Abstract
Harvesting has been demonstrated to cause rapid, yield-decreasing trait change towards slower somatic growth and earlier maturation in wild populations. These changes are largely considered to result from direct, density-independent harvest selection on traits. Here, we show that exact same trait changes may also indirectly result from a harvest-induced relaxation of density-dependent (K) natural selection for faster growth and delayed maturation. We exposed 12 pond populations of medaka fish (Oryzias latipes) to contrasted size-selective harvesting during 5 years, and show that harvesting effectively changed juvenile natural mortality from density-dependent to density-independent. We then laboratory-reared medaka progeny under contrasted food levels mimicking the environmental effects of a harvest-induced density gradient. Interaction between past harvest regime and present food environment on progeny traits revealed that harvest-induced trait changes in medaka resulted from selection in a low-food environment only, i.e., were driven by relaxed K-selection only, not by direct harvest selection. Feeding trials further demonstrated that trait changes were associated with reorganizations in rates of food acquisition, assimilation and allocation that were contingent upon the food environments. This is the first study to demonstrate that harvesting can induce undesirable distortions of natural selection that impair productivity traits. We conclude that sustaining harvesting yields over extended time scales requires a preservation of high population densities.