More management, less damage? With increasing population size, economic costs of managing geese to minimize yield losses may outweigh benefits.

Conflicts between farmers and geese are intensifying; yet, it remains unclear how interactions between goose population size and management regimes affect yield loss and economic costs. We investigate the cost-effectiveness of accommodation and scaring areas in relation to barnacle goose (Branta leucopsis) population size. We use an existing individual-based model of barnacle geese foraging in nature, accommodation, and scaring areas in Friesland, the Netherlands, to study the most cost-effective management under varying population sizes (i.e., between 20 and 200% of the current size). Our study shows that population size non-linearly affects yield loss costs and total costs per goose. The most cost-effective management scenario for intermediate to large populations is to avoid scaring of geese. For small populations, intensive scaring resulted in minimized yield loss costs and total costs, but also substantially lower goose body mass. Our results strongly suggest that scaring becomes a less effective management measure as goose populations increase.

Migratory swans individually adjust their autumn migration and winter range to a warming climate.

In response to climate warming, migratory animals can alter their migration so that different events in the annual cycle are better aligned in space and time with suitable environmental conditions. Although such responses have been studied extensively during spring migration and the breeding season, much less is known about the influence of temperature on movements throughout autumn migration and how those movements result in a winter range and shifts therein. We use multi-year GPS tracking data to quantify how daily autumn movement and annual winter distance from the breeding grounds are related to temperature in the Western Palearctic Bewick's swan, a long-lived migratory waterbird whose winter range has shifted more than 350 km closer to the breeding grounds since 1970 due to individuals increasingly 'short-stopping' their autumn migration. We show that the migratory movement of swans is driven by lower temperatures throughout the autumn season, with individuals during late autumn moving only substantially when temperatures drop below freezing. As a result, there is large flexibility in their annual winter distance as a response to winter temperature. On average, individuals overwinter 118 km closer to the breeding grounds per 1°C increase in mean December-January temperature. Given the observed temperature increase in the Bewick's swan winter range during the last decades, our results imply that the observed range shift is for a substantial part driven by individual responses to a warming climate. We thus present an example of individual flexibility towards climatic conditions driving the range shift of a migratory species. Our study adds to the understanding of the processes that shape autumn migration decisions, winter ranges and shifts therein, which is crucial to be able to predict how climate change may impact these processes in the future.

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.

Rapid formation of new migration route and breeding area by Arctic geese.

Many Arctic-breeding animals are at risk from local extirpation associated with habitat constriction and alterations in phenology in their Arctic environment as a result of rapid global warming.1 Migratory species face additional increasing anthropogenic pressures along their migratory routes such as habitat destruction, droughts, creation of barriers, and overexploitation.2,3 Such species can only persist if they adjust their migration, timing of breeding, and range.4 Here, we document both the abrupt (∼10 years) formation of a new migration route and a disjunct breeding population of the pink-footed goose (Anser brachyrhynchus) on Novaya Zemlya, Russia, almost 1,000 km away from the original breeding grounds in Svalbard. The population has grown to 3,000-4,000 birds, explained by intrinsic growth and continued immigration from the original route. The colonization was enabled by recent warming on Novaya Zemlya. We propose that social behavior of geese, resulting in cultural transmission of migration behavior among conspecifics as well as in mixed-species flocks, is key to this fast development and acts as a mechanism enabling ecological rescue in a rapidly changing world.

Energetic and behavioral consequences of migration: an empirical evaluation in the context of the full annual cycle.

Seasonal migrations are used by diverse animal taxa, yet the costs and benefits of migrating have rarely been empirically examined. The aim of this study was to determine how migration influences two ecological currencies, energy expenditure and time allocated towards different behaviors, in a full annual cycle context. We compare these currencies among lesser black-backed gulls that range from short- (< 250 km) to long-distance (> 4500 km) migrants. Daily time-activity budgets were reconstructed from tri-axial acceleration and GPS, which, in conjunction with a bioenergetics model to estimate thermoregulatory costs, enabled us to estimate daily energy expenditure throughout the year. We found that migration strategy had no effect on annual energy expenditure, however, energy expenditure through time deviated more from the annual average as migration distance increased. Patterns in time-activity budgets were similar across strategies, suggesting migration strategy does not limit behavioral adjustments required for other annual cycle stages (breeding, molt, wintering). Variation among individuals using the same strategy was high, suggesting that daily behavioral decisions (e.g. foraging strategy) contribute more towards energy expenditure than an individual's migration strategy. These findings provide unprecedented new understanding regarding the relative importance of fine versus broad-scale behavioral strategies towards annual energy expenditures.

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.

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.

Concurrent shifts in wintering distribution and phenology in migratory swans: Individual and generational effects.

Range shifts and phenological change are two processes by which organisms respond to environmental warming. Understanding the mechanisms that drive these changes is key for optimal conservation and management. Here we study both processes in the migratory Bewick's swan (Cygnus columbianus bewickii) using different methods, analysing nearly 50 years of resighting data (1970-2017). In this period the wintering area of the Bewick's swans shifted eastwards ('short-stopping') at a rate of ~13 km/year, thereby shortening individual migration distance on an average by 353 km. Concurrently, the time spent at the wintering grounds has reduced ('short-staying') by ~38 days since 1989. We show that individuals are consistent in their migratory timing in winter, indicating that the frequency of individuals with different migratory schedules has changed over time (a generational shift). In contrast, for short-stopping we found evidence for both individual plasticity (individuals decrease their migration distances over their lifetime) and generational shift. Additional analysis of swan resightings with temperature data showed that, throughout the winter, Bewick's swans frequent areas where air temperatures are c. 5.5°C. These areas have also shifted eastwards over time, hinting that climate warming is a contributing factor behind the observed changes in the swans' distribution. The occurrence of winter short-stopping and short-staying suggests that this species is to some extent able to adjust to climate warming, but benefits or repercussions at other times of the annual cycle need to be assessed. Furthermore, these phenomena could lead to changes in abundance in certain areas, with resulting monitoring and conservation implications. Understanding the processes and driving mechanisms behind population changes therefore is important for population management, both locally and across the species range.

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.

Less is more: On-board lossy compression of accelerometer data increases biologging capacity.

GPS-tracking devices have been used in combination with a wide range of additional sensors to study animal behaviour, physiology and interaction with their environment. Tri-axial accelerometers allow researchers to remotely infer the behaviour of individuals, at all places and times. Collection of accelerometer data is relatively cheap in terms of energy usage, but the amount of raw data collected generally requires much storage space and is particularly demanding in terms of energy needed for data transmission. Here, we propose compressing the raw accelerometer (ACC) data into summary statistics within the tracking device (before transmission) to reduce data size, as a means to overcome limitations in storage and energy capacity. We explored this type of lossy data compression in the accelerometer data of tagged Bewick's swans Cygnus columbianus bewickii collected in spring 2017. Using software settings in which bouts of 2 s of both raw ACC data and summary statistics were collected in parallel but with different bout intervals to keep total data size comparable, we created the opportunity for a direct comparison of time budgets derived by the two data collection methods. We found that the data compression in our case yielded a six times reduction in data size per bout, and concurrent, similar decreases in storage and energy use of the device. We show that with the same accuracy of the behavioural classification, the freed memory and energy of the device can be used to increase the monitoring effort, resulting in a more detailed representation of the individuals' time budget. Rare and/or short behaviours, such as daily roost flights, were picked up significantly more when collecting summary statistics instead of raw ACC data (but note differences in sampling rate). Such level of detail can be of essential importance, for instance to make a reliable estimate of the energy budgets of individuals. In conclusion, we argue that this type of lossy data compression can be a well-considered choice in study situations where limitations in energy and storage space of the device pose a problem. Ultimately, these developments can allow for long-term and nearly continuous remote monitoring of the behaviour of free-ranging animals.

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.

A mechanistic assessment of the relationship between gut morphology and endozoochorous seed dispersal by waterfowl.

Many plants and invertebrates rely on internal transport by animals for long-distance dispersal. Their dispersal capacity is greatly influenced by interactions with the animal's digestive tract. Omnivorous birds adjust their digestive tract morphology to seasonally variable diets. We performed feeding trials in waterfowl to unravel how changing organ size, in combination with seed size, affects dispersal potential. We subjected captive mallards to mimics of summer (animal-based), winter (plant-based), and intermediate diets, and analyzed gut passage of seeds before and after the treatment (trials 1 and 2). To test the effect of gut morphology on seed digestion, we measured digestive organ size after euthanasia. Three hours before euthanasia, differently sized seeds were fed to test how seed size affects gut passage by determining their relative position in the digestive tract (trial 3). Trials 1 and 2 showed that intact seed passage was lower in the plant-based than in the animal-based diet group. Retention time changed only within groups, decreasing in the animal-based, and increasing in the plant-based diet group. No post-diet differences in organ size were detected, probably due to large between-individual variation within groups. Digestive tract measures did not explain variation in seed survival or retention time. Trial 3 revealed that small seeds pass the digestive tract rapidly, while large seeds are retained longer, particularly in the gizzard. Differential retention in the gizzard, the section where seeds can be destroyed, is likely why larger seeds have a lower probability to pass the digestive tract intact. Our results confirm that rapid, flexible adaptation to diet shifts affects seed digestion in waterfowl, although we could not conclusively relate this to organ size. Large interindividual variation in digestive efficiency between mallards feeding on the same diet may provide opportunities for seed dispersal in the field throughout the annual cycle.

The roles of migratory and resident birds in local avian influenza infection dynamics.

Migratory birds are an increasing focus of interest when it comes to infection dynamics and the spread of avian influenza viruses (AIV). However, we lack detailed understanding migratory birds' contribution to local AIV prevalence levels and their downstream socio-economic costs and threats.To explain the potential differential roles of migratory and resident birds in local AIV infection dynamics, we used a susceptible-infectious-recovered (SIR) model. We investigated five (mutually non- exclusive) mechanisms potentially driving observed prevalence patterns: 1) a pronounced birth pulse (e.g. the synchronised annual influx of immunologically naïve individuals), 2) short-term immunity, 3) increase of susceptible migrants, 4) differential susceptibility to infection (i.e. transmission rate) for migrants and residents, and 5) replacement of migrants during peak migration.SIR models describing all possible combinations of the five mechanisms were fitted to individual AIV infection data from a detailed longitudinal surveillance study in the partially migratory mallard duck (Anas platyrhynchos). During autumn and winter, the local resident mallard community also held migratory mallards that exhibited distinct AIV infection dynamics.Replacement of migratory birds during peak migration in autumn was found to be the most important mechanism driving the variation in local AIV infection patterns. This suggests that a constant influx of migratory birds, likely immunological naïve to locally circulating AIV strains, is required to predict the observed temporal prevalence patterns and the distinct differences in prevalence between residents and migrants.Synthesis and applications. Our analysis reveals a key mechanism that could explain the amplifying role of migratory birds in local avian influenza virus infection dynamics; the constant flow and replacement of migratory birds during peak migration. Aside from monitoring efforts, in order to achieve adequate disease management and control in wildlife - with knock-on effects for livestock and humans, - we conclude that it is crucial, in future surveillance studies, to record host demographical parameters such as population density, timing of birth and turnover of migrants.

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.

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.

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.

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.

Combining modelling tools to evaluate a goose management scheme.

Many goose species feed on agricultural land, and with growing goose numbers, conflicts with agriculture are increasing. One possible solution is to designate refuge areas where farmers are paid to leave geese undisturbed. Here, we present a generic modelling tool that can be used to designate the best locations for refuges and to gauge the area needed to accommodate the geese. With a species distribution model, locations are ranked according to goose suitability. The size of the area to be designated as refuge can be chosen by including more or less suitable locations. A resource depletion model is then used to estimate whether enough resources are available within the designated refuge to accommodate all geese, taking into account the dynamics of food resources, including depletion by geese. We illustrate this with the management scheme for pink-footed goose Anser brachyrhynchus implemented in Norway. Here, all geese can be accommodated, but damage levels appear to depend on weather, land use and refuge size.

Movement patterns of a keystone waterbird species are highly predictable from landscape configuration.

BACKGROUND: Movement behaviour is fundamental to the ecology of animals and their interactions with other organisms, and as such contributes to ecosystem dynamics. Waterfowl are key players in ecological processes in wetlands and surrounding habitats through predator-prey interactions and their transportation of nutrients and other organisms. Understanding the drivers of their movement behaviour is crucial to predict how environmental changes affect their role in ecosystem functioning. Mallards (Anas platyrhynchos) are the most abundant duck species worldwide and important dispersers of aquatic invertebrates, plants and pathogens like avian influenza viruses. By GPS tracking of 97 mallards in four landscape types along a gradient of wetland availability, we identified patterns in their daily movement behaviour and quantified potential effects of weather conditions and water availability on the spatial scale of their movements. RESULTS: We demonstrate that mallard movement patterns were highly predictable, with regular commuting flights at dusk and dawn between a fixed day roost and one or several fixed nocturnal foraging sites, linked strongly to surface water. Wind and precipitation hardly affected movement, but flight distances and home range sizes increased when temperatures dropped towards zero. Flight distances and home range sizes increased exponentially with decreasing availability of freshwater habitat. Total shoreline length and the number of water bodies in the landscape surrounding the roost were the best predictors of the spatial scale of daily mallard movements. CONCLUSIONS: Our results show how mallards may flexibly adjust the spatial scale of their movements to wetland availability in the landscape. This implies that mallards moving between discrete habitat patches continue to preserve biotic connectivity in increasingly fragmented landscapes. The high predictability of mallard movement behaviour in relation to landscape features makes them reliable dispersal vectors for organisms to adapt to, and allows prediction of their ecological role in other landscapes.