Not-so-great tits: early-life environment drives long-term decrease in adult body mass in a wild bird population

Body mass is a key organismal characteristic that impacts many physiological and ecological processes and often a strong determinant of fitness. Recent studies have documented temporal phenotypic changes in this trait in many populations, but identifying the mechanisms underpinning these changes can be difficult. Here, we use 47 years of data to analyse how adult and nestling body mass have changed over time in a great tit Parus major population in Wytham Woods (UK). Further, we link those changes to three environmental variables previously recognised as drivers of body mass: temperature, intra- and inter-specific competition and temporal mismatch with a key prey during breeding, winter moth Operophtera brumata caterpillars. Temporal analyses of adult body mass revealed contrasting dynamics at the between- and within-cohort levels, mirroring Simpsons Paradox. At the population level we report a marked decrease in body mass in adults between 1978 and 2024 (-0.042 Haldanes), and show that this results from phenotypic plasticity, driven by a negative between-cohort trend likely reflecting carry-over effects of the early environment. Within cohorts, however, trends were consistently positive likely reflecting an age-dependent mass increase. The change in adults was paralleled by a change in nestling body mass (-0.036 Haldanes). Nestling mass was negatively associated with estimated intensity of intraspecific competition, as well as inter-specific competition from blue tits Cyanistes caeruleus, as quantified by local population density. These effects carried over to adulthood, as shown by a negative association between adult mass and the population density experienced at early life. Temperature during development and mismatch with the caterpillar food supply, despite being associated with adult and nestling mass, did not explain the observed declines in mass, largely because these have not changed over time. Overall, our results illustrate the potential for effects mediated early in development to carry-over into long-term phenotypic change at later life history stages, and emphasise the value of considering multiple effects as drivers of phenotypic change in natural populations.