Resilience of alternative stable states during the recovery of shallow lakes from eutrophication: Lake Veluwe as a case study

In this paper we analyze a long-term dataset on the recovery from eutrophication of Lake Veluwe (The Netherlands). Clear hysteresis was observed in a number of ecosystem variables: the route to recovery differed significantly from the route that led to loss of clear water. The macrophyte dominated state disappeared in the late 1960s at TP above 0.20 mg l−1, whereas its return occurred at less than 0.10 mg TP l−1. Several regime shifts resulting in the occurrence of three alternative stable states were observed over a period of 30 years. The turbid state showed resistance to change, despite a strong and prompt reduction in Chl-a following reduction of external P-loading. The most important component that determined hysteresis in the return to clear water was not internal P-loading, but a high level of nonalgal light attenuation (through sediment resuspension) maintained by the interaction between wind and benthivorous fish. Although Chara was able to re-colonize the most shallow parts of the lake, recovery stalled and for a number of years clear (above charophyte beds) and turbid (deeper parts of the lake) water co-existed, as a separate alternative state on route to full recovery. Lake-wide clear water was re-established after bream density had been reduced substantially. This allowed a return of zebra mussels to the lake, whose high filtration capacity helped in maintaining clear water. In this study, we were able to identify the main drivers of hysteresis and regime shifts, although formal demonstration of cause and effect was not possible on the basis of field data alone. We argue that resilience of the present clear water state of Lake Veluwe very much depends on sizable populations of a few keystone species, especially Chara (stoneworts) and Dreissena (zebra mussels), and that careful management of these species is equally important as control of nutrients. Lake management should strive to maintain and strengthen resilience of the ecosystem, and this should offer protection against a renewed collapse of the clear state