Proximate and evolutionary sources of variation in offspring energy expenditure in songbirds

Aim Understanding variation in offspring energy expenditure is important because energy is critical for growth and development. Weather may exert proximate effects on offspring energy expenditure, but in altricial species these might be masked by parental care and huddling with siblings. Such effects are particularly important to understand given changing global weather patterns, yet studies of wild offspring in the presence of parental care are lacking. Offspring energy expenditure may also vary among species due to evolved responses to environmental selection pressures, requiring studies at both proximate and ultimate levels. Location USA, South Africa, Malaysia. Time period 2016-2019. Major taxa studied Songbirds. Methods We used the doubly-labelled water technique to estimate nestling daily energy expenditure of 54 songbird species across three continents. We used Bayesian phylogenetic mixed models to test proximate and evolutionary causes of variation in offspring energy expenditure while accounting for phylogeny and phylogenetic uncertainty. Results Offspring energy expenditure increased with more rainfall and colder air temperatures, but decreased among offspring in broods with more siblings. Across species, nestling and adult mortality, but not growth rate, were positively associated with offspring energy use. Main conclusions Weather had clear proximate effects on offspring energy expenditure and parents were either unable or unwilling to fully offset these effects. However, the decrease in offspring energy use when huddling with more siblings demonstrated a modulating effect of life history traits. For example, high nest predation rates favour reduced parental care and can force offspring to spend more energy coping with environmental conditions. Furthermore, reduced energy expenditure is thought to facilitate increased longevity, which is increasingly realized with lower extrinsic mortality rates, providing an explanation for the positive association between adult mortality and offspring energy expenditure. Ultimately, both proximate and evolutionary influences need to be considered to better understand causes of offspring energetics.