and phytoplankton biomass and decreasing zooplankton biomass. The increase in phytoplankton biomass is further supported by a bottom-up contribution due to the cumulative response to high anthropogenic nutrient load and the concurrent shift of the physical system to the “cold climate regime” following its ∼20-year persistence in the “warm climate regime”. The end of the 1980s signifies the depletion of small planktivores and the transition to a gelatinous carnivore-controlled system. By the end of the 1990s, small planktivore populations take over control of the system again. Concomitantly, their top-down pressure when combined with diminishing anthropogenic nutrient load and more limited nutrient supply into the surface waters due to stabilizing effects of relatively warm winter conditions switched the “high production” regime of phytoplankton to its background “low production” regime. The Black Sea regime-shifts appear to be sporadic events forced by strong transient decadal perturbations, and therefore differ from the multi-decadal scale cyclical events observed in pelagic ocean ecosystems under low-frequency climatic forcing. The Black Sea observations illustrate that eutrophication and extreme fishery exploitation can indeed induce hysteresis in large marine ecosystems, when they can exert sufficiently strong forcing onto the system. They further illustrate the link between the disruption of the top predators, proliferation of new predator stocks, and regime-shift events. Examples of these features have been reported for some aquatic ecosystems, but are extremely limited for large marine ecosystems.