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.

Environmental refuges from disease in host-parasite interactions under global change.

The physiological performance of organisms depends on their environmental context, resulting in performance-response curves along environmental gradients. Parasite performance-response curves are generally expected to be broader than those of their hosts due to shorter generation times and hence faster adaptation. However, certain environmental conditions may limit parasite performance more than that of the host, thereby providing an environmental refuge from disease. Thermal disease refuges have been extensively studied in response to climate warming, but other environmental factors may also provide environmental disease refuges which, in turn, respond to global change. Here, we (1) showcase laboratory and natural examples of refuges from parasites along various environmental gradients, and (2) provide hypotheses on how global environmental change may affect these refuges. We strive to synthesize knowledge on potential environmental disease refuges along different environmental gradients including salinity and nutrients, in both natural and food-production systems. Although scaling up from single host-parasite relationships along one environmental gradient to their interaction outcome in the full complexity of natural environments remains difficult, integrating host and parasite performance-response can serve to formulate testable hypotheses about the variability in parasitism outcomes and the occurrence of environmental disease refuges under current and future environmental conditions.

Polygamy and purifying selection in birds.

Good genes theories of sexual selection predict that polygamy will be associated with more efficient removal of deleterious alleles (purifying selection), due to the alignment of sexual selection with natural selection. On the other hand, runaway selection theories expect no such alignment of natural and sexual selection, and may instead predict less efficient purifying selection in polygamous species due to higher reproductive variance. In an analysis of polymorphism data extracted from 150-bird genome assemblies, we show that polygamous species carry significantly fewer nonsynonymous polymorphisms, relative to synonymous polymorphisms, than monogamous bird species (p = .0005). We also show that this effect is independent of effective population size, consistent with the alignment of natural selection with sexual selection and "good genes" theories of sexual selection. Further analyses found no impact of polygamy on genetic diversity, while polygamy in females (polyandry) had a marginally significant impact (p = .045). We also recapitulate previous findings that smaller body mass and greater geographic range size are associated with more efficient purifying selection, more intense GC-biased gene conversion, and greater genetic diversity.

Postnatal growth rate varies with latitude in range-expanding geese: The role of plasticity and day length.

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 Russian Barents 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. 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. 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. 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 re-distribute and adapt to new breeding environments.

Seasonal variation in sex-specific immunity in wild birds.

Whilst the immune system often varies seasonally and exhibits differences between males and females, the general patterns in seasonality and sex differences across taxa have remained controversial. Birds are excellent model organisms to assess these patterns, because the immune system of many species is well characterised. We conducted a meta-analysis using 41 wild bird species from 24 avian families to investigate sex differences and seasonal (breeding/non-breeding) variations in immune status, including white blood cell counts, phytohaemagglutinin (PHA) test, bacteria-killing ability (BKA), haemolysis and haemagglutination assays. We found male-biased macrophage concentration, BKA and haemolysis titers, but only during the breeding season. Sex-specific heterophil concentrations, heterophil/lymphocyte ratios and PHA responses differed between breeding and non-breeding, suggesting larger changes in males than in females. Importantly, sex differences in immune status are stronger during the breeding period than during the non-breeding period. Taken together, our study suggests that both seasonal variation and sex differences in immune system are common in birds, although their associations are more complex than previously thought.

Ecological insights from three decades of animal movement tracking across a changing Arctic.

The Arctic is entering a new ecological state, with alarming consequences for humanity. Animal-borne sensors offer a window into these changes. Although substantial animal tracking data from the Arctic and subarctic exist, most are difficult to discover and access. Here, we present the new Arctic Animal Movement Archive (AAMA), a growing collection of more than 200 standardized terrestrial and marine animal tracking studies from 1991 to the present. The AAMA supports public data discovery, preserves fundamental baseline data for the future, and facilitates efficient, collaborative data analysis. With AAMA-based case studies, we document climatic influences on the migration phenology of eagles, geographic differences in the adaptive response of caribou reproductive phenology to climate change, and species-specific changes in terrestrial mammal movement rates in response to increasing temperature.

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.

Heterothermy in growing king penguins.

A drop in body temperature allows significant energy savings in endotherms, but facultative heterothermy is usually restricted to small animals. Here we report that king penguin chicks (Aptenodytes patagonicus), which are able to fast for up to 5 months in winter, undergo marked seasonal heterothermy during this period of general food scarcity and slow-down of growth. They also experience short-term heterothermy below 20 °C in the lower abdomen during the intense (re)feeding period in spring, induced by cold meals and adverse weather. The heterothermic response involves reductions in peripheral temperature, reductions in thermal core volume and temporal abandonment of high core temperature. Among climate variables, air temperature and wind speed show the strongest effect on body temperature, but their effect size depends on physiological state. The observed heterothermy is remarkable for such a large bird (10 kg before fasting), which may account for its unrivalled fasting capacity among birds.

Predation danger can explain changes in timing of migration: the case of the barnacle goose.

Understanding stopover decisions of long-distance migratory birds is crucial for conservation and management of these species along their migratory flyway. Recently, an increasing number of Barnacle geese breeding in the Russian Arctic have delayed their departure from their wintering site in The Netherlands by approximately one month and have reduced their staging duration at stopover sites in the Baltic accordingly. Consequently, this extended stay increases agricultural damage in The Netherlands. Using a dynamic state variable approach we explored three hypotheses about the underlying causes of these changes in migratory behavior, possibly related to changes in (i) onset of spring, (ii) potential intake rates and (iii) predation danger at wintering and stopover sites. Our simulations showed that the observed advance in onset of spring contradicts the observed delay of departure, whereas both increased predation danger and decreased intake rates in the Baltic can explain the delay. Decreased intake rates are expected as a result of increased competition for food in the growing Barnacle goose population. However, the effect of predation danger in the model was particularly strong, and we hypothesize that Barnacle geese avoid Baltic stopover sites as a response to the rapidly increasing number of avian predators in the area. Therefore, danger should be considered as an important factor influencing Barnacle goose migratory behavior, and receive more attention in empirical studies.

Skipping the Baltic: the emergence of a dichotomy of alternative spring migration strategies in Russian barnacle geese.

1. Since the early 1990s, an increasing proportion of barnacle geese, Branta leucopsis, bound for breeding sites in the Russian Arctic delay their departure from the wintering quarters in the Wadden Sea by 4 weeks. These late-migrating geese skip spring stopover sites in the Baltic traditionally used by the entire population. 2. Individual geese from an arctic colony tracked by satellite or light-level geolocators during spring migration in 2004 and 2005 predominantly followed the new strategy, but a minority still maintained the traditional pattern. Despite a spread of more than 50 days in departure date from the Wadden Sea, both early and late departing females laid their eggs within the short time-window conferring breeding success. 3. The spread of these new migration routines coincided with a strong increase of overall numbers and the exploitation of new spring staging resources in the Wadden Sea. Counts from Estonia demonstrate that numbers have levelled off recently at the Baltic staging sites, suggesting that the capacity of these staging sites in spring has been reached. Although onset of spring affects migratory timing in barnacle geese, it cannot explain the observed delay in departure from the wintering grounds. 4. We hypothesize that the new migratory strategy has evolved in response to increased competition for food at spring staging sites in the Baltic. According to an analytical model of optimal migration, the geese should skip the Baltic whenever the energy deposition rate falls below 88% of the Wadden Sea value.

Evaluation of the deuterium dilution method to estimate body composition in the barnacle goose: accuracy and minimum equilibration time.

We examined body composition in barnacle geese (Branta leucopsis) by proximate carcass analysis and by deuterium isotope dilution. We studied the effect of isotope equilibration time on the accuracy of total body water (TBW) estimates and evaluated models to predict fat-free mass (FFM) and fat mass (FM) from different measurements varying in their level of invasiveness. Deuterium enrichment determined at 45, 90, and 180 min after isotope injection did not differ significantly. At all sampling intervals, isotope dilution spaces (TBW(d)) consistently overestimated body water determined by carcass analysis (TBW(c)). However, variance in the deviation from actual TBW was higher at the 45-min sampling interval, whereas variability was the same at 90 and 180 min, indicating that 90 min is sufficient time to allow for adequate equilibration. At 90 min equilibration time, deuterium isotope dilution overestimated TBW(c) by 7.1% +/= 2.6% (P < 0.001, paired t-test, n=20). This overestimate was consistent over the range of TBW studied, and TBW(c) could thus be predicted from TBW(d) (r2=0.976, P<0.001). Variation in TBW(c) and TBW(d) explained, respectively, 99% and 98% of the variation in FFM. FM could be predicted with a relative error of ca. 10% from TBW estimates in combination with body mass (BM). In contrast, BM and external body measurements allowed only poor prediction. Abdominal fat fresh mass was highly correlated to total FM and, if the carcass is available, allows simple means of fat prediction without dissecting the entire specimen.