Detecting and measuring senescence in wild birds: experience with long-lived seabirds.

This paper points out and discusses several practical and methodological problems that arise in attempts to detect and measure senescent declines in survival or breeding performance of wild animals, with specific emphasis on long-lived seabirds. Birds have no anatomical markers of age, so studies of age-related biology require marking individuals at the time of hatching and following them throughout their lives. Seabirds live longer than the working lifespan of biologists, and longer than the turnover times of study techniques or theories of senescence. Seabirds are exposed to changing environmental and demographic conditions and cannot be assumed to be in demographic equilibrium. Sample sizes of the oldest age-classes are always small, requiring either marking very large numbers of birds at hatching or continuing studies of old birds over many years. Incomplete sampling requires the use of mark-recapture models that have only been developed in the last 20years. Mortality selection resulting from demographic heterogeneity (selective survival of high-quality individuals) can offset or confound the effects of senescent changes within individuals. Many of these problems are amenable to solution and will be probably solved within a few years. In the meantime, this paper recommends that reviewers should be cautious about accepting published reports of senescent declines in natural populations.