Parental provisioning drives brain size in birds

Larger brains should be adaptive because they support numerous eco- and socio-cognitive benefits, but these benefits explain only a modest part of the interspecific variation in brain size. Notably underexplored are the high energetic costs of developing brains, and thus the possible role of parental provisioning in the evolution of adult brain size. We explore this idea in birds, which show considerable variation in both socio-ecological traits and the energy transfer from parents to offspring. Comparative analyses of 1,176 bird species show that the combination of adult body mass, mode of development at hatching, relative egg mass, and the time spent provisioning the young in combination strongly predict relative brain size across species. Adding adult eco- and socio-cognitive predictors only marginally adds explanatory value. We therefore conclude that parental provisioning enabled bird species to evolve into skill-intensive niches, reducing interspecific competition and consequently promoting survival prospects and population stability. Critically, parental provisioning also explains why precocial bird species have smaller brains than altricial ones. Finally, these results suggest that the cognitive adaptations that provide the behavioral flexibility to improve reproductive success and survival are intrinsically linked to successful parental provisioning. Our findings also suggest that the traditionally assessed cognitive abilities may not predict relative brain size.

Hierarchical fear: parental behaviour and corticosterone release mediate nestling growth in response to predation risk

Nestling development, a critical life-stage for altricial songbirds, is highly vulnerable to predation, particularly for open-cup nesting species. Since nest predation risk increases cumulatively with time, rapid growth may be an adaptive response that promotes early fledging. However, greater predation risk can reduce parental provisioning rate as a risk aversion strategy and subsequently constrain nestling growth, or directly elicit a physiological response in nestlings with adaptive or detrimental effects on development rate. Despite extensive theory, evidence for the relative strength of these effects on nestling development in response to prevailing predation risk and the underlying mechanisms remain unclear. For an alpine population of horned lark (Eremophila alpestris), we elevated perceived predation risk (decoys/playback) during the nestling stage to assess the influence of predator cues and parental care on nestling wing growth and the glucocorticoid hormone corticosterone. We used piecewise path analysis to test a hypothesized causal response structure composed of direct and indirect pathways. Nestlings under greater perceived predation risk reduced corticosterone and increased wing growth, resulting in an earlier age at fledge. This represented both a direct response that was predator-specific, and an indirect response dependent on parental provisioning rate. Parents that reduced provisioning rate most severely in response to predator cues had smaller nestlings with greater corticosterone. Model comparisons indicated the strongest support for a directed, causal influence of corticosterone on nestling wing growth, highlighting corticosterone as a potential physiological mediator of the nestling growth response to predation risk. Finally, cold temperatures prior to the experiment constrained wing growth closer to fledge, illustrating the importance of considering the combined influence of weather and predation risk across developmental stages. We present the first study to separate the direct and indirect effects of predation risk on nestling development in a causal, hierarchical framework that incorporates corticosterone as an underlying mechanism and provides experimental evidence for an adaptive developmental response to predation risk in ground-nesting songbirds.

Endothermy, offspring size and evolution of parental provisioning in vertebrates

Mammals and birds provide food for their young after birth, but such provisioning is almost absent in other vertebrates. A recent theory suggested that, in addition to thermoregulation, the large discrepancy in size between adult and young ectothermic vertebrates precludes them from providing for their young, whereas the relatively large offspring of endotherms are easier to provision. I show here that reptile neonates and hatchlings are as large as those of mammals and birds. Differences in size between adults and young thus cannot explain the lack of parental provisioning in reptiles. I suggest that the large size at birth is the ancestral condition in amniotes as a whole and that provisioning has thus evolved after endothermy.

Effects of manipulated levels of predation threat on parental provisioning and nestling begging

Parental provisioning behavior is a major determinant of offspring growth and survival, but high provisioning rates might come at the cost of increased predation threat. Parents should thus adjust provisioning activity according to current predation threat levels. Moreover, life-history theory predicts that response to predation threat should be correlated with investment in current reproduction. We experimentally manipulated perceived predation threat in free-living great tits (Parus major) by presenting parents with a nest predator model while monitoring different aspects of provisioning behavior and nestling begging. Experiments were conducted in 2 years differing greatly in ecological conditions, including food availability. We further quantified male territorial aggressiveness and male and female exploratory tendency. Parents adjusted provisioning according to current levels of threat in an apparently adaptive way. They delayed nest visits during periods of elevated perceived predation threat and subsequently compensated for lost feeding opportunities by increasing provisioning once the immediate threat had diminished. Nestling begging increased after elevated levels of predation threat, but returned to baseline levels by the end of the experiment, suggesting that parents had fully compensated for lost feeding opportunities. There was no evidence for a link between male exploration behavior or aggressiveness and provisioning behavior. In contrast, fast-exploring females provisioned at higher rates, but only in the year with poor environmental conditions, which might indicate a greater willingness to invest in current reproduction in general. Future work should assess whether these personality-related differences in delivery rates under harsher conditions came at a cost of reduced residual reproductive value.