When does greater mortality increase population size? The long history and diverse mechanisms underlying the hydra effect

The phenomenon of a population increasing in response to an increase in its per-capita mortality rate has recently been termed the 'hydra effect'. This article reviews and unifies previous work on this phenomenon. Some discrete models of density-dependent growth were shown to exhibit hydra effects in 1954, but the topic was then ignored for decades. Here the history of research on the hydra effect is reviewed, and the key factors producing it are explored. Mortality that precedes overcompensatory density dependence always has the potential to produce hydra effects. Even when mortality follows density dependence, hydra effects may occur in unstable systems due to changes in the amplitude and/or form of population cycles. An increase in resource productivity due to lower consumption rates following increased consumer mortality can also produce a hydra effect. Lower consumption can come about as the result of increased satiation of the consumers or changes in behaviour of either consumer or resource species that reduce the mean attack rate. Changes in species composition of a resource community may also decrease the average attack rate. Population structure can promote hydra effects by allowing separation of the timing of density dependence and mortality, although stage-specific density dependence usually decreases hydra effects.