Persistence, spread and the drift paradox.

We derive conditions for persistence and spread of a population where individuals are either immobile or dispersing by advection and diffusion through a one-dimensional medium with a unidirectional flow. Reproduction occurs only in the stationary phase. Examples of such systems are found in rivers and streams, marine currents, and areas with prevalent wind direction. In streams, a long-standing question, dubbed 'the drift paradox', asks why aquatic insects faced with downstream drift are able to persist in upper stream reaches. For our two-phase model, persistence of the population is guaranteed if, at low population densities, the local growth rate of the stationary component of the population exceeds the rate of entry of individuals into the drift. Otherwise the persistence condition involves all the model parameters, and persistence requires a critical (minimum) domain size. We calculate the rate at which invasion fronts propagate up- and downstream, and show that persistence and ability to spread are closely connected: if the population cannot advance upstream against the flow, it also cannot persist on any finite spatial domain. By studying two limiting cases of our model, we show that residence in the immobile state always enhances population persistence. We use our findings to evaluate a number of mechanisms previously proposed in the ecological literature as resolutions of the drift paradox.