Balancing selection is frequently invoked as a mechanism to maintain variation within and across populations. However, rigorous tests demonstrating balancing selection operating in nature are scarce, particularly on complex traits, which frequently display high levels of variation. Leveraging a foca

Fitness trade-offs across episodes of selection and environments influence life-history evolution and adaptive population divergence. Documenting these trade-offs remains challenging as selection can vary in magnitude and direction through time and space. Here, we evaluate fitness trade-offs at the