Proactive vs. reactive multi-brooding: divergent drivers of double brooding and replacement clutching in varied tits
摘要
Multibrooding is a common avian reproductive strategy in which individuals produce multiple broods within a single breeding season. Theoretically, this behavior enhances fitness by maximizing reproductive output under favorable conditions. However, within a species, not all individuals adopt this tactic, prompting questions about the factors motivating multiple breeding attempts. In this study, we investigated the ecological and individual factors influencing the initiation of second clutches in the Varied Tit (Sittiparus varius), a species capable of multibrooding. We explicitly distinguished between replacement clutches and true second clutches (double brooding) to identify their distinct drivers. Our results indicated that double brooding was a proactive strategy primarily driven by an earlier first egg-laying date. Path analysis further showed that the effect of age on double brooding was entirely indirect, mediated through earlier breeding phenology. In contrast, replacement clutching following nest failure was a reactive, time-constrained tactic, associated with an earlier nest failure date and heavier average egg mass in the initial failed clutch. Once a second breeding attempt was initiated, both strategies exhibited convergent adjustments in clutch size, highlighting the late-season environmental constraints on breeding decisions. This study highlights distinct evolutionary pathways for reproductive flexibility: double brooding serves as a proactive mechanism for older-experienced individuals to maximize reproductive output, whereas replacement clutching serves as a time-limited reactive compensatory response following failure.