Carry-over effects and plasticity to temperature shape phenology across life stages and generations

Climate change is advancing spring phenology in temperate systems, with the potential to disrupt synchrony between trophic levels. Predicting these shifts requires understanding not only direct plastic responses to temperature, but also how plasticity at one stage carries over to shape timing at subsequent stages. We experimentally quantified direct thermal plasticity in phenology and its carry-over effects across the full life cycle of the winter moth (Operophtera brumata), a holometabolous insect whose fitness relies on synchrony with host plant budburst. Using a fully factorial split-clutch rearing experiment across four temperature treatments, we exposed individuals to contrasting conditions at each life stage and used structural equation modelling to partition direct and carry-over effects on phenology. Each life stage showed distinct plastic responses to temperature. Carry-over effects transmitted approximately 0.38 days/day of plastic advance to the next life stage on average, with the remainder absorbed by compensatory changes in the duration of the subsequent life stage. Together, these results show that carry-over effects propagate plastic responses across the life cycle, which are partially buffered by compensatory changes in developmental duration. Accurate predictions of phenological shifts under climate change therefore require models that account for carry-over effects and developmental compensation across life stages.