Invasion of a Top Predator into an Epipelagic Ecosystem can bring a Paradoxical Top-Down Trophic Control

We apply mathematical modeling to explore different scenarios of invasion of a top predator (carnivorous zooplankton or planktivorous fish) into an epipelagic plankton ecosystem. We use a ‘minimal’ model of three nonlinear ordinary differential equations (nutrient–phytoplankton–herbivores) with the top predator density as a time-dependent parameter. The ecosystem shows different types of response, which can be described in terms of top-down trophic control. Our investigation indicates that under certain conditions the plankton ecosystem model demonstrates a surprising kind of response: in a wide range of realistic ecosystem parameters the invasion of the top predator leads to a prominent increase in the average density of zooplankton and to a resulting decrease of phytoplankton density. This phenomenon is opposite to the ‘typical’ top–down control when the carnivore pressure decreases zooplankton density which, in turn, increases phytoplankton biomass. We call the revealed type of top-down control ‘paradoxical’. Examples of such a response in natural aquatic ecosystems were reported earlier but no clear explanation has been provided hitherto. In this paper, we analyze possible mechanisms of ‘paradoxical top–down control’ and show that it can occur in eutrophic epipelagic ecosystems subject to high rate of cross-pycnocline exchange.