Climate warming challenges animals to advance their timing of reproduction [1], but many animals appear to be unable to advance at the same rate as their food species [2, 3]. As a result, mismatches can arise between the moment of largest food requirements for their offspring and peak food availability [4-6], with important fitness consequences [7]. For long-distance migrants, adjustment of phenology to climate warming may be hampered by their inability to predict the optimal timing of arrival at the breeding grounds from their wintering grounds [8]. Arrival can be advanced if birds accelerate migration by reducing time on stopover sites [9, 10], but a recent study suggests that most long-distance migrants are on too tight a schedule to do so [11]. This may be different for capital-breeding migrants, which use stopovers not only to fuel migration but also to acquire body stores needed for reproduction [12-14]. By combining multiple years of tracking and reproduction data, we show that a long-distance migratory bird (the barnacle goose, Branta leucopsis) accelerates its 3,000 km spring migration to advance arrival on its rapidly warming Arctic breeding grounds. As egg laying has advanced much less than arrival, they still encounter a phenological mismatch that reduces offspring survival. A shift toward using more local resources for reproduction suggests that geese first need to refuel body stores at the breeding grounds after accelerated migration. Although flexibility in body store use allows migrants to accelerate migration, this cannot solve the time constraint they are facing under climate warming.

Rapid climate warming is driving organisms to advance timing of reproduction with earlier springs, but the rate of advancement shows large variation, even among populations of the same species. In this study, we investigated how the rate of advancement in timing of reproduction with a warming climate varies for barnacle goose (Branta leucopsis) populations breeding at different latitudes in the Arctic. We hypothesized that populations breeding further North are generally more time constrained and, therefore, produce clutches earlier relative to the onset of spring than southern populations. Therefore, with increasing temperatures and a progressive relief of time constraint, we expected latitudinal differences to decrease. For the years 2000–2016, we determined the onset of spring from snow cover data derived from satellite images, and compiled data on egg laying date and reproductive performance in one low-Arctic and two high-Arctic sites. As expected, high-Arctic geese laid their eggs earlier relative to snowmelt than low-Arctic geese. Contrary to expectations, advancement in laying dates was similar in high- and low-Arctic colonies, at a rate of 27% of the advance in date of snowmelt. Although advancement of egg laying did not fully compensate for the advancement of snowmelt, geese laying eggs at intermediate dates in the low Arctic were the most successful breeders. In the high Arctic, however, early nesting geese were the most successful breeders, suggesting that high-Arctic geese have not advanced their laying dates sufficiently to earlier springs. This indicates that high-Arctic geese especially are vulnerable to negative effects of climate warming.Electronic supplementary materialThe online version of this article (10.1007/s00442-019-04533-7) contains supplementary material, which is available to authorized users.

Breeding output of geese, measured as the proportion of juveniles in autumn or winter flocks, is lower in years with a late onset of spring in some species, but higher in at least one other species. Here we argue that this is because the timing of spring affects different stages of the reproductive cycle differently in different species. Because the effects on 2 different stages are opposite, the combined effects can result in either a positive or a negative overall effect. These stages are the pre-laying, laying, and nesting phase on the one hand; and the hatchling, fledgling, and juvenile phase on the other hand. The first phase is predominantly positively affected by an early snowmelt, with higher breeding propensity, clutch size, and nest success. The second phase in contrast is negatively affected by early snowmelt because of a mismatch with a nutrient food peak, leading to slow gosling growth and reduced survival. We argue that recognition of this chain of events is crucial when one wants to predict goose productivity and eventually goose population dynamics. In a rapidly warming Arctic, the negative effects of a mismatch might become increasingly important.

1. The postnatal growth period is a crucial life stage, with potential lifelong effects on an animal's fitness. How fast animals grow depends on their life-history strategy and rearing environment, and interspecific comparisons generally show higher growth rates at higher latitudes. However, to elucidate the mechanisms behind this gradient in growth rate, intraspecific comparisons are needed. Recently, barnacle geese expanded their Arctic breeding range from the RussianBarents Sea coast southwards, and now also breed along the Baltic and North Sea coasts. Baltic breeders shortened their migration, while barnacle geese breeding along the North Sea stopped migrating entirely.3. We collected cross-sectional data on gosling tarsus length, head length and body mass, and constructed population-specific growth curves to compare growth rates among three populations (Barents Sea, Baltic Sea and North Sea) spanning 17° in latitude. 4. Growth rate was faster at higher latitudes, and the gradient resembled the latitudinal gradient previously observed in an interspecific comparison of precocial species. Differences in day length among the three breeding regions could largely explain the observed differences in growth rate. In the Baltic, and especially in the Arctic population, growth rate was slower later in the season, most likely because of the stronger seasonal decline in food quality. 5. Our results suggest that differences in postnatal growth rate between the Arctic and temperate populations are mainly a plastic response to local environmental conditions. This plasticity can increase the individuals' ability to cope with annual variation in local conditions, but can also increase the potential to redistribute and adapt to new breeding environments.