From gestation to weaning: Combining robust design and multi-event models unveils cost of lactation in a large herbivore.

The cost of current reproduction on survival or future reproduction is one of the most studied trade-offs governing resource distribution between fitness components. Results have often been clouded, however, by the existence of individual heterogeneity, with high-quality individuals able to allocate energy to several functions simultaneously, at no apparent cost. Surprisingly, it has also rarely been assessed within a breeding season by breaking down the various reproductive efforts of females from gestation to weaning, even though resource availability and energy requirements vary greatly. We filled this gap by using an intensively monitored population of Pyrenean chamois and by expanding a new methodological approach integrating robust design in a multi-event framework. We distinguished females that gave birth or not, and among reproducing females whether they lost their kid or successfully raised it until weaning. We estimated spring and summer juvenile survival, investigated whether gestation, lactation or weaning incurred costs on the next reproductive occasion, and assessed how individual heterogeneity influenced the detection of such costs. Contrary to expectations if trade-offs occur, we found a positive relationship between gestation and adult survival suggesting that non-breeding females are in poor condition. Costs of reproduction were expressed through negative relationships between lactation and both subsequent breeding probability and spring juvenile survival. Such costs could be detected only once individual heterogeneity (assessed as two groups contrasting good vs. poor breeders) and time variations in juvenile survival were accounted for. Early lactation decreased the probability of future reproduction, providing quantitative evidence of the fitness cost of this period recognized as the most energetically demanding in female mammals and critical for neonatal survival. The new approach employed made it possible to estimate two components of kid survival that are often considered practically unavailable in free-ranging populations, and also revealed that reproductive costs appeared only when contrasting the different stages of reproductive effort. From an evolutionary perspective, our findings stressed the importance of the temporal resolution at which reproductive cost is studied, and also provided insights on the reproductive period during which internal and external factors would be expected to have the greatest fitness impact.