1. Department of Zoology, University of Wisconsin–Madison, Madison, Wisconsin 53706, USA
2. Myvatn Research Station and Institute of Biology, University of Iceland, Sturlugata 7, IS-101 Reykjavik, Iceland
3. School of Biological Sciences, Royal Holloway, University of London, Egham, Surrey TW20 0EX, UK

Correspondence to: Anthony R. Ives¹ Correspondence and requests for materials should be addressed to A.R.I. (Email: arives@wisc.edu).

Abstract

Complex dynamics are often shown by simple ecological models^{1, 2} and have been clearly demonstrated in laboratory^{3, 4} and natural systems^{5, 6, 7, 8, 9}. Yet many classes of theoretically possible dynamics are still poorly documented in nature. Here we study long-term time-series data of a midge, Tanytarsus gracilentus (Diptera: Chironomidae), in Lake Myvatn, Iceland. The midge undergoes density fluctuations of almost six orders of magnitude. Rather than regular cycles, however, these fluctuations have irregular periods of 4–7 years, indicating complex dynamics. We fit three consumer–resource models capable of qualitatively distinct dynamics to the data. Of these, the best-fitting model shows alternative dynamical states in the absence of environmental variability; depending on the initial midge densities, the model shows either fluctuations around a fixed point or high-amplitude cycles. This explains the observed complex population dynamics: high-amplitude but irregular fluctuations occur because stochastic variability causes the dynamics to switch between domains of attraction to the alternative states. In the model, the amplitude of fluctuations depends strongly on minute resource subsidies into the midge habitat. These resource subsidies may be sensitive to human-caused changes in the hydrology of the lake, with human impacts such as dredging leading to higher-amplitude fluctuations. Tanytarsus gracilentus is a key component of the Myvatn ecosystem, representing two-thirds of the secondary productivity of the lake¹⁰ and providing vital food resources to fish and to breeding bird populations^{11, 12}. Therefore the high-amplitude, irregular fluctuations in midge densities generated by alternative dynamical states dominate much of the ecology of the lake.

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