Earlier springs increase goose breeding propensity and nesting success at Arctic but not at temperate latitudes.

Intermittent breeding is an important tactic in long-lived species that trade off survival and reproduction to maximize lifetime reproductive success. When breeding conditions are unfavourable, individuals are expected to skip reproduction to ensure their own survival. Breeding propensity (i.e. the probability for a mature female to breed in a given year) is an essential parameter in determining reproductive output and population dynamics, but is not often studied in birds because it is difficult to obtain unbiased estimates. Breeding conditions are especially variable at high latitudes, potentially resulting in a large effect on breeding propensity of Arctic-breeding migratory birds, such as geese. With a novel approach, we used GPS-tracking data to determine nest locations, breeding propensity and nesting success of barnacle geese, and studied how these varied with breeding latitude and timing of arrival on the breeding grounds relative to local onset of spring. Onset of spring at the breeding grounds was a better predictor of breeding propensity and nesting success than relative timing of arrival. At Arctic latitudes (>66° N), breeding propensity decreased from 0.89 (95% CI: 0.65-0.97) in early springs to 0.22 (95% CI: 0.06-0.55) in late springs, while at temperate latitudes, it varied between 0.75 (95% CI: 0.38-0.93) and 0.89 (95% CI: 0.41-0.99) regardless of spring phenology. Nesting success followed a similar pattern and was lower in later springs at Arctic latitudes, but not at temperate latitudes. In early springs, a larger proportion of geese started breeding despite arriving late relative to the onset of spring, possibly because the early spring enabled them to use local resources to fuel egg laying and incubation. While earlier springs due to climate warming are considered to have mostly negative repercussions on reproductive success through phenological mismatches, our results suggest that these effects may partly be offset by higher breeding propensity and nesting success.

Year-round activity levels reveal diurnal foraging constraints in the annual cycle of migratory and non-migratory barnacle geese.

Performing migratory journeys comes with energetic costs, which have to be compensated within the annual cycle. An assessment of how and when such compensation occurs is ideally done by comparing full annual cycles of migratory and non-migratory individuals of the same species, which is rarely achieved. We studied free-living migratory and resident barnacle geese belonging to the same flyway (metapopulation), and investigated when differences in foraging activity occur, and when foraging extends beyond available daylight, indicating a diurnal foraging constraint in these usually diurnal animals. We compared foraging activity of migratory (N = 94) and resident (N = 30) geese throughout the annual cycle using GPS-transmitters and 3D-accelerometers, and corroborated this with data on seasonal variation in body condition. Migratory geese were more active than residents during most of the year, amounting to a difference of over 370 h over an entire annual cycle. Activity differences were largest during the periods that comprised preparation for spring and autumn migration. Lengthening days during spring facilitated increased activity, which coincided with an increase in body condition. Both migratory and resident geese were active at night during winter, but migratory geese were also active at night before autumn migration, resulting in a period of night-time activity that was 6 weeks longer than in resident geese. Our results indicate that, at least in geese, seasonal migration requires longer daily activity not only during migration but throughout most of the annual cycle, with migrants being more frequently forced to extend foraging activity into the night.

Mismatch-induced growth reductions in a clade of Arctic-breeding shorebirds are rarely mitigated by increasing temperatures.

In seasonal environments subject to climate change, organisms typically show phenological changes. As these changes are usually stronger in organisms at lower trophic levels than those at higher trophic levels, mismatches between consumers and their prey may occur during the consumers' reproduction period. While in some species a trophic mismatch induces reductions in offspring growth, this is not always the case. This variation may be caused by the relative strength of the mismatch, or by mitigating factors like increased temperature-reducing energetic costs. We investigated the response of chick growth rate to arthropod abundance and temperature for six populations of ecologically similar shorebirds breeding in the Arctic and sub-Arctic (four subspecies of Red Knot Calidris canutus, Great Knot C. tenuirostris and Surfbird C. virgata). In general, chicks experienced growth benefits (measured as a condition index) when hatching before the seasonal peak in arthropod abundance, and growth reductions when hatching after the peak. The moment in the season at which growth reductions occurred varied between populations, likely depending on whether food was limiting growth before or after the peak. Higher temperatures led to faster growth on average, but could only compensate for increasing trophic mismatch for the population experiencing the coldest conditions. We did not find changes in the timing of peaks in arthropod availability across the study years, possibly because our series of observations was relatively short; timing of hatching displayed no change over the years either. Our results suggest that a trend in trophic mismatches may not yet be evident; however, we show Arctic-breeding shorebirds to be vulnerable to this phenomenon and vulnerability to depend on seasonal prey dynamics.

Ontogenetic niche shifts as a driver of seasonal migration.

Ontogenetic niche shifts have helped to understand population dynamics. Here we show that ontogenetic niche shifts also offer an explanation, complementary to traditional concepts, as to why certain species show seasonal migration. We describe how demographic processes (survival, reproduction and migration) and associated ecological requirements of species may change with ontogenetic stage (juvenile, adult) and across the migratory range (breeding, non-breeding). We apply this concept to widely different species (dark-bellied brent geese (Branta b. bernicla), humpback whales (Megaptera novaeangliae) and migratory Pacific salmon (Oncorhynchus gorbuscha) to check the generality of this hypothesis. Consistent with the idea that ontogenetic niche shifts are an important driver of seasonal migration, we find that growth and survival of juvenile life stages profit most from ecological conditions that are specific to breeding areas. We suggest that matrix population modelling techniques are promising to detect the importance of the ontogenetic niche shifts in maintaining migratory strategies. As a proof of concept, we applied a first analysis to resident, partial migratory and fully migratory populations of barnacle geese (Branta leucopsis). We argue that recognition of the costs and benefits of migration, and how these vary with life stages, is important to understand and conserve migration under global environmental change.

Climate warming may affect the optimal timing of reproduction for migratory geese differently in the low and high Arctic.

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.

Potential for an Arctic-breeding migratory bird to adjust spring migration phenology to Arctic amplification.

Arctic amplification, the accelerated climate warming in the polar regions, is causing a more rapid advancement of the onset of spring in the Arctic than in temperate regions. Consequently, the arrival of many migratory birds in the Arctic is thought to become increasingly mismatched with the onset of local spring, consequently reducing individual fitness and potentially even population levels. We used a dynamic state variable model to study whether Arctic long-distance migrants can advance their migratory schedules under climate warming scenarios which include Arctic amplification, and whether such an advancement is constrained by fuel accumulation or the ability to anticipate climatic changes. Our model predicts that barnacle geese Branta leucopsis suffer from considerably reduced reproductive success with increasing Arctic amplification through mistimed arrival, when they cannot anticipate a more rapid progress of Arctic spring from their wintering grounds. When geese are able to anticipate a more rapid progress of Arctic spring, they are predicted to advance their spring arrival under Arctic amplification up to 44 days without any reproductive costs in terms of optimal condition or timing of breeding. Negative effects of mistimed arrival on reproduction are predicted to be somewhat mitigated by increasing summer length under warming in the Arctic, as late arriving geese can still breed successfully. We conclude that adaptation to Arctic amplification may rather be constrained by the (un)predictability of changes in the Arctic spring than by the time available for fuel accumulation. Social migrants like geese tend to have a high behavioural plasticity regarding stopover site choice and migration schedule, giving them the potential to adapt to future climate changes on their flyway.

Exploring the drivers of variation in trophic mismatches: A systematic review of long-term avian studies.

Many organisms reproduce in seasonal environments, where selection on timing of reproduction is particularly strong as consumers need to synchronize reproduction with the peaked occurrence of their food. When a consumer species changes its phenology at a slower rate than its resources, this may induce a trophic mismatch, that is, offspring growing up after the peak in food availability, potentially leading to reductions in growth and survival. However, there is large variation in the degree of trophic mismatches as well as in its effects on reproductive output.Here, we explore the potential causes for variation in the strength of trophic mismatches in published studies of birds. Specifically, we ask whether the changes in the degree of mismatch that have occurred over time can be explained by a bird's (a) breeding latitude, (b) migration distance, and/or (c) life-history traits.We found that none of these three factors explain changes in the degree of mismatch over time. Nevertheless, food phenology did advance faster at more northerly latitudes, while shifts in bird phenology did not show a trend with latitude.We argue that the lack of support in our results is attributable to the large variation in the metrics used to describe timing of food availability. We propose a pathway to improve the quantification of trophic mismatches, guided by a more rigorous understanding of links between consumers and their resources.

Nocturnal foraging lifts time constraints in winter for migratory geese but hardly speeds up fueling.

Climate warming advances the optimal timing of breeding for many animals. For migrants to start breeding earlier, a concurrent advancement of migration is required, including premigratory fueling of energy reserves. We investigate whether barnacle geese are time constrained during premigratory fueling and whether there is potential to advance or shorten the fueling period to allow an earlier migratory departure. We equipped barnacle geese with GPS trackers and accelerometers to remotely record birds' behavior, from which we calculated time budgets. We examined how time spent foraging was affected by the available time (during daylight and moonlit nights) and thermoregulation costs. We used an energetic model to assess onset and rates of fueling and whether geese can further advance fueling by extending foraging time. We show that, during winter, when facing higher thermoregulation costs, geese consistently foraged at night, especially during moonlit nights, in order to balance their energy budgets. In spring, birds made use of the increasing day length and gained body stores by foraging longer during the day, but birds stopped foraging extensively during the night. Our model indicates that, by continuing nighttime foraging throughout spring, geese may have some leeway to advance and increase fueling rate, potentially reaching departure body mass 4 days earlier. In light of rapid climatic changes on the breeding grounds, whether this advancement can be realized and whether it will be sufficient to prevent phenological mismatches remains to be determined.

Arctic Geese Tune Migration to a Warming Climate but Still Suffer from a Phenological Mismatch.

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.

Forage plants of an Arctic-nesting herbivore show larger warming response in breeding than wintering grounds, potentially disrupting migration phenology.

During spring migration, herbivorous waterfowl breeding in the Arctic depend on peaks in the supply of nitrogen-rich forage plants, following a "green wave" of grass growth along their flyway to fuel migration and reproduction. The effects of climate warming on forage plant growth are expected to be larger at the Arctic breeding grounds than in temperate wintering grounds, potentially disrupting this green wave and causing waterfowl to mistime their arrival on the breeding grounds. We studied the potential effect of climate warming on timing of food peaks along the migratory flyway of the Russian population of barnacle geese using a warming experiment with open-top chambers. We measured the effect of 1.0-1.7°C experimental warming on forage plant biomass and nitrogen concentration at three sites along the migratory flyway (temperate wintering site, temperate spring stopover site, and Arctic breeding site) during 2 months for two consecutive years. We found that experimental warming increased biomass accumulation and sped up the decline in nitrogen concentration of forage plants at the Arctic breeding site but not at temperate wintering and stop-over sites. Increasing spring temperatures in the Arctic will thus shorten the food peak of nitrogen-rich forage at the breeding grounds. Our results further suggest an advance of the local food peak in the Arctic under 1-2°C climate warming, which will likely cause migrating geese to mistime their arrival at the breeding grounds, particularly considering the Arctic warms faster than the temperate regions. The combination of a shorter food peak and mistimed arrival is likely to decrease goose reproductive success under climate warming by reducing growth and survival of goslings after hatching.

Analyzing time-ordered event data with missed observations.

A common problem with observational datasets is that not all events of interest may be detected. For example, observing animals in the wild can difficult when animals move, hide, or cannot be closely approached. We consider time series of events recorded in conditions where events are occasionally missed by observers or observational devices. These time series are not restricted to behavioral protocols, but can be any cyclic or recurring process where discrete outcomes are observed. Undetected events cause biased inferences on the process of interest, and statistical analyses are needed that can identify and correct the compromised detection processes. Missed observations in time series lead to observed time intervals between events at multiples of the true inter-event time, which conveys information on their detection probability. We derive the theoretical probability density function for observed intervals between events that includes a probability of missed detection. Methodology and software tools are provided for analysis of event data with potential observation bias and its removal. The methodology was applied to simulation data and a case study of defecation rate estimation in geese, which is commonly used to estimate their digestive throughput and energetic uptake, or to calculate goose usage of a feeding site from dropping density. Simulations indicate that at a moderate chance to miss arrival events (p = 0.3), uncorrected arrival intervals were biased upward by up to a factor 3, while parameter values corrected for missed observations were within 1% of their true simulated value. A field case study shows that not accounting for missed observations leads to substantial underestimates of the true defecation rate in geese, and spurious rate differences between sites, which are introduced by differences in observational conditions. These results show that the derived methodology can be used to effectively remove observational biases in time-ordered event data.

A century of ecosystem change: human and seabird impacts on plant species extirpation and invasion on islands.

We used 116 years of floral and faunal records from Mandarte Island, British Columbia, Canada, to estimate the indirect effects of humans on plant communities via their effects on the population size of a surface-nesting, colonial seabird, the Glaucous-winged gull (Larus glaucescens). Comparing current to historical records revealed 18 extirpations of native plant species (32% of species historically present), 31 exotic species introductions, and one case of exotic introduction followed by extirpation. Contemporary surveys indicated that native species cover declined dramatically from 1986 to 2006, coincident with the extirpation of 'old-growth' conifers. Because vegetation change co-occurred with an increasing gull population locally and regionally, we tested several predictions from the hypothesis that the presence and activities of seabirds help to explain those changes. Specifically, we predicted that on Mandarte and nearby islands with gull colonies, we should observe higher nutrient loading and exotic plant species richness and cover than on nearby islands without gull colonies, as a consequence of competitive dominance in species adapted to high soil nitrogen and trampling. As predicted, we found that native plant species cover and richness were lower, and exotic species cover and richness higher, on islands with versus without gull colonies. In addition, we found that soil carbon and nitrogen on islands with nesting gulls were positively related to soil depth and exotic species richness and cover across plots and islands. Our results support earlier suggestions that nesting seabirds can drive rapid change in insular plant communities by increasing nutrients and disturbing vegetation, and that human activities that affect seabird abundance may therefore indirectly affect plant community composition on islands with seabird colonies.