The seasonal decline of avian clutch size may result from the conflict between the advantage of early breeding (greater offspring value) and the advantage of a delay in lay date (improved body condition and hence clutch size). We tested predictions of a condition-dependent individual optimization model based on this trade-off (Rowe et al. 1994) in a long-distance migrant, the greater snow goose (Chen caerulescens atlantica), using data on condition, migration, and reproductive decisions of individuals. We closely tracked radio-marked females at their main spring staging area and on their breeding grounds. Our results were consistent with predictions of the model. Early-arriving females had a longer prelaying period and initiated their nests earlier than late arrivals. After controlling statistically for arrival date, we determined that females with high premigration condition had an earlier lay date than those in low condition. After controlling for the seasonal decline (i.e., lay date), we observed that clutch size was not related to premigration condition. Moreover, we took advantage of an unplanned manipulation of the prebreeding condition that occurred during our long-term study. We found that a reduction in condition caused a delay in lay date. However, after controlling for the seasonal decline, it did not affect clutch size. Our study indicates that geese simultaneously adjust their lay date and clutch size according to their premigration condition and migratory behavior as predicted by the condition-dependent optimization model.
Summary: We investigated the hypothesis that cyclic lemming populations indirectly affect arctic‐nesting greater snow geese (Anser caerulescens atlanticus L.) through the behavioural and numerical responses of shared predators. The study took place on Bylot Island in the Canadian High Arctic during two lemming cycles. We recorded changes in foraging behaviour and activity rate of arctic foxes, parasitic jaegers, glaucous gulls and common ravens in a goose colony during one lemming cycle and we monitored denning activity of foxes for 7 years. We also evaluated the total response of predators (i.e. number of eggs depredated). Arctic foxes were more successful in attacking lemmings than goose nests because predators were constrained by goose nest defence. Predators increased their foraging effort on goose eggs following a lemming decline. Activity rates in the goose colony varied 3·5‐fold in arctic foxes and 4·8‐fold in parasitic jaegers, and were highest 2 and 3 years after the lemming peak, respectively. The breeding output of arctic foxes appeared to be driven primarily by lemming numbers. Predators consumed 19–88% of the annual goose nesting production and egg predation intensity varied 2·7‐fold, being lowest during peak lemming years. Arctic foxes and parasitic jaegers were the key predators generating marked annual variation in egg predation. Our study provides strong support for short‐term, positive indirect effects and long‐term, negative indirect effects of lemming populations on arctic‐nesting geese. The outcome between these opposing indirect effects is probably an apparent competition between rodents and many terrestrial arctic‐nesting birds.
Summary: For many species, human‐induced disturbances can severely influence an individual's pay‐off; however, energy‐cost variations from different disturbance types have rarely been reported. We evaluated the dynamic behavioural responses of staging greater snow geese Anser caerulescens atlanticus to different types of disturbance in southern Quebec, Canada, between 1997 and 2000. We specifically considered the impact of a unique measure, a spring conservation hunt implemented in agricultural habitats in 1999. We tracked 237 radio‐tagged females for 2764 h and recorded 697 take‐offs following fortuitous disturbance, scaring and hunting in three regions characterized by different habitats. Geese used cornfields in south‐western Quebec, Scirpus marshes and hayfields in the upper St Lawrence estuary, and Spartina marshes and hayfields in the lower estuary. Overall, disturbance levels increased in the upper and lower estuary during years with hunting, mostly through an increase in hunting and scaring activities. The probability of geese returning to a refuge after disturbance in agricultural habitats increased in years with hunting except in the corn‐growing region. The short‐term energy gain of geese resuming feeding after disturbance was less than before disturbance, and this difference was greater in years with hunting. Distances flown after disturbance decreased with flock size and were longer after scaring and hunting than after fortuitous disturbances in the Scirpus region. Overall, habitat use varied among years and associated estimated energy gain decreased markedly in both years with hunting in the Spartina and corn‐growing region, but did not change in the Scirpus region. Changes in behaviour due to disturbance, and especially those associated with hunting, probably contributed to the reduced body condition of staging greater snow geese during years with hunting. Synthesis and applications. From a methodological viewpoint, we highlight the importance of tracking the behaviour of individual animals after disturbance to properly evaluate its impact. From a conservation perspective, we provide empirical arguments to limit the hunting of breeding waterfowl during their prenuptial migration in order to facilitate their fattening and forthcoming reproduction. From a management standpoint, we suggest that a side‐effect of disturbance induced by spring hunting to control overabundant populations may be reduced fattening and breeding output among birds that survive. Together, these data emphasize further the importance of measuring the direct and indirect effects of disturbance rather than assuming effects from the incidence of the disturbance alone.