Fledging size and survival in snow geese: Timing is everything (or is it?)

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Abstract
In many birds, body size at fledging is assumed to predict accurately the probability of subsequent survival, and size at fledging is often used as a proxy variable in analyses attempting to assess the pattern of natural selection on body size. However, in some species, size at fledging can vary significantly as a function of variation in the environmental component of growth. Such developmental plasticity has been demonstrated in several species of Arctic-breeding geese. In many cases, slower growth and reduced size at fledging has been suggested as the most parsimonious explanation for reduced post-fledging survival in goslings reared under poor environmental conditions. However, simply quantifying a relationship between mean size at fledging and mean survival rate (Francis et al ., 1992) may obscure the pattern of selection on the interaction of the genetic and environmental components of growth. The hypothesis that selection operates on the environmental component of body size at fledging, rather than the genetic component of size per se, was tested using data from the long-term study of Lesser Snow Geese ( Anser c. caerulescens ) breeding at La Pérouse Bay, Manitoba, Canada. Using data from female goslings measured at fledging, post-fledging survival rates were estimated using combined live encounter and dead recovery data (Burnham, 1993). To control for the covariation between growth and environmental factors, survival rates were constrained to be functions of individual covariation of size at fledging, and various measures of the timing of hatch; in all Arctic-breeding geese studied to date, late hatching goslings grow significantly more slowly than do early hatching goslings. The slower growth of late-hatching goslings has been demonstrated to reflect systematic changes in the environmental component of growth, and thus controlling for hatch date controls for a significant proportion of variation in the environmental component of growth. The relationship between size at fledging, hatch date and survival was found to be significantly non-linear; among early hatching goslings, there was little indication of significant differences in survival rate among large and small goslings. However, with increasingly later hatch dates, there was progressively greater mortality selection against smaller, slower growing goslings in most years. This would appear to suggest that body size matters, but not absolutely; small size leads to reduced survival for late-hatching goslings only at La Pe´rouse Bay. Since at least some of the variation in size among goslings for a given hatch date reflects genetic differences, this suggests selection may favour larger size at fledging, albeit only among late-hatching goslings.