1Nestling development, a critical life-stage for altricial songbirds, is highly vulnerable to 2 predation, particularly for open-cup nesting species. Since nest predation risk increases 3 cumulatively with time, rapid growth may be an adaptive response that promotes early fledging. 4 However, greater predation risk can reduce parental provisioning rate as a risk aversion strategy 5 and subsequently constrain nestling growth, or directly elicit a physiological response in 6 nestlings with adaptive or detrimental effects on development rate. Despite extensive theory, 7 evidence for the relative strength of these effects on nestling development in response to 8 prevailing predation risk and the underlying mechanisms remain unclear. For an alpine 9 population of horned lark (Eremophila alpestris), we elevated perceived predation risk 10 (decoys/playback) during the nestling stage to assess the influence of predator cues and parental 11 care on nestling wing growth and the glucocorticoid hormone corticosterone. We used piecewise 12 path analysis to test a hypothesized causal response structure composed of direct and indirect 13 pathways. Nestlings under greater perceived predation risk reduced corticosterone and increased 14 wing growth, resulting in an earlier age at fledge. This represented both a direct response that 15 was predator-specific, and an indirect response dependent on parental provisioning rate. Parents 16 that reduced provisioning rate most severely in response to predator cues had smaller nestlings 17 with greater corticosterone. Model comparisons indicated the strongest support for a directed, 18 causal influence of corticosterone on nestling wing growth, highlighting corticosterone as a 19 potential physiological mediator of the nestling growth response to predation risk. Finally, cold 20 temperatures prior to the experiment constrained wing growth closer to fledge, illustrating the 21 importance of considering the combined influence of weather and predation risk across 22developmental stages. We present the first study to separate the direct and indirect effects of 23 3 predation risk on nestling development in a causal, hierarchical framework that incorporates 24 corticosterone as an underlying mechanism and provides experimental evidence for an adaptive 25 developmental response to predation risk in ground-nesting songbirds. 26 27