Although it has been shown that phenology can respond to temporal environmental variation in free ranging populations of several species, little is known about the mechanisms of these responses and their effects on demography, and in particular on survival. Exploring phenological responses and their associated consequences on survival can be achieved at two distinct scales: the population scale, which focusses on a set of common responses to environmental conditions, and the individual scale, focusing on the relative position of each individual in the distribution of survival and phenology under particular conditions. In this study, we apply capture-mark-recapture multistate modelling on a 38-year monitoring dataset of blue tits (Cyanistes caeruleus) to investigate the effects of breeding phenology and some demographic covariates (breeding density, average and individual breeding success) on adult survival, at both population and individual scales. Our analysis revealed that (i) at the population scale, early breeding years are followed by lower average adult survival. (ii) At the individual level, earlier breeders within the population have higher subsequent survival than later breeders, although this relationship is reversed in years with very harsh conditions, e.g. warm spring and high breeding density. (iii) High individual relative breeding success is also associated with higher subsequent survival and explains more survival variation than relative phenology. Overall, our study indicates that, although earlier breeding is associated with a survival cost at the population level, substantial intrapopulation hererogeneity shapes a positive association between earlier breeding, breeding success and survival at the individual level.

中文翻译：

---

早期生殖与更高或更低的存活率相关吗？蓝雀个体和人口规模之间的对立结果

尽管已经表明物候学可以对几种物种的自由散居种群的时间环境变化做出反应，但对于这些反应的机制及其对人口统计学尤其是生存的影响知之甚少。探索物候响应及其对生存的影响可以通过两个不同的规模来实现：人口规模（专注于对环境条件的一系列共同反应）和个人规模（专注于每个人在人类分布中的相对位置）在特定条件下的生存率和物候。在这项研究中，我们对蓝山雀（Cyanistes caeruleus）的38年监测数据集应用了捕获标记-捕获-多状态建模，以研究育种物候和某些人口统计学协变量（育种密度，种群和个体规模的成年存活率）。我们的分析表明，（i）在种群规模上，繁殖早期是成年人平均存活率较低的时期。（ii）在个体层面上，种群中较早的育种者比较晚的育种者具有更高的后继存活率，尽管这种关系在条件极为恶劣的年份（如温暖的春季和高繁殖密度）中会逆转。（iii）较高的个人相对育种成功率也与较高的随后存活率有关，并且比相对物候学解释了更多的存活变异。总体而言，我们的研究表明，尽管早期繁殖与种群水平的生存成本相关，但大量的种群内异质性在早期繁殖之间形成了正相关关系，