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 1.We investigated the hypothesis that cyclic lemming populations indirectly affect arcticnesting greater snow geese ( Anser caerulescens atlanticus L.) through the behavioural and numerical responses of shared predators. 2. 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). 3. 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. 4. 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. 5. 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. 6. 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.

Quantifying the costs and benefits of migration distance is critical to understanding the evolution of long-distance migration. In migratory birds, life history theory predicts that the potential survival costs of migrating longer distances should be balanced by benefits to lifetime reproductive success, yet quantification of these reproductive benefits in a controlled manner along a large geographical gradient is challenging. We measured a controlled effect of predation risk along a 3350-kilometer south-north gradient in the Arctic and found that nest predation risk declined more than twofold along the latitudinal gradient. These results provide evidence that birds migrating farther north may acquire reproductive benefits in the form of lower nest predation risk.

The strategy of relying extensively on stored nutrient reserves for reproduction (capital breeding) was thought to be common in large‐bodied birds breeding in harsh environments, such as arctic‐nesting geese, but this view has been challenged recently. Our objective was to model inputs to the eggs from stored reserves and from local food plants in Greater Snow Geese (Chen caerulescens atlantica) breeding in the high Arctic, using a new approach based on stable‐isotope signatures. Snow Geese and their eggs were collected during laying from 1999 to 2001 (N = 66 females and 110 eggs). We analyzed the isotopic signature (δ13C and δ15N) of egg constituents (lipid‐free yolk, yolk lipid, and albumen), goose tissues (lipid‐free breast muscles, abdominal fat, and whole liver) and of the food plants eaten by laying geese in the Arctic (graminoids and forbs). We applied a two‐isotope mixing model approach to delineate nutrient input to eggs quantitatively. Differences in the isotopic signature of endogenous reserves and arctic food plants were relatively large (5.3–8.0‰ for Δδ13C and 7.5‰ for Δδ15N) because reserves were accumulated in southern staging areas where geese feed in farmlands and estuarine habitats. The percentage of egg nutrients derived from exogenous sources (food consumed in the Arctic) was higher than from endogenous (body) reserves and varied little among the three years. Isotopic signatures indicated that endogenous reserves contributed 33% of lipid‐free yolk nutrients, 27% of albumen, and 20% of yolk lipid, on average. Isotopic signatures of egg constituents of individual females were more strongly related to those of liver than endogenous sources (breast muscles or abdominal fat), indicating that the endogenous isotopic signature was diluted by a dietary input in the liver. We also found evidence of seasonal variation in the use of endogenous reserves. Late‐laying females apparently invested proportionally more endogenous reserves in their eggs than did early layers, but not those laying larger clutches. We conclude that Greater Snow Geese use a mixed capital/income breeding strategy. Our study shows that isotopic composition of tissues can be used to infer the contribution of exogenous vs. endogenous sources of nutrients for egg formation where inputs differ isotopically.

The suggested link between lemming cycles and reproductive success of arctic birds is caused by potential effects of varying predation pressure (the Alternative Prey Hypothesis, APH) and protective association with birds of prey (the Nesting Association Hypothesis, NAH). We used data collected over two complete lemming cycles to investigate how fluctuations in lemming density were associated with nesting success of greater snow geese (Anser caerulescens atlanticus) in the Canadian High Arctic. We tested predictions of the APH and NAH for geese breeding at low and high densities. Goose nesting success varied from 22% to 91% between years and the main egg predator was the arctic fox (Alopex lagopus). Nesting associations with snowy owls (Nyctea scandiaca) were observed but only during peak lemming years for geese nesting at low density. Goose nesting success declined as distance from owls increased and reached a plateau at 550 m. Artificial nest experiments indicated that owls can exclude predators from the vicinity of their nests and thus reduce goose egg predation rate. Annual nest failure rate was negatively associated with rodent abundance and was generally highest in low lemming years. This relationship was present even after excluding goose nests under the protective influence of owls. However, nest failure was inversely density‐dependent at high breeding density. Thus, annual variations in nest density influenced the synchrony between lemming cycles and oscillations in nesting success. Our results suggest that APH is the main mechanism linking lemming cycles and goose nesting success and that nesting associations during peak lemming years (NAH) can enhance this positive link at the local level. The study also shows that breeding strategies used by birds (the alternative prey) could affect the synchrony between oscillations in avian reproductive success and rodent cycles.

Arctic wildlife is often presented as being highly at risk in the face of current climate warming. We use the long-term (up to 24 years) monitoring records available on Bylot Island in the Canadian Arctic to examine temporal trends in population attributes of several terrestrial vertebrates and in primary production. Despite a warming trend (e.g. cumulative annual thawing degree-days increased by 37% and snow-melt date advanced by 4–7 days over a 23-year period), we found little evidence for changes in the phenology, abundance or productivity of several vertebrate species (snow goose, foxes, lemmings, avian predators and one passerine). Only primary production showed a response to warming (annual above-ground biomass of wetland graminoids increased by 123% during this period). We nonetheless found evidence for potential mismatches between herbivores and their food plants in response to warming as snow geese adjusted their laying date by only 3.8 days on average for a change in snow-melt of 10 days, half of the corresponding adjustment shown by the timing of plant growth (7.1 days). We discuss several reasons (duration of time series, large annual variability, amplitude of observed climate change, nonlinear dynamic or constraints imposed by various rate of warming with latitude in migrants) to explain the lack of response by herbivores and predators to climate warming at our study site. We also show how length and intensity of monitoring could affect our ability to detect temporal trends and provide recommendations for future monitoring.

In seasonal environments, breeding events must be synchronized with resource peaks to ensure production and growth of offspring. As changes in climate may affect trophic levels differentially, we hypothesized that a lack of synchrony between chick hatch and resource peaks could decrease growth rates in chicks of shorebirds nesting in the High Arctic. To test this hypothesis, we compared growth curves of chicks hatching in synchrony with peak periods of food abundance to those hatching outside of these peak periods. We also tested for changes in lay dates of shorebirds in the Canadian Arctic using recent and historical data. Mean daily temperatures during the laying period increased since the 1950s by up to 1.5 °C, and changes in lay dates were apparent for three shorebird species, yet differences in median lay dates between 1954 and 2005–2008 were only significant for White-rumped Sandpiper ( Calidris fuscicollis (Viellot, 1819)). During 2005–2008, there was only 1 year of relatively high synchrony between hatch and resource peaks. Asynchrony between hatch and peaks in Tipulidae biomass reduced growth rates in chicks of Baird’s Sandpiper (Calidris bairdii (Coues, 1861)). As anticipated changes in climate may decouple phenological events, the effects of asynchrony on growth rates of arctic-nesting birds warrant further investigation.

Summary1. Optimality theory predicts that both timing of arrival and arrival state on the breeding area will determine reproductive timing and investment in migratory organisms. We tested this idea using a condition-dependent individual optimization model (Ardea 68, 1980, 225 and The American Naturalist 143, 1994, 698) in common eider ducks through descriptive data, path analyses and experimental manipulation. 2. Our results support the causal pathways drawn from the optimization model indicating that individuals adjust their reproductive decisions as a function of their arrival date and body condition at arrival. 3. Independent of body condition, early-arriving females had a longer pre-laying period, but still initiated their nests earlier, and produced larger clutches than late-arriving birds. Independent of arrival date, females in good condition laid earlier than those in poor condition. Manipulation of pre-laying female body condition confirmed that the relationship between condition and laying date was causal. 4. Female common eiders appear to optimize reproductive decisions in response to both their external (i.e. environmental conditions affecting the egg-value) and internal (i.e. body condition) states. These adjustments seem to minimize the fitness costs of reproduction, in which higher clutch size is not associated with an apparent lower survival or future breeding probability. 5. Our study emphasizes the importance of (i) simultaneously considering the timing of migration, the state of individuals and the seasonal change in egg-value to understand clearly birds' breeding decisions and (ii) appreciating the potential proximate and ultimate factors explaining why some individuals delay breeding and ⁄ or produce small clutches.