Bart A. Nolet

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.

In recent decades, conflict between geese and agriculture has increased. Management practices to limit this conflict include concentrating geese in protected areas, derogation shooting or population reduction. To justify such management, we need to understand their effects on goose-related damages, which requires an understanding of how yield loss is influenced by goose abundance and species interactions.
We combined data from monthly goose counts and GPS-tracked geese to estimate grazing pressures by barnacle, white-fronted and greylag geese on agricultural grassland in Fryslân, the Netherlands. Using linear mixed models, we related this to damages assessed by professional inspectors.
Our results show a positive nonlinear relationship between yield loss and barnacle goose grazing pressure, where assessed damage increases with a decelerating rate as grazing pressure increases. For white-fronted geese, we find a negative relationship, while for greylag geese both positive and negative relationships occur. For each species, the relationship is influenced by the abundance of the other two.
For barnacle geese, the relationship can be explained by selection of fields offering the best balance between food intake and energy expenditure, and by grass regrowth, with highest grazing pressures occurring over a longer time period. The results for the other species are likely due to spatial and temporal differences in foraging preferences compared to barnacle geese, where larger species avoid areas with highest damages.
Synthesis and applications. Our results suggest that decreasing herbivore abundance may not translate directly to decreased yield loss, and management tools such as population reduction or derogation shooting should be used with care. Management aimed at concentrating geese in refuges could help to alleviate farmer–goose conflict, although further studies are required to determine if it would lead to damage reduction. We also find that not all species contribute equally to agricultural damage; care should be taken to ensure wildlife management targets the right species.

In the present Anthropocene, wild animals are globally affected by human activity. Consumer fireworks during New Year (NY) are widely distributed in W-Europe and cause strong disturbances that are known to incur stress responses in animals. We analyzed GPS tracks of 347 wild migratory geese of four species during eight NYs quantifying the effects of fireworks on individuals. We show that, in parallel with particulate matter increases, during the night of NY geese flew on average 5–16 km further and 40–150 m higher, and more often shifted to new roost sites than on previous nights. This was also true during the 2020–2021 fireworks ban, despite fireworks activity being reduced. Likely to compensate for extra flight costs, most geese moved less and increased their feeding activity in the following days. Our findings indicate negative effects of NY fireworks on wild birds beyond the previously demonstrated immediate response.

Many angiosperms rely on vertebrates for seed dispersal via gut passage, an interaction that has been traditionally classified as a mutualism. The seed dispersal effectiveness (SDE) framework provides a mechanistic approach to evaluate evolutionary and ecological characteristics of animal-mediated seed dispersal, by synthesising the quantity and the quality of the dispersal that a plant species receives from each of its animal dispersers. However, the application of the SDE framework has been largely restricted to plant–frugivore interactions, whereas animal-mediated seed dispersal results from plant–disperser interactions that cover a continuum from pure mutualisms to antagonisms. This biases ecological and evolutionary knowledge on plant–disperser interactions. Here, we propose an extended SDE framework (‘eSDE') that allows comparing plant–disperser interactions in the full mutualism–antagonism continuum ranging from pure mutualisms (frugivores) to conditional mutualisms (scatter-hoarding granivores and folivores) and antagonisms (pure granivores). We present the eSDE framework, and use examples to illustrate how it can be applied to compare effectiveness among plant–disperser interaction types. Our initial comparison based on available data suggests that vertebrate species differ more in the number of seeds they deposit away from the mother plant (quantity), than in the effects such dispersal processes have on seed fate (quality). Scatter-hoarding granivores provide the most effective dispersal due to high removal rates, closely followed by frugivores due to high deposition rates. Folivores and pure granivores provide low quantity dispersal, but of high and moderate quality, respectively. These early comparative insights illustrate the necessity and usefulness of more standardized data collection protocols, for which we provide recommendations. Applying the eSDE framework can reveal broad-scale patterns across and within plant–disperser interaction types, which will advance our evolutionary understanding of plant–animal interactions. This will provide new insights into the consequence of anthropogenic impacts on vertebrate-mediated seed dispersal in a world in which plant–animal interactions are increasingly threatened.

In animal ecology, energy expenditure is used for assessing the consequences of different behavioural strategies, life-history events or environments. Animals can also influence energy expenditure through instantaneous behavioural responses to their external environment. It is therefore of interest to measure energy expenditure of free-ranging animals across seasons and at high temporal resolutions. Heart rate has historically been used for this, but requires invasive surgery for long-term use. Dynamic body acceleration (DBA) is an alternative proxy for energy expenditure that is simpler to deploy, yet few studies have examined how it performs over extended time periods, or for species using different locomotory modes, especially passive modes like soaring flight. We measured DBA alongside heart rate in free-ranging lesser black-backed gulls, a seabird that moves using flapping flight, soaring and walking, and rests on both land and water. Our objectives were to compare the relative changes in DBA and heart rate among and within behaviours and to examine how accelerometers can be used to estimate daily energy expenditure by comparing DBA to time-energy budgets (TEBs). DBA and heart rate were sampled concurrently at 2.5- and 5-min intervals throughout the breeding season, though measurements were not exactly synchronised. Behaviour was identified from accelerometer measurements, and DBA and heart rate were averaged over bouts of consistent behaviour. Heart rate was converted to metabolic rate using an allometric calibration, after confirming its fit using metabolic measurements taken in captivity and values from existing literature. Both proxies showed similar changes among behaviours, though DBA overestimated costs of floating, likely due to waves. However, relationships between DBA and heart rate were weak within a behaviour mode, possibly due to the lack of synchrony between proxy measurements. On daily scales, DBA and TEBs perform comparably for estimating daily energy expenditure. Accelerometery methods deviated from a 1:1 relationship with heart rate because acceleration could not measure variation in resting metabolic costs. We conclude DBA functions well for detecting energy expenditure arising from activity costs, including during soaring flight. We discuss scenarios where one method (DBA vs. TEBs) may be preferred over the other. Read the free Plain Language Summary for this article on the Journal blog.

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.

Migratory birds may need to cross barriers such as seas, without opportunities to rest or refuel. Waterbirds, unlike land birds, can stop at sea to rest or wait for better winds and thus may be less selective for supportive winds at departure and tolerate larger drift. However, pay-offs of waiting are likely to depend on circumstances (e.g. pressure for well-timed arrival, wind availability and travelling with/without juvenile brood), thus migratory behaviour during barrier crossings is expected to differ between seasons. We studied pink-footed geese Anser brachyrhynchus crossing the Barents Sea (ca 650 km), in spring and autumn during 2018–2020, using 94 GPS-tracks of 38 individuals, with annotated ERA5 weather data. We found that 1) especially in autumn, geese selected supportive winds for departure; 2) in spring, geese experienced lower wind support and more crosswinds than in autumn, leading to 23% longer routes, 60% longer durations, 93% longer air distances and 45% higher ratios of air-to-ground distances; 3) in both seasons, geese had more tailwinds in the first part of crossings, and in spring when deviating more from the shortest route; 4) geese stopped at sea more often in spring (mean 11×) than autumn (3×), in spring during earlier stages of crossings, but in both seasons, spent half of the crossing time at sea, during which they still continued to approach their destination slowly; 5) stops at sea happened mostly in adverse winds, warmer air, higher air humidity and on calmer water and, in autumn, took longer without juvenile brood. We conclude that for migrating pink-footed geese, Arctic capital breeders, the importance of time and energy can shift en route and that seasonal differences in wind support, flying and stopping behaviour and the pressure for a well-timed arrival cause the Barents Sea to be a larger barrier in spring than in autumn.

Climate change is driving worldwide shifts in the distribution of biodiversity, and fundamental changes to global avian migrations. Some arctic‐nesting species may shorten their migration distance as warmer temperatures allow them to winter closer to their high‐latitude breeding grounds. However, such decisions are not without risks, since this intensifies pressure on resources when they are used for greater periods of time. In this study, we used an individual‐based model to predict how future changes in food abundance, winter ice coverage, and human disturbance could impact an Arctic/sub‐Arctic breeding goose species, black brant (Branta bernicla nigricans, Lawrence 1846), and their primary food source, common eelgrass (Zostera marina L.), at the Izembek Lagoon complex in southwest Alaska. Brant use the site during fall and spring migrations, and increasingly, for the duration of winter. The model was validated by comparing predictions to empirical observations of proportion of geese surviving, proportion of geese emigrating, mean duration of stay, mean rate of mass gain/loss, percentage of time spent feeding, number of bird days, peak population numbers, and distribution across the complex. The model predicted that reductions >50% of the current decadal (2007–2015) mean of eelgrass biomass, which have been observed in some years, or increases in the number of brant, could lead to a reduction in the proportion of birds that successfully migrate to their breeding grounds from the site. The model also predicted that access to eelgrass in lagoons other than Izembek was critical for overwinter survival and spring migration of brant, if overall eelgrass biomass was 50% of the decadal mean biomass. Geese were typically predicted to be more vulnerable to environmental change during winter and spring, when eelgrass biomass is lower, and thermoregulatory costs for the geese are higher than in fall. We discuss the consequences of these predictions for goose population trends in the face of natural and human drivers of change.

Many species of large herbivore rely on agricultural land for their feeding habitats, but available food resources are highly variable in space and time. The conservation and management of farmland-dependant herbivores would therefore benefit from predictions about how species will respond to changes in their environment. We developed an individual-based model (IBM) to provide such predictions for three overwintering avian herbivore species that feed on agricultural land: Bewick's swans (Cygnus columbianus bewickii), whooper swans (Cygnus cygnus), and mute swans (Cygnus olor). Our validated model predicted how potential future changes in food availability and competition would affect (i) the proportion of the current swan population that could be supported, (ii) the proportion of swans that successfully departed on migration at the end of winter, (iii) swan daily foraging effort, and (iv) late winter crop biomasses. Regardless of competitor numbers or food availability, all individuals were predicted to avoid starvation and depart successfully. Individual swans offset higher competition and reduced food availability by increasing the proportion of daylight spent foraging. Our simulations indicate that swans have considerable capacity to buffer against losses of food resources and increased competition by increasing their foraging effort, but this may result in additional grazing damage to agricultural crops. Our findings suggest that the recent c.40% decline in Bewick's swan numbers was unlikely to be linked to changes in winter food resources or competition.

Sensors, such as accelerometers, in tracking devices allow for detailed bio-logging to understand animal behaviour, even in remote places where direct observation is difficult. To study breeding in birds remotely, one needs to understand how to recognize a breeding event from tracking data, and ideally validate this by direct observation. We tagged 49 adult female Pink-footed Geese (Anser brachyrhynchus) with transmitter neckbands in Finland in spring of 2018 and 2019, and in Svalbard in summer 2018, and validated inferences from tracking by field observations of nesting sites and family status in 2018-2020 (54 spring-summer tracks). We estimated nesting locations by taking the median coordinates of GPS-fixes at which the goose was motionless (overall dynamic body acceleration, ODBA<1) on days with a daily median ODBA<1, which approached the real nesting locations closely (within 1.6-3.7 m, n=6). The start of nesting was defined as the first day on which the goose spent >75% of time within 50 m of the nest, because nest site attendances steeply increased within one day to above this threshold. Nesting duration (number of consecutive days with >75% nest site attendance) ranged between 3-44 days (n=28), but was 30-34 days in confirmed successful nests (n=9). The prolonged nesting of 39-44 days (n=3) suggested incubation on unhatchable egg(s). Nest losses before hatching time occurred mostly in day 3-10 and 23-29 of nesting, periods with an increased frequency of nest site recesses. As alternative method, allowing for non-simultaneous GPS and accelerometer data, we show that nesting days were classified with 98.6% success by two general characteristics of breeding: low body motion (daily median ODBA) and low geographic mobility (daily SD of latitude). Median coordinates on nesting days approached real nest sites closely (within 0.8-3.6 m, n=6). When considering only geographic mobility (allowing for GPS data only) nesting locations were similarly accurate, but some short nesting attempts were undetected and non-breeding tracks misclassified. We show that nesting attempts, as short as 3 days, and nesting success can be detected remotely with good precision using GPS-tracking and accelerometry. Our method may be generalized to other (precocial) bird species with similar incubation behaviour.

Climate warming in the Arctic has led to warmer and earlier springs, and as a result, many food resources for migratory animals become available earlier in the season, as well as become distributed further northwards. To optimally profit from these resources, migratory animals are expected to arrive earlier in the Arctic, as well as shift their own spatial distributions northwards. Here, we review literature to assess whether Arctic migratory birds and mammals already show shifts in migration timing or distribution in response to the warming climate. Distribution shifts were most prominent in marine mammals, as expected from observed northward shifts of their resources. At least for many bird species, the ability to shift distributions is likely constrained by available habitat further north. Shifts in timing have been shown in many species of terrestrial birds and ungulates, as well as for polar bears. Within species, we found strong variation in shifts in timing and distributions between populations. Our review thus shows that many migratory animals display shifts in migration timing and spatial distribution in reaction to a warming Arctic. Importantly, we identify large knowledge gaps especially concerning distribution shifts and timing of autumn migration, especially for marine mammals. Our understanding of how migratory animals respond to climate change appears to be mostly limited by the lack of long-term monitoring studies.

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.

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 this is because the timing of spring affects different stages of the reproductive cycle differently in different species. Because the effects on two 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 on the other hand 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.

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.

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 or 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 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. By 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 6 time 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.

Migratory species are threatened worldwide by climate change, overexploitation, and habitat changes. Availability of suitable habitat is important for flying migrants, and in particular for large birds that use the energetically expensive flapping flight mode, such as the Bewick’s Swan (Cygnus columbianus bewickii). Bewick’s Swans largely feed on aquatic macrophytes during migration that may disappear when nutrient levels, waves, and turbidity exceed certain thresholds. Macrophyte collapse has been suggested as a reason for the sharp decline of the Bewick’s Swan population during 1995–2015. We used Bewick’s Swans fitted with GPS/GSM neck collars including an accelerometer and water sensor to record the occurrence of aquatic foraging in remote stopovers along their migratory route. We concentrated on spring migration, when stopovers are longer than during autumn, and focused on four key sites identified in earlier tracking studies. Within these sites, we identified areas that are protected based on the World Database on Protected Areas (WDPA). During three years (2017–2019), we obtained a total of 51 complete spring tracks of adult female Bewick’s Swans. Most swans showed aquatic foraging along the coast of Estonia and in Dvina Bay, and about half in the Gulf of Finland and Cheskaya Bay. In Estonia and in the Gulf of Finland, swans predominantly used protected zones, but in Dvina Bay swans also foraged extensively in areas that are currently not protected according to WDPA. No protected areas occur in Cheskaya Bay. Macrophyte vegetation is threatened by ongoing or planned construction works in the Gulf of Finland and Dvina Bay, and by future oil and gas exploitation in Cheskaya Bay. Our study shows how migrants can be used as sentinels to pinpoint areas that require protection in order to maintain a chain of suitable stopovers on their migration.

The ability of a species to adapt to environmental change is ultimately reflected in its vital rates – i.e. survival and reproductive success of individuals. Together, vital rates determine trends in numbers, commonly monitored using counts of species abundance. Rapid changes in abundance can give rise to concern, leading to calls for research into the biological mechanisms underlying variations in demography. For the northwest European population of Bewick's swan Cygnus columbianus bewickii, there have been major changes in the population trends recorded during nearly five decades of monitoring (1970–2016). The total number of birds increased to a maximum of ca 30 000 in 1995 and subsequently decreased to about 18 000 individuals in 2010. Such large fluctuation in population numbers is rare in long‐lived species and understanding the drivers of this population change is crucial for species management and conservation. Using the integrated population model (IPM) framework, we analysed three demographic datasets in combination: population counts, capture–mark–resightings (CMR) and the proportion of juveniles in winter over a period of ~50 years. We found higher apparent breeding success in the years when the population had a positive growth rate compared to years with a negative growth rate. Moreover, no consistent trend in adult and yearling survival, and an increasing trend in juvenile survival was found. A transient life‐table response experiment showed that apparent breeding success and adult survival contributed most to the variation in population trend. We explored possible explanatory variables for the different demographic rates and found a significant association between juvenile survival both with the water level in lakes during autumn migration, which affects food accessibility for the swans, and with summer temperatures. Such associations are important for understanding the dynamics of species with fluctuating population sizes, and thus for informing management and conservation decisions.

Accelerometers in tracking devices are increasingly used to identify behaviour leading to detailed insights into the lives of free-ranging animals. To make proper use of an accelerometer, their settings and signals need to be tested and calibrated. Calibration of an accelerometer can be done by directly observing an individual animal of the species of interest, while an accelerometer is simultaneously measuring the movements of this individual. In case direct observations are difficult to obtain, this procedure can be performed with captive individuals. This study sought to calibrate the accelerometer sensor in GPS/GSM neck-collars in Bewick’s swans Cygnus columbianus bewickii with observations of captive individuals in three zoos in the Netherlands. Using a random forest classification model, five behavioural classes were classified with an overall accuracy of 91%. An additional behavioural class (aquatic foraging) was identified based on a water sensor that was also included in the GPS/GSM collars. This classification was subsequently applied to accelerometer data from 12 free-ranging Bewick’s swans equipped with these neck-collars to identify their behaviour during two spring migrations (2017 and 2018). The resulting time-activity budgets were in general agreement with current knowledge based on fragmentary field observations of Bewick’s swan flocks along the flyway. The study shows how observations of zoo individuals can be instrumental to derive time-activity budgets of free-ranging individuals that can contribute to further research into the ecology of the species.

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.

Background
For the conservation and management of migratory species that strongly decrease or increase due to anthropological impacts, a clear delineation of populations and quantification of possible mixing (migratory connectivity) is crucial. Usually, population exchange in migratory species is only studied in breeding or wintering sites, but we considered the whole annual cycle in order to determine important stages and sites for population mixing in an Arctic migrant.

Methods
We used 91 high resolution GPS tracks of Western Palearctic greater white-fronted geese (Anser A. albifrons) from the North Sea and Pannonic populations to extract details of where and when populations overlapped and exchange was possible. Overlap areas were calculated as dynamic Brownian bridges of stopover, nest and moulting sites.

Results
Utilisation areas of the two populations overlapped only somewhat during spring and autumn migration stopovers, but much during moult. During this stage, non-breeders and failed breeders of the North Sea population intermixed with geese from the Pannonic population in the Pyasina delta on Taimyr peninsula. The timing of use of overlap areas was highly consistent between populations, making exchange possible. Two of our tracked geese switched from the North Sea population flyway to the Pannonic flyway during moult on Taimyr peninsula or early during the subsequent autumn migration. Because we could follow one of them during the next year, where it stayed in the Pannonic flyway, we suggest that the exchange was long-term or permanent.

Conclusions
We have identified long-distance moult migration of failed or non-breeders as a key phenomenon creating overlap between two flyway populations of geese. This supports the notion of previously suggested population exchange and migratory connectivity, but outside of classically suggested wintering or breeding sites. Our results call for consideration of moult migration and population exchange in conservation and management of our greater white-fronted geese as well as other waterfowl populations.

“Everything changes and nothing stands still” (Heraclitus). Here we review three major improvements to freshwater aquatic ecosystem models — and ecological models in general — as water quality scenario analysis tools towards a sustainable future. To tackle the rapid and deeply connected dynamics characteristic of the Anthropocene, we argue for the inclusion of eco-evolutionary, novel ecosystem and social-ecological dynamics. These dynamics arise from adaptive responses in organisms and ecosystems to global environmental change and act at different integration levels and different time scales. We provide reasons and means to incorporate each improvement into aquatic ecosystem models. Throughout this study we refer to Lake Victoria as a microcosm of the evolving novel social-ecological systems of the Anthropocene. The Lake Victoria case clearly shows how interlinked eco-evolutionary, novel ecosystem and social-ecological dynamics are, and demonstrates the need for transdisciplinary research approaches towards global sustainability.

While many species suffer from human activities, some like geese benefit and may show range expansions. In some cases geese (partially) gave up migration and started breeding at wintering and stopover grounds. Range expansion may be facilitated and accompanied by physiological changes, especially when associated with changes in migratory behaviour. Interspecific comparisons found that migratory tendency is associated with a higher basal or resting metabolic rate (RMR). We compared RMR of individuals belonging to a migratory and a sedentary colony of barnacle geese Branta leucopsis. The migratory colony is situated in the traditional arctic breeding grounds (Russia), whereas the sedentary colony has recently been established in the now shared wintering area (the Netherlands). We measured RMR by oxygen consumption (urn:x-wiley:00301299:media:oik12892:oik12892-math-0001) during two ontogenetic phases (juvenile growth and adult wing moult). We also investigated juvenile growth rates and adult body mass dynamics.

Mass‐independent urn:x-wiley:00301299:media:oik12892:oik12892-math-0002 was 13.6% lower in goslings from the sedentary colony than in goslings from the migratory colony. Similarly, in adult geese, mass‐independent urn:x-wiley:00301299:media:oik12892:oik12892-math-0003 was 15.5% lower in sedentary than in migratory conspecifics. Goslings in the Netherlands grew 36.2% slower than goslings in Russia, while we found no differences in body dimensions in adults. Adult geese from both colonies commenced wing moult with similar body stores, but whereas Russian barnacle geese maintained this level throughout moult, body stores in geese from the Netherlands fell, being 8.5% lower half‐way through the moult.

We propose that the colony differences in resting metabolic rate, growth rate and body mass dynamics during moult can be explained by environmental and behavioural differences. The less stringent time constraints combined with poorer foraging opportunities allow for a smaller ‘metabolic machinery’ in non‐migratory geese. Our analysis suggests that range expansion may be associated with changes in physiology, especially when paired with changes in migratory tendency.

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, 5, 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, 13, 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.

Agricultural crops have become increasingly important foraging habitats to geese and swans in northern Europe, and a recent climate-driven expansion in the area of maize fields has led to a rapid increase in the exploitation of this habitat. However, due to the novelty of maize foraging in this region, little is known about the abundance and energetic value of this resource to foraging birds. In this study we quantify food availability, intake rates and energetic profitability of the maize stubble habitat, and describe the value of this increasingly cultivated crop to wintering geese and swans in the region. Our results indicate that the maize resource varies considerably among fields and years, but also that the energetic returns from maize foraging is substantial. As such, fields with extensive spill allow foraging birds to fulfill their daily energetic demands in 4 h of active foraging. Both the area of cultivated maize fields and the importance of this habitat to foraging birds are expected to increase in years to come. This may alleviate conflicts with other more vulnerable crops such as winter cereals, and have the potential to affect migratory decisions, site use and population dynamics of geese and swans wintering in northern Europe.

Abstract 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.

Many migratory goose populations have thrived over the past decades and their reliance on agricultural resources has often led to conflicts. Control and management measures are sought after but since migratory geese use several sites in their annual cycle, local management actions should consider their potential effects further down the flyway.
We used a behaviour‐based migration model to illustrate the consequences of management actions involving hunting, derogation shooting and scaring at single or multiple locations along the flyway, considering various mechanisms of how geese might perceive shooting/hunting. Furthermore, as a proxy for the agricultural damage caused, we calculated the per capita biomass consumption between scenarios—both over time and cumulatively.
We found that hunting, shooting and scaring can result in a suite of direct and indirect consequences on migration and foraging behaviour. Most importantly, hunting/shooting on a particular site had implications not only for the behaviour at the actual site but also for behaviour at, and use of, other sites. Furthermore, the specific consequences of shooting/hunting could be counter‐intuitive, that is, aggravate rather than alleviate agricultural damage, depending on where along the migration route changes had taken place and the mechanisms through which hunting/shooting was assumed to affect geese.
Synthesis and applications. Management plans are being discussed or implemented for several migratory goose populations and often include shooting, hunting or scaring at one or multiple locations. Using a behaviour‐based model, we assessed the consequences of such local management measures and found that they can indeed lead to a reduction of agricultural conflicts locally but may also aggravate the conflict or shift it to other sites along the flyway. Thus, we recommend the use of these models to scrutinize the efficiency of specific management measures and to assist in identifying an international management regime that minimizes conflicts on a flyway level while still maintaining migratory populations.

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 of 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 in 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. Apart 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.

Following increases in numbers during the second half of the 20th century, several Arctic-breeding migrant bird species are now undergoing sustained population declines. These include the northwest European population of Bewick's Swan Cygnus columbianus bewickii, which declined from c. 29 000 birds on the wintering grounds in 1995 to 18 000 in 2010. It is unclear whether this decrease reflects reduced survival, emigration to a different area, or a combination of both. Furthermore, the environmental drivers of any demographic changes are also unknown. We therefore used an information-theoretic approach in RMark to analyse a dataset of 3929 individually marked and resighted Bewick's Swans to assess temporal trends and drivers of survival between the winters of 1970/71 and 2014/2015, while accounting for effects of age, sex and different marker types. The temporal trend in apparent survival rates over our study period was best explained by different survival rates for each decade, with geometric mean survival rates highest in the 1980s (leg-ring marked birds = 0.853, 95% confidence interval (CI) 0.830–0.873) and lowest in the 2010s (leg-ring = 0.773, 95% CI 0.738–0.805; neck-collar = 0.725, 95% CI 0.681–0.764). Mean (±95% CI) resighting probabilities over the study period were higher for birds marked with neck-collars (0.91 ± 0.01) than for those marked with leg-rings (0.70 ± 0.02). Weather conditions in different areas across the flyway, food resources on the winter grounds, density-dependence and the growth of numbers at a relatively new wintering site (the Evros Delta in Greece) all performed poorly as explanatory variables of apparent survival. None of our 18 covariates accounted for more than 7.2% of the deviance associated with our survival models, with a mean of only 2.2% of deviance explained. Our results provide long-term demographic information needed to help conservationists understand the population dynamics of Bewick's Swans in northwest Europe.

Long-distance migratory birds rely on the acquisition of body stores to fuel their migration and reproduction. Breeding success depends
on the amount of body stores acquired prior to migration, which is thought to increase with access to food at the fueling site. Here, we
studied how food abundance during fueling affected time budgets and reproductive success. In a regime of plenty, we expected that
1) limitations on food harvesting would become lifted, allowing birds to frequently idle, and 2) birds would reach sufficient fuel loads,
such that departure weight would no longer affect reproductive success. Our study system comprised brent geese (Branta b. bernicla)
staging on high-quality agricultural pastures. Fueling conditions were assessed by a combination of high-resolution GPS tracking,
acceleration-based behavioral classification, thermoregulation modeling, and measurements of food digestibility and excretion rates.
Mark-resighting analysis was used to test for correlations between departure weight and offspring recruitment. Our results confirm
that birds loafed extensively, actively postponed fueling in early spring, and took frequent digestion pauses, suggesting that traditional
time constraints on harvest and fueling rates are absent on modern-day fertilized grasslands. Nonetheless, departure weight remained
correlated with recruitment success. The persistence of this correlation after a prolonged stopover with access to abundant highquality
food, suggests that between-individual differences in departure condition are not so much enforced by food quality and availability
during stopover, but reflect individual quality and longer-lived life-history traits, such as health status and digestive capacity,
which may be developed before the fueling period.

Broad‐scale land conversions and fertilizer use have dramatically altered the available staging area for herbivorous long‐distance migrants. Instead of natural land, these birds rely increasingly on pastures for migratory fuelling and stopover, often conflicting with farming practices. To predict and manage birds’ future habitat use, the relative advantages and disadvantages of natural (e.g. saltmarsh, intertidal) versus anthropogenic staging sites for foraging need to be understood.
We compared the migratory staging of brent geese on saltmarsh and pasture sites in spring. Food quality (nitrogen and fibre content), antagonistic behaviour, and body weight were quantified at nearby sites in simultaneous seasons. Individuals were tracked with high‐resolution GPS and accelerometers to compare timing of migration and time budgets during fuelling.
On pastures, birds rested more and experienced higher ingestion rates, similar or superior food quality and reduced antagonistic interactions than on saltmarsh.
Brent geese using fertilized grasslands advanced their fuelling and migration schedules compared to those using saltmarsh. Pasture birds reached heavy weights earlier, departed sooner, and arrived in the Arctic earlier.
Intertidal mudflats were frequently visited by saltmarsh birds during the day, and available food there (algae, some seagrass) was of higher quality than terrestrial resources. Availability of intertidal resources was an important factor balancing the otherwise more favourable conditions on pastures relative to saltmarsh.
Synthesis and applications. Disadvantages of longer foraging effort, more antagonistic interactions and delayed fuelling schedules on traditional saltmarshes may cause geese to exchange this traditional niche in favour of pastures, especially in a warming climate that requires advancement of migratory schedules. However, due to its high quality, intertidal forage can complement terrestrial foraging, potentially removing the incentive for habitat switches to pastures. The relatively high quality of green algae and seagrass, and birds’ remarkable preference for these resources when available, provides a key for managers to create landscapes that can sustain this specialist’s intertidal lifestyle. To keep natural habitats attractive to staging geese with the purpose of preventing conflicts with farming practices, management actions should focus on conservation and restoration of saltmarsh and especially intertidal habitat.

Background
Tracking devices have enabled researchers to study unique aspects of behavior in birds. However, it has become clear that attaching these devices to birds often affects their survival and behavior. While most studies only focus on negative effects on return rates, tracking devices can also affect the behavior under study, and it is therefore important to measure potential negative effects of tracking device attachment on the full range of behavioral aspects of birds. At the same time, we should aim to improve our current attachment methods to reduce these effects.

Results
We used a modified harness to attach tracking devices to a total of 111 individuals of three goose species (Greater White-fronted Geese, Brent Geese, and Barnacle Geese) to study their migratory behavior. By creating control groups of birds marked with colored leg bands, geolocators, and/or neck collars, we were able to compare return rates, body condition, and migratory and reproductive behavior, thus allowing a much broader comparison than return rates alone. Birds with harness-attached tracking devices had lower return rates, which could partly be explained by increased rates of divorce, but is likely also the result of reduced survival induced by the harness and device. A comparison between Barnacle Geese equipped with harness-attached tracking devices and individuals fitted with geolocators attached to leg bands showed that birds equipped with tracking devices were only slightly delayed in timing of migration and reproduction and otherwise were not affected in reproductive output.

Conclusions
We argue that tracking devices can be used for studies on migration timing. Nevertheless, given the effect of tracking devices on survival and divorce rate, which may differ between sexes and species, we stress that researchers should carefully consider which birds to tag in order to reduce potential negative effects.

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.

Economic development and energy exploration are increasing in the Arctic. Important breeding habitats for many waterbird species, which have previously been relatively undisturbed, are now being subjected to these anthropogenic pressures. The conservation of the habitats and the species they support is a significant challenge for sustainable development. Even if governments and corporates operating in this fragile environment are committed to sustainable development, there is little information available to avoid, mitigate and manage environmental risk and impacts. Taking a risk management perspective, we followed the International Finance Corporations (IFC) Performance Standard 6 (PS6) criteria on Environmental and Social Sustainability and developed an approach to identify "critical habitat", as defined in IFC PS6, for waterbird species breeding in the Arctic. While the range of these waterbirds is roughly known, more accuracy is needed for proper risk assessment. We have therefore gone a step further by modelling suitable habitat within these ranges. Depending on the relevance of the species for IFC PS6 and the level of certainty we separated the classes likely and potential critical habitat. We tested the approach for Russian breeding populations of five Anatidae species (White-fronted Goose Anser albifrons, Lesser White-fronted Goose Anser erythropus, Brent Goose Branta bernicla, Redbreasted Goose Branta ruficollis and Bewick's Swan Cygnus columbianus bewicki). Likely critical habitats were identified through a review of literature and available data for these waterbird species and multi-species congregations. To address the information gap for most of the Russian Arctic a species distribution modelling approach was used. The outputs of this approach were labelled as potential critical habitat, indicating the lower level of certainty than likely critical habitat. Based on existing information the amount of likely critical habitat is estimated to be at least x,xxx,xxx km2. For the five Anatidae species, X,XXX,XXX km2 potential critical habitat was identified; 95% of these areas were outside of the area boundaries of likely critical habitat for the species. Insufficient data in the east of the study area did affect the results, as some areas known to support breeding populations were not identified as suitable. Conversely, species' distributions may be overpredicted in other areas; It should also be recognized that the analyzed species currently have depressed populations and may therefore only utilize a proportion of suitable habitat available. For risk assessment purposes however, it is better to predict false positives, rather than false negatives. The study indicates that there are large areas in the Arctic that are potentially important for each of the Anatidae species modelled, but are not yet recognised as key important areas. The results confirm that there is still much to learn about waterbird distribution and abundance in the Russian Arctic. Synthesis and applications: The critical habitat maps produced do not just provide a new source of information for the economic development sector, but provide it in a way that is relevant to the sector and directly applicable. The maps are useful for initial risk assessments of potential developments, to identify likely impacts and to consider mitigation options, in accordance with IFC PS6. Risk assessors should exercise caution and detailed surveys for any development in areas predicted to be suitable for each species should be carried out.

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.

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.

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.

Dispersal via animals (zoochory) is a primary mechanism for seed exchange between habitat patches. Recent studies have established that many plant species can survive waterbird gut passage. To quantify the patterns and consequences of waterbird-mediated dispersal, information on ingestion and gut passage must be combined with bird movement data. Such analysis has recently revealed seed dispersal kernels by migrating waterbirds. However, since many waterbird populations are largely resident, and migrating populations spend only a minor part of the main dispersal season (autumn–winter) on active migration, daily regional-scale movements probably cause more frequent dispersal.
We synthesized high-resolution empirical data on landscape-scale movements and seed gut passage times in a key disperser species, the mallard (Anas platyrhynchos), using a spatially explicit, mechanistic model to quantify dispersal distributions resulting from daily autumn–winter movements. We evaluated how landscape composition and seed traits affect these dispersal patterns.
The model indicates that mallards generate highly clumped seed deposition patterns, dispersing seeds primarily between core areas used for foraging and resting. Approximately 34% of all dispersed seeds are transported to communal roost areas, which may function as reservoirs for mallard-dispersed species, and 7% are transported between foraging areas. Landscape-dependent movement patterns strongly affect the dispersal distributions, resulting in multi-modal dispersal kernels, with dispersal distances increasing with fragmentation of freshwater foraging habitat. Seed size-related gut retention times determine the proportion of seeds being dispersed away from the ingestion area, with larger seeds (20 mm3) having a 8–10% higher potential for long-distance dispersal than smaller seeds (0·2 mm3), if surviving gut passage. However, twice as many small seeds will finally accomplish long-distance dispersal due to their higher gut passage survival.
Synthesis. Firstly, this study reveals how seed dispersal patterns resulting from daily waterfowl movements are shaped by landscape-dependent differences in movement patterns. Secondly, seed survival appears more important than retention time in determining the scale of long-distance dispersal by non-migrating mallards. We conclude that the frequent flights of staging waterbirds result in directed dispersal over distances inversely related to wetland availability, indicating that they maintain landscape connectivity across a range from wet to increasingly dry landscapes.

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.

In 2014, H5N8 clade 2.3.4.4 highly pathogenic avian influenza (HPAI) viruses of the A/Goose/Guangdong/1/1996 lineage emerged in poultry and wild birds in Asia, Europe and North America. Here, wild birds were extensively investigated in the Netherlands for HPAI H5N8 virus (real-time polymerase chain reaction targeting the matrix and H5 gene) and antibody detection (haemagglutination inhibition and virus neutralisation assays) before, during and after the first virus detection in Europe in late 2014. Between 21 February 2015 and 31 January 2016, 7,337 bird samples were tested for the virus. One HPAI H5N8 virus-infected Eurasian wigeon (Anas penelope) sampled on 25 February 2015 was detected. Serological assays were performed on 1,443 samples, including 149 collected between 2007 and 2013, 945 between 14 November 2014 and 13 May 2015, and 349 between 1 September and 31 December 2015. Antibodies specific for HPAI H5 clade 2.3.4.4 were absent in wild bird sera obtained before 2014 and present in sera collected during and after the HPAI H5N8 emergence in Europe, with antibody incidence declining after the 2014/15 winter. Our results indicate that the HPAI H5N8 virus has not continued to circulate extensively in wild bird populations since the 2014/15 winter and that independent maintenance of the virus in these populations appears unlikely.

Background

GPS and accelerometer tracking presently revolutionises the fields of ecology and animal behaviour. However, the effects of tag characteristics like weight, attachment and data quality on study outcomes and animal welfare are important to consider. In this study, we compare how different tag attachment types influence the behaviour of a group of tagged large waterbirds, GPS accuracy and behaviour classification success from accelerometer data.

Results

Both neckband and backpack tags had similar effects on the behaviour of six captive Canada geese (Branta canadensis), increasing the amount of discomfort behaviour in relation to untagged individuals. Both treatment groups also slightly decreased the amount of foraging, but the duration of neither vigilance nor resting was affected. GPS positions that were filtered with classical GPS platform settings (i.e. smoothing) were more accurate than positions improved by satellite-based differential augmentation. Tag attachment, however, did not induce any differences in position accuracy of both data types. Behaviour classification success was generally similar for neckband and backpack tags. But in detail, behaviours mainly performed by the head like foraging and vigilance were better detected from accelerometer data of neckband tags, whereas behaviours like resting and walking were more successfully detected from backpack tag data.

Conclusion

Our findings suggest that the use of neckband or backpack tags for tracking large waterbirds and their behaviour largely depends on which behaviours are most important to detect. However, for wildlife tracking studies, factors like tag retention time are also of great importance, especially for animals like some goose species that are known to quickly destroy backpack tags. For future studies, we advise to carefully evaluate not only tag weight, but also attachment methods and data quality, because the right choice depends on the research question. This will improve the scope of wildlife tracking even more for various scientific, conservation and management applications.

In different ecosystems herbivores highly prefer particular plant species. This is often explained in a stoichiometric framework of nutrient-based plant adaptations to herbivory. We hypothesize that such super-palatability can also arise as an evolutionary by-product of osmoregulatory adaptations of plants to stressful environmental conditions, as salinity, drought and cold. Here, we investigate in a coastal salt marsh why some plant species are highly preferred by migratory brent geese Branta bernicla bernicla in spring while others are avoided. This salt marsh is an important spring staging site for the geese. Sufficient energy storage in a short period is critical to enable their northward migration to Siberia and subsequent reproduction. We test if geese prefer plants that balance their internal osmotic potential with the saline environment through energy-rich soluble sugars over plant species that use (compartmentalized) salts for this. We find that plant nitrogen and acid detergent fiber content, classic predictors of herbivore preferences, poorly explain which plants the geese prefer. Instead, plant species that are highly preferred by the geese adapt to salinity by high soluble sugar concentrations while avoided species do this by high plant salt concentrations. Thus, the type of osmoregulatory adaptation to stress displayed by different plant species is a good predictor for the food preference of geese on this salt marsh. We suggest that variation in other types of osmoregulation-based stress adaptations, as plant cold adaptations in tundras and plant drought adaptations in savannas, have similar important consequences for trophic interactions. This article is protected by copyright. All rights reserved.

Population declines among migratory Arctic-breeding birds are a growing concern for conservationists. To inform the conservation of these declining populations, we need to understand how demographic rates such as breeding success are influenced by combinations of extrinsic and intrinsic factors. In this study we examined inter-annual variation and long-term trends in two aspects of the breeding success of a migratory herbivore, the Bewick's Swan (Cygnus columbianus bewickii), which is currently undergoing a population decline: (i) the percentage of young within the wintering population and (ii) mean brood size. We used an information-theoretic approach to test how these two measures of productivity were influenced over a 26 year period by 12 potential explanatory variables, encompassing both environmental (e.g. temperature) and intrinsic (e.g. pair-bond duration) factors. Swan productivity exhibited sensitivity to both types of explanatory variable. Fewer young were observed on the wintering grounds in years in which the breeding period (May to September) was colder and predator (Arctic Fox) abundance was higher. The percentage of young within the wintering population also showed negative density-dependence. Inter-annual variance in mean swan brood size was best explained by a model comprised of the negative degree days during the swan breeding period, mean pair-bond duration of all paired swans (i.e. mean pair duration), and an interaction between these two variables. In particular, mean pair duration had a strong positive effect on mean brood size. However, we found no long-term directional trend in either measure of breeding success, despite the recent decline in the NW European population. Our results highlight that inter-annual variability in breeding success is sensitive to the combined effects of both intrinsic and extrinsic factors.

Breeding success of many Arctic-breeding bird populations varies with lemming cycles due to prey switching behavior of generalist predators. Several bird species breed on islands to escape from generalist predators like Arctic fox Vulpes lagopus, but little is known about how these species interact. We studied brent geese Branta bernicla bernicla that share islands with gulls (Larus spec.) in Taimyr, Siberia (Russia). On one hand, gulls are egg predators, which occasionally steal an egg when incubating geese leave the nest for foraging bouts. On the other hand, gulls import marine resources to the islands, enriching the soil with their guano. We considered three hypotheses regarding clutch size of brent geese after partial nest predation. According to the “predator proximity hypothesis”, clutch size is expected to be smallest close to gulls, because of enhanced predator exposure. Conversely, clutch size is expected to be largest close to gulls, because of the supposedly better feeding conditions close to gulls, which might reduce nest recess times of geese and hence egg predation risk (“guano hypothesis”). Furthermore, gulls may defend their nesting territory, and thus nearby goose nests might benefit from this protection against other gulls (“nest association hypothesis”). We mapped goose and gull nests toward the end of the goose incubation period. In accordance with the latter two hypotheses, goose clutch size decreased with distance to the nearest gull nest in all but the lemming peak year. In the lemming peak year, clutch size was consistently high, indicating that partial nest predation was nearly absent. By mapping food quantity and quality, we found that nitrogen availability was indeed higher closer to gull nests, reflecting guanofication. Unlike predicted by the nest association hypothesis, a predation pressure experiment revealed that egg predation rate decreased with distance to the focal gull nests. We therefore propose that higher food availability close to gulls enables female geese to reduce nest recess time, limiting egg predation by gulls.

Predicting the environmental impact of a proposed development is notoriously difficult, especially when future conditions fall outside the current range of conditions. Individual-based approaches have been developed and applied to predict the impact of environmental changes on wintering and staging coastal bird populations. How many birds make use of staging sites is mostly determined by food availability and accessibility, which in the case of many waterbirds in turn is affected by water level. Many water systems are regulated and water levels are maintained at target levels, set by management authorities. We used an individual-based modelling framework (MORPH) to analyse how different target water levels affect the number of migratory Bewick’s swans Cygnus columbianus bewickii staging at a shallow freshwater lake (Lauwersmeer, the Netherlands) in autumn. As an emerging property of the model, we found strong non-linear responses of swan usage to changes in water level, with a sudden drop in peak numbers as well as bird-days with a 0.20 m rise above the current target water level. Such strong non-linear responses are probably common and should be taken into account in environmental impact assessments.

The migration strategy of many capital breeders is to garner body stores along the flyway at distinct stopover sites. The rate at which they can fuel is likely to be strongly influenced by a range of factors, such as physiology, food availability, time available for foraging and perceived predation. We analysed the foraging behaviour and fuel accumulation of pink-footed geese, an Arctic capital breeder, at their mid- flyway spring stopover site and evaluated to what extent their behaviour and fuelling were related to physiological and external factors and how it differed from other stopovers along the flyway. We found that fuel accumulation rates of geese at the mid-flyway site were limited by habitat availability rather than by digestive constraints. However, as the time available for foraging increased over the stopover season, geese were able to keep constant fuelling rate. Putting this in perspective, geese increased their daily net energy intake along the flyway corresponding to the increase in time available for foraging. The net energy intake per hour of foraging remained the same. Geese showed differences in their reaction to predators/disturbance between the sites, taking higher risks particularly at the final stopover site. Hence, perceived predation along the flyway may force birds to postpone the final fuel accumulation to the last stopover along the flyway. Flexibility in behaviour appears to be an important trait to ensure fitness in this capital breeder. Our findings are based on a new, improved method for estimating fuel accumulation of animals foraging in heterogeneous landscapes based on data obtained from satellite telemetry and habitat specific intake rates.

1.With increasing numbers of many herbivorous waterfowl species, often foraging on farmland, the conflict with agriculture has intensified. One popular management tool is to scare birds off the land, often in association with shooting. However, the energy costs of flying are considerably higher than those of resting. Therefore, when birds fly off after a disturbance, they use extra energy that subsequently needs to be compensated.

2.We used the white-fronted goose Anser albifrons, the most common (grass-eating) species wintering in western Europe, as a model species. We measured flight durations by high-frequency accelerometer recordings over 2 × 24 h in 9 focal geese that were only incidentally disturbed. We also made direct observations on these days to determine whether the flight durations were reliably recorded. Using both a simple and a more realistic model of the energy balance, we calculated the extra grass consumption resulting from additional intentional disturbances.

3.On average, the geese flew daily 2 × 323 s (from and to their roosting sites at 3200 m), and furthermore took to the air 5.3 times during a day (and 1.9 times a night). Multiplied with the average flight durations of 195 s, this gives a total flying time of almost 0.6 h day-1 and a total foraging time of 7.4 h day-1. The extra foraging time needed to compensate for additional intentional disturbances strongly depends on the frequency of such disturbances and the following flight duration. If, for example, flights when intentionally disturbed are twice as long (2 × 195 s), the extra foraging time will be 3.7% day-1 (2.3–3.2% day-1 in the more realistic model) for each intentional disturbance, and the geese will no longer be able to cover their energy requirements when intentionally disturbed six times per day.

4.Synthesis and applications. Recent experiments suggest that geese have to be scared frequently in order to reduce goose visitation to particular fields. With an intentional disturbance rate e.g. of five times a day, the birds’ compensation for the increased energy expenditure will lead to a higher overall consumption of grass of 11.5–16 % day-1. Accommodation schemes have to take such increases in total grass consumption into account when deciding on the refuge areas to be set aside.

Dispersal of seeds by animals is an important mechanism regulating plant diversity, range expansions and invasions. Many birds, mammals, fish, and reptiles regularly ingest, transport and excrete viable seeds (known as endozoochory). The effectiveness of endozoochory is modelled in dispersal kernels: functions that describe seed shadows in the landscape by combining movement of animals with experimentally obtained seed retention times and survival. Currently, dispersal kernels use experimental data from resting animals, yet only moving animals disperse seeds. Although physical activity is known to affect digestive processes, little is known on how and to what extent this may influence current estimates of endozoochory. Activity may either prolong seed retention in the animal's gut (locomotion-priority mode hypothesis) or may not affect seed excretion rate (digestion-priority mode hypothesis), and may affect seed survival and germination positively or negatively. We tested how activity alters dispersal estimates in fish. We compared the seed dispersal potential of two riparian plant species (Carex acuta and C. riparia) by the common carp (Cyprinus carpio) subjected to three different activity levels: low (basal metabolic rate, BMR), medium (2×BMR), or high activity (3×BMR). Physical activity of the fish did not affect the number of intact retrieved seeds over 15 h of activity, but significantly affected seed retrieval patterns over time for both seed species. More active fish started seed excretion about 1 h later and kept excreting seeds at least 2 h longer. Effects of gut passage on germination could only be tested for C. acuta, where it reduced the percentage of germinating seeds by 22%, independent of the activity level. Seeds ingested by the fish germinated on average 3.5 days later than non-ingested control seeds. Seed retention times did not affect the timing of germination. Our results support the locomotion-priority mode hypothesis, and show that modelling dispersal kernels using parameters from inactive fish may underestimate potential dispersal distances. Because a trade-off between physical activity and digestive physiology is likely common in animals, it should be taken into account in future modelling of endozoochorous seed dispersal kernels. This article is protected by copyright. All rights reserved.

The migration timing of birds can be controlled by endogenous parameters. However, little is known about how environmental parameters influence the timing of migration and which have the greatest influence at different stages of migration. In this study we identified the main environmental parameters that correlate with the timing of the last stage of spring migration for the barnacle goose, Branta leucopsis. GPS tracking data were registered for 12 barnacle geese (in 2008–2010) on the Russian flyway and 17 (2006–2010) on the Svalbard flyway. A linear mixed-effect model and principal component analysis were used to retrieve statistically significant parameters. Departure date from the last staging site on the Russian flyway was related to daylength, temperature, cloud cover and barometric pressure, and on the Svalbard flyway to a food availability index and daylength. Arrival date at the Russian breeding site was related to cloud cover and barometric pressure en route and the food availability index and temperature at the breeding site. For the Svalbard flyway, temperature and cloud cover en route and the food availability index, wind, temperature and cloud cover at the breeding site were significantly related to arrival date at the breeding site. Our study highlights the importance of environmental parameters including food, weather and daylength for the last stage of goose spring migration. We found different priorities in selecting the environmental parameters in migration timing decisions between Svalbard and Russian barnacle geese which fly over sea and over land, respectively. Identifying the key factors that act as cues during the final stages of spring migration is important when assessing the possible effects of climate change on the timing of migration for a highly selective herbivore such as the barnacle goose.

According to migration theory and several empirical studies, long-distance migrants are more time-limited during spring migration and should therefore migrate faster in spring than in autumn. Competition for the best breeding sites is supposed to be the main driver, but timing of migration is often also influenced by environmental factors such as food availability and wind conditions.

Using GPS tags, we tracked 65 greater white-fronted geese Anser albifrons migrating between western Europe and the Russian Arctic during spring and autumn migration over six different years. Contrary to theory, our birds took considerably longer for spring migration (83 days) than autumn migration (42 days). This difference in duration was mainly determined by time spent at stopovers.

Timing and space use during migration suggest that the birds were using different strategies in the two seasons: In spring they spread out in a wide front to acquire extra energy stores in many successive stopover sites (to fuel capital breeding), which is in accordance with previous results that white-fronted geese follow the green wave of spring growth. In autumn they filled up their stores close to the breeding grounds and waited for supportive wind conditions to quickly move to their wintering grounds. Selection for supportive winds was stronger in autumn, when general wind conditions were less favourable than in spring, leading to similar flight speeds in the two seasons. In combination with less stopover time in autumn this led to faster autumn than spring migration.

White-fronted geese thus differ from theory that spring migration is faster than autumn migration. We expect our findings of different decision rules between the two migratory seasons to apply more generally, in particular in large birds in which capital breeding is common, and in birds that meet other environmental conditions along their migration route in autumn than in spring.

Until the 1990s, herbivory on aquatic vascular plants was considered to be of minor importance, and the predominant view was that freshwater and marine macrophytes did not take part in the food web: their primary fate was the detritivorous pathway. In the last 25 years, a substantial body of evidence has developed that shows that herbivory is an important factor in the ecology of vascular macrophytes across freshwater and marine habitats. Herbivores remove on average 40-48% of plant biomass in freshwater and marine ecosystems, which is typically 5-10 times greater than reported for terrestrial ecosystems. This may be explained by the lower C:N stoichiometry found in submerged plants. Herbivores affect plant abundance and species composition by grazing and bioturbation and therewith alter the functioning of aquatic ecosystems, including biogeochemical cycling, carbon stocks and primary production, transport of nutrients and propagules across ecosystem boundaries, habitat for other organisms and the level of shoreline protection by macrophyte beds. With ongoing global environmental change, herbivore impacts are predicted to increase. There are pressing needs to improve our management of undesirable herbivore impacts on macrophytes (e.g. leading to an ecosystem collapse), and the conflicts between people associated with the impacts of charismatic mega-herbivores. While simultaneously, the long-term future of maintaining both viable herbivore populations and plant beds should be addressed, as both belong in complete ecosystems and have co-evolved in these long before the increasing influence of man. Better integration of the freshwater, marine, and terrestrial herbivory literatures would greatly benefit future research efforts.

Highly pathogenic avian influenza (HPAI) A(H5N8) viruses that emerged in poultry in east Asia since 2010 spread to Europe and North America by late 2014. Despite detections in migrating birds, the role of free-living wild birds in the global dispersal of H5N8 virus is unclear. Here, wild bird sampling activities in response to the H5N8 virus outbreaks in poultry in the Netherlands are summarised along with a review on ring recoveries. HPAI H5N8 virus was detected exclusively in two samples from ducks of the Eurasian wigeon species, among 4,018 birds sampled within a three months period from mid-November 2014. The H5N8 viruses isolated from wild birds in the Netherlands were genetically closely related to and had the same gene constellation as H5N8 viruses detected elsewhere in Europe, in Asia and in North America, suggesting a common origin. Ring recoveries of migratory duck species from which H5N8 viruses have been isolated overall provide evidence for indirect migratory connections between East Asia and Western Europe and between East Asia and North America. This study is useful for better understanding the role of wild birds in the global epidemiology of H5N8 viruses. The need for sampling large numbers of wild birds for the detection of H5N8 virus and H5N8-virus-specific antibodies in a variety of species globally is highlighted, with specific emphasis in north-eastern Europe, Russia and northern China.

According to the green wave hypothesis, herbivores follow the flush of spring growth of forage plants during their spring migration to northern breeding grounds. In this study we compared two green wave indices for predicting the timing of the spring migration of avian herbivores: the satellite-derived green wave index (GWI), and an index of the rate of acceleration in temperature (GDDjerk). The GWI was calculated from MODIS normalized difference vegetation index (NDVI) satellite imagery and GDDjerk from gridded temperature data using products from the global land data assimilation system (GLDAS). To predict the timing of arrival at stopover and breeding sites, we used four years (2008–2011) of tracking data from 12 GPS-tagged barnacle geese, a long-distance herbivorous migrant, wintering in the Netherlands, breeding in the Russian Arctic. The stopover and breeding sites for these birds were identified and the relations between date of arrival with the date of 50% GWI and date of peak GDDjerk at each site were analyzed using mixed effect linear regression. A cross-validation method was used to compare the predictive accuracy of the GWI and GDDjerk indices. Significant relationships were found between the arrival dates at the stopover and breeding sites for the dates of 50% GWI as well as the peak GDDjerk (p < 0.01). The goose arrival dates at both stopover and breeding sites were predicted more accurately using GWI (R2cv = 0.68, RMSDcv = 5.9 and R2cv= 0.71, RMSDcv = 3.9 for stopover and breeding sites, respectively) than GDDjerk. The GDDjerk returned a lower accuracy for prediction of goose arrival dates at stopover ( R2cv = 0.45, RMSDcv = 7.79) and breeding sites (R2cv = 0.55, RMSDcv = 4.93). The positive correlation between the absolute residual values of the GDDjerk model and distance to the breeding sites showed that this index is highly sensitive to latitude. This study demonstrates that the satellite-derived green wave index (GWI) can accurately predict the timing of goose migration, irrespective of latitude and therefore is suggested as a reliable green wave index for predicting the timing of avian herbivores spring migration.

Animal movements may contribute to the spread of pathogens. In the case of avian influenza virus, [migratory] birds have been suggested to play a role in the spread of some highly pathogenic strains (e.g. H5N1, H5N8), as well as their low pathogenic precursors which circulate naturally in wild birds. For a better understanding of the emergence and spread of both highly pathogenic (HPAIV) and low pathogenic avian influenza virus (LPAIV), the potential effects of LPAIVs on bird movement need to be evaluated. In a key host species, the mallard Anas platyrhynchos, we tested whether LPAIV infection status affected daily local (< 100 m) and regional (> 100 m) movements by comparing movement behaviour 1) within individuals (captured and sampled at two time points) and 2) between individuals (captured and sampled at one time point). We fitted free-living adult males with GPS loggers throughout the autumn LPAIV infection peak, and sampled them for LPAIV infection at logger deployment and at logger removal on recapture. Within individuals, we found no association between LPAIV infection and daily local and regional movements. Among individuals, daily regional movements of LPAIV infected mallards in the last days of tracking were lower than those of non-infected birds. Moreover, these regional movements of LPAIV infected birds were additionally reduced by poor weather conditions (i.e. increased wind and/or precipitation and lower temperatures). Local movements of LPAIV infected birds in the first days of tracking were higher when temperature decreased. Our study thus demonstrates that bird-assisted dispersal rate of LPAIV may be lower on a regional scale than expected on the basis of the movement behaviour of non-infected birds. Our study underlines the importance of understanding the impact of pathogen infection on host movement in order to assess its potential role in the emergence and spread of infectious diseases.

Plant populations in fragmented ecosystems rely largely on internal dispersal by animals. To unravel the mechanisms underlying this mode of dispersal, an increasing number of experimental feeding studies is carried out. However, while physical activity is known to affect vertebrate digestive processes, almost all current knowledge on mechanisms of internal seed dispersal has been obtained from experiments with resting animals. We investigated how physical activity of the mallard (Anas platyrhynchos), probably the quantitatively most important biotic dispersal agent in aquatic habitats in the entire Northern Hemisphere, affects gut passage survival and retention time of ingested plant seeds. We fed seeds of nine common wetland plants to mallards trained to subsequently swim for 6 hours in a flume tank at different swimming speeds (activity levels). We compared gut passage survival and retention times of seeds against a control treatment with mallards resting in a conventional dry cage. Intact gut passage of seeds increased significantly with mallard activity (up to 80% in the fastest swimming treatment compared to the control), identifying reduced digestive efficiency due to increased metabolic rates as a mechanism enhancing the dispersal potential of ingested seeds. Gut passage speed was modestly accelerated (13% on average) by increased mallard activity, an effect partly obscured by the interaction between seed retention time and probability of digestion. Gut passage acceleration will be more pronounced in digestion-resilient seed species, thereby modulating their dispersal distances. Our findings imply that seed dispersal potential by mallards calculated from previous experiments with resting birds is highly underestimated, while dispersal distances may be overestimated for some plant species. Similar effects of physical activity on digestive efficiency of mammals suggests that endozoochorous dispersal of plant seeds by vertebrates is more effective and plays a quantitatively more important ecological role in both terrestrial and aquatic ecosystems than previously thought.

Recently, Lévy walks have been put forward as a new paradigm for animal search and many cases have been made for its presence in nature. However, it remains debated whether Lévy walks are an inherent behavioural strategy or emerge from the animal reacting to its habitat. Here, we demonstrate signatures of Lévy behaviour in the search movement of mud snails (Hydrobia ulvae) based on a novel, direct assessment of movement properties in an experimental set-up using different food distributions. Our experimental data uncovered clusters of small movement steps alternating with long moves independent of food encounter and landscape complexity. Moreover, size distributions of these clusters followed truncated power laws. These two findings are characteristic signatures of mechanisms underlying inherent Lévy-like movement. Thus, our study provides clear experimental evidence that such multi-scale movement is an inherent behaviour rather than resulting from the animal interacting with its environment.

The increasing spatiotemporal accuracy of Global Navigation Satellite Systems (GNSS) tracking systems opens the possibility to infer animal behaviour from tracking data. We studied the relationship between high-frequency GNSS data and behaviour, aimed at developing an easily interpretable classification method to infer behaviour from location data. Behavioural observations were carried out during tracking of cows (Bos Taurus) fitted with high-frequency GPS (Global Positioning System) receivers. Data were obtained in an open field and forested area, and movement metrics were calculated for 1 min, 12 s and 2 s intervals. We observed four behaviour types (Foraging, Lying, Standing and Walking). We subsequently used Classification and Regression Trees to classify the simultaneously obtained GPS data as these behaviour types, based on distances and turning angles between fixes. GPS data with a 1 min interval from the open field was classified correctly for more than 70% of the samples. Data from the 12 s and 2 s interval could not be classified successfully, emphasizing that the interval should be long enough for the behaviour to be defined by its characteristic movement metrics. Data obtained in the forested area were classified with a lower accuracy (57%) than the data from the open field, due to a larger positional error of GPS locations and differences in behavioural performance influenced by the habitat type. This demonstrates the importance of understanding the relationship between behaviour and movement metrics, derived from GNSS fixes at different frequencies and in different habitats, in order to successfully infer behaviour. When spatially accurate location data can be obtained, behaviour can be inferred from high-frequency GNSS fixes by calculating simple movement metrics and using easily interpretable decision trees. This allows for the combined study of animal behaviour and habitat use based on location data, and might make it possible to detect deviations in behaviour at the individual level.

1.Herbivorous birds are hypothesized to migrate in spring along a seasonal gradient of plant profitability towards their breeding grounds (green wave hypothesis). For Arctic-breeding species in particular, following highly profitable food is important, so that they can replenish resources along the way and arrive in optimal body condition to start breeding early. 2.We compared the timing of migratory movements of Arctic-breeding geese on different flyways to examine whether flyways differed in the predictability of spring conditions at stopovers, and whether this was reflected in the degree to which birds were following the green wave. 3.Barnacle geese (Branta leucopsis) were tracked with solar Argos/GPS PTTs from their wintering grounds to breeding sites in Greenland (N = 7), Svalbard (N = 21) and the Barents Sea (N = 12). The numerous stopover sites of all birds were combined into a set of 16 general stopover regions. 4.The predictability of climatic conditions along the flyways was calculated as the correlation and slope between onsets of spring at consecutive stopovers. These values differed between sites, mainly because of the presence or absence of ecological barriers. Goose arrival at stopovers was more closely tied to the local onset of spring when predictability was higher and when geese attempted breeding that year. 5.All birds arrived at early stopovers after the onset of spring and arrived at the breeding grounds before the onset of spring, thus overtaking the green wave. This is in accordance with patterns expected for capital breeders: first they must come into condition; at intermediate stopovers arrival with the food quality peak is important to stay in condition and at the breeding grounds early arrival is favoured so that hatching of young can coincide with the peak of food quality. 6.Our results suggest that a chain of correlations between climatic conditions at subsequent stopovers enables geese to closely track the green wave. However, the birds’ precision of migratory timing seems uninfluenced by ecological barriers, indicating partly fixed migration schedules. These might become non-optimal due to climate warming and preclude accurate timing of long-distance migrants in the future.

Understanding how climate change and increasing human impacts may exert pressure on ecosystems and threaten biodiversity requires efficient monitoring programs. Indicator species have been proposed as useful tools, and predators and their diet may be particularly suitable. The vast and remote arctic tundra represents a good case study as shifts in ecosystem states are presently occurring, and monitoring is a major challenge. Here we assess what stable isotopes reflecting the diet of the arctic fox, a widespread and highly flexible top predator, can contribute to effective monitoring of the vertebrate prey basis of Arctic tundra. We used data collected over 2–5 years from six sites in the Eurasian Arctic and Greenland. Stable isotope signatures of arctic fox winter fur reflected both spatial and temporal variability in the composition of the vertebrate prey basis. Clear contrasts were apparent in the importance of marine resources, as well as of small rodents and their multiannual density fluctuations. Some important resources could however not be separated because of confounding isotopic signatures. Moreover, except for preferred prey, the proportions of prey in the diet may not necessarily reflect the relative importance of species in the community of available prey. Knowing these limitations, we suggest that the arctic fox diet as inferred from stable isotopes could serve as one of several key targets in ecosystem-based monitoring programs.

Food-hoarding patterns range between larder hoarding (a few large caches) and scatter hoarding (many small caches), and are, in essence, the outcome of a hoard size–number trade-off in pilferage risk. Animals that scatter hoard are believed to do so, despite higher costs, to reduce loss of cached food to competitors against which they cannot defend their food reserves (henceforth: superior competitors). We tested the underlying assumption that the cost of having more caches under scatter hoarding, thus increasing the likelihood of cache encounter by superior competitors, is outweighed by the benefit of having small caches that are less likely to be detected upon encounter by superior competitors. We carried out a controlled experiment in which we distributed a fixed number of acorns over a fixed number of patches within a fixed area, varying cache size and cache depth, thus mimicking alternative hoarding patterns. We then recorded cache pilferage by a fixed number of wild boar, a well-known pilferer of acorn caches. The time wild boar needed to pilfer the first cache was shortest for scatter hoarding, but the time needed to pilfer all caches was slightly longer for scatter hoarding than for larder hoarding. Overall, however, the rate of pilferage did not differ between scatter hoarding and larder hoarding, and was not affected by cache depth. We conclude that the effects of alternative hoarding patterns on reducing cache pilferage by wild boar were smaller than expected, and that superior competitors may thus not be important drivers of scatter hoarding. Instead, other factors, such as conspecific pilferage or the risk of cross-contamination of food items in large caches, which can also cause catastrophic loss of food reserves, may be more important drivers of scatter hoarding.

In clonal plants, vegetative parts may outcompete seeds in the absence of disturbance, limiting the build-up of genotypic diversity through repeated seedling recruitment (RSR). Herbivory may provide disturbance and trigger establishment of strong colonizers (seeds) at the expense of strong competitors (clonal propagules). In the clonal aquatic fennel pondweed (Potamogeton pectinatus), two distinct herbivore guilds may modify the dynamics of propagation. In winter, Bewick's swans may deplete patches of tubers, promoting seedling establishment in spring. In summer, seed consumption by waterfowl can reduce the density of viable seeds but grazing may also reduce tuber production and hence facilitate seedling establishment. This study is among the first to experimentally test herbivore impact on plant genotypic diversity. We assess the separate and combined effects of both herbivore guilds on genotypic diversity and structure of fennel pondweed beds. Using microsatellites, we genotyped P. pectinatus from an exclosure experiment and assessed the contribution of herbivory, dispersal, and sexual reproduction to the population genetic structure. Despite the predominance of clonal propagation in P. pectinatus, we found considerable genotypic diversity. Within the experimental blocks, kinship among genets decreased with geographic distance, clearly identifying a role for RSR in the maintenance of genotypic diversity within the fennel pondweed beds. However, over a period of 5 years, none of the herbivory treatments affected genotypic diversity. Hence, sexual reproduction on a local scale is important in this putatively clonal plant and possibly sufficient to ensure a relatively high genotypic diversity even in the absence of herbivores. Although we cannot preclude a role of herbivory in shaping genotypic diversity of a clonal plant, after 5 years of exclusion of the two investigated herbivore guilds no measurable effect on genotypic diversity was detected

The abundance of primary producers is controlled by bottom-up and top-down forces. Despite the fact that there is consensus that the abundance of freshwater macrophytes is strongly influenced by the availability of resources for plant growth, the importance of top-down control by vertebrate consumers is debated, because field studies yield contrasting results. We hypothesized that these bottom-up and top-down forces may interact, and that consumer impact on macrophyte abundance depends on the nutrient status of the water body. To test this hypothesis, experimental ponds with submerged vegetation containing a mixture of species were subjected to a fertilization treatment and we introduced consumers (mallard ducks, for 8 days) on half of the ponds in a full factorial design. Over the whole 66-day experiment fertilized ponds became dominated by Elodea nuttallii and ponds without extra nutrients by Chara globularis. Nutrient addition significantly increased plant N and P concentrations. There was a strong interactive effect of duck presence and pond nutrient status: macrophyte biomass was reduced (by 50 %) after the presence of the ducks on fertilized ponds, but not in the unfertilized ponds. We conclude that nutrient availability interacts with top-down control of submerged vegetation. This may be explained by higher plant palatability at higher nutrient levels, either by a higher plant nutrient concentration or by a shift towards dominance of more palatable plant species, resulting in higher consumer pressure. Including nutrient availability may offer a framework to explain part of the contrasting field observations of consumer control of macrophyte abundance.http://link.springer.com/journal/442

In the vast majority of migratory bird species studied so far, spring migration has been found to proceed faster than autumn migration. In spring, selection pressures for rapid migration are purportedly higher, and migratory conditions such as food supply, daylength, and/or wind support may be better than in autumn. In swans, however, spring migration appears to be slower than autumn migration. Based on a comparison of tundra swan Cygnus columbianus tracking data with long-term temperature data from wheather stations, it has previously been suggested that this was due to a capital breeding strategy (gathering resources for breeding during spring migration) and/or to ice cover constraining spring but not autumn migration. Here we directly test the hypothesis that Bewick's swans Cygnus columbianus bewickii follow the ice front in spring, but not in autumn, by comparing three years of GPS tracking data from individual swans with concurrent ice cover data at five important migratory stop-over sites. In general, ice constrained the swans in the middle part of spring migration, but not in the first (no ice cover was present in the first part) nor in the last part. In autumn, the swans migrated far ahead of ice formation, possibly in order to prevent being trapped by an early onset of winter. We conclude that spring migration in swans is slower than autumn migration because spring migration speed is constrained by ice cover. This restriction to spring migration speed may be more common in northerly migrating birds that rely on freshwater resources.

Many migrating herbivores rely on plant biomass to fuel their life cycles and have adapted to following changes in plant quality through time. The green wave hypothesis predicts that herbivorous waterfowl will follow the wave of food availability and quality during their spring migration. However, testing this hypothesis is hampered by the large geographical range these birds cover. The satellite-derived normalized difference vegetation index (NDVI) time series is an ideal proxy indicator for the development of plant biomass and quality across a broad spatial area. A derived index, the green wave index (GWI), has been successfully used to link altitudinal and latitudinal migration of mammals to spatio-temporal variations in food quality and quantity. To date, this index has not been used to test the green wave hypothesis for individual avian herbivores. Here, we use the satellite-derived GWI to examine the green wave hypothesis with respect to GPS-tracked individual barnacle geese from three flyway populations (Russian n = 12, Svalbard n = 8, and Greenland n = 7). Data were collected over three years (2008–2010). Our results showed that the Russian and Svalbard barnacle geese followed the middle stage of the green wave (GWI 40–60%), while the Greenland geese followed an earlier stage (GWI 20–40%). Despite these differences among geese populations, the phase of vegetation greenness encountered by the GPS-tracked geese was close to the 50% GWI (i.e. the assumed date of peak nitrogen concentration), thereby implying that barnacle geese track high quality food during their spring migration. To our knowledge, this is the first time that the migration of individual avian herbivores has been successfully studied with respect to vegetation phenology using the satellite-derived GWI. Our results offer further support for the green wave hypothesis applying to long-distance migrants on a larger scale.

Ecological theory uses Brownian motion as a default template for describing ecological movement, despite limited mechanistic underpinning. The generality of Brownian motion has recently been challenged by empirical studies that highlight alternative movement patterns of animals, especially when foraging in resource-poor environments. Yet, empirical studies reveal animals moving in a Brownian fashion when resources are abundant. We demonstrate that Einstein's original theory of collision-induced Brownian motion in physics provides a parsimonious, mechanistic explanation for these observations. Here, Brownian motion results from frequent encounters between organisms in dense environments. In density-controlled experiments, movement patterns of mussels shifted from Lévy towards Brownian motion with increasing density. When the analysis was restricted to moves not truncated by encounters, this shift did not occur. Using a theoretical argument, we explain that any movement pattern approximates Brownian motion at high-resource densities, provided that movement is interrupted upon encounters. Hence, the observed shift to Brownian motion does not indicate a density-dependent change in movement strategy but rather results from frequent collisions. Our results emphasize the need for a more mechanistic use of Brownian motion in ecology, highlighting that especially in rich environments, Brownian motion emerges from ecological interactions, rather than being a default movement pattern.

1.Similar to other infectious diseases, the prevalence of low pathogenic avian influenza viruses (LPAIV) has been seen to exhibit marked seasonal variation. However, mechanisms driving this variation in wild birds have yet to be tested. We investigated the validity of three previously suggested drivers for the seasonal dynamics in LPAIV infections in wild birds: (i) host density, (ii) immunologically naïve young and (iii) increased susceptibility in migrants. 2.To address these questions, we sampled a key LPAIV host species, the mallard Anas platyrhynchos, on a small spatial scale, comprehensively throughout a complete annual cycle, measuring both current and past infection (i.e. viral and seroprevalence, respectively). 3.We demonstrate a minor peak in LPAIV prevalence in summer, a dominant peak in autumn, during which half of the sampled population was infected, and no infections in spring. Seroprevalence of antibodies to a conserved gene segment of avian influenza virus (AIV) peaked in winter and again in spring. 4.The summer peak of LPAIV prevalence coincided with the entrance of unfledged naïve young in the population. Moreover, juveniles were more likely to be infected, shed higher quantities of virus and were less likely to have detectable antibodies to AIV than adult birds. The arrival of migratory birds, as identified by stable hydrogen isotope analysis, appeared to drive the autumn peak in LPAIV infection, with both temporal coincidence and higher infection prevalence in migrants. Remarkably, seroprevalence in migrants was substantially lower than viral prevalence throughout autumn migration, further indicating that each wave of migrants amplified local AIV circulation. Finally, while host abundance increased throughout autumn, it peaked in winter, showing no direct correspondence with either of the LPAIV infection peaks. 5.At an epidemiologically relevant spatial scale, we provide strong evidence for the role of migratory birds as key drivers for seasonal epizootics of LPAIV, regardless of their role as vectors of these viruses. This study exemplifies the importance of understanding host demography and migratory behaviour when examining seasonal drivers of infection in wildlife populations.

Keywords: food intake rate; giving-up density; habitat switch; parental costs; social dominance After reproducing successfully, birds with extended parental care form family groups. Despite being the dominant social unit, such family groups have been reported to switch to alternative habitat earlier than adults without offspring, with potential negative carry-over effects for the next breeding season. Here we test a proposed mechanism for this earlier habitat switch, namely a low foraging efficiency in juveniles. Such a test is best performed under controlled conditions because in the field families may occupy food patches of a different quality than singles or pairs without young. We studied this mechanism in Bewick's Swans Cygnus columbianus bewickii, which trample (or ‘treadle’) for food buried in the sediment. The gross intake rate of juveniles was as low as 60% of that of adults, depending on the burial depth of the food. Trampling effort did not differ between age classes, but differences in intake rate were related to body size, suggesting that larger or heavier birds were trampling more efficiently. Corresponding giving-up densities in the field were calculated to be c. 60% higher for juveniles than for adults. Our findings are consistent with the hypothesis that the lower foraging efficiency of juveniles may be responsible for the segregation of family groups from adults without offspring.

Many migratory birds use a chain of stopover sites to fuel their migration. Under time-minimizing migration, fuelling time and giving-up density at stopovers are predicted to depend on fuelling conditions. Fluctuations in food accessibility likely lead to changes in fuelling conditions, which should in turn be reflected in fuelling time and giving-up density. During their migration, Bewick’s Swans Cygnus columbianus bewickii refuel on belowground tubers of Fennel Pondweed Potamogeton pectinatus in shallow lakes. We studied giving-up density and stop-over use (expressed in bird-days) of Bewick's Swans at an autumn stopover site (Lauwersmeer, The Netherlands) during 1995–2008, as dependent on local environmental conditions. High water levels were hypothesized to restrict access to tuber stocks. High water levels at the stopover site were predicted to lead to higher giving-up densities and less bird-days spent at the stopover. Annual variation in giving-up densities and number of bird-days was strongly associated with year-to-year differences in initial tuber biomass density and number of days with high water levels. As predicted, giving-up density increased and bird-days decreased with the number of days with high water level. We conclude that, in line with time-minimizing migration, changes in fuelling conditions may lead to underuse of a stopover site. Underuse of stopovers by migratory birds has been reported before but only in the sense that more food was left at stopover sites than at wintering sites. In contrast, in our case, dealing with a given stopover site, more food is left behind in some years than in other years.

1. The huge changes in population sizes of Arctic-nesting geese offer a great opportunity to study population limitation in migratory animals. In geese, population limitation seems to have shifted from wintering to summering grounds. There, in the Arctic, climate is rapidly changing, and this may impact reproductive performance, and perhaps population size of geese, both directly (e.g. by changes in snow melt) or indirectly (e.g. by changes in trophic interactions). 2. Dark-bellied brent geese (Branta bernicla bernicla L.) increased 20-fold since the 1950s. Its reproduction fluctuates strongly in concert with the 3-year lemming cycle. An earlier analysis, covering the growth period until 1988, did not find evidence for density dependence, but thereafter the population levelled off and even decreased. The question is whether this is caused by changes in lemming cycles, population density or other factors like carry-over effects. 3. Breeding success was derived from proportions of juveniles. We used an information-theoretical approach to investigate which environmental factors best explained the variation in breeding success over nearly 50 years (1960–2008). We subsequently combined GLM predictions of breeding success with published survival estimates to project the population trajectory since 1991 (year of maximum population size). In this way, we separated the effects of lemming abundance and population density on population development. 4. Breeding success was mainly dependent on lemming abundance, the onset of spring at the breeding grounds, and the population size of brent goose. No evidence was found for carryover effects (i.e. effects of conditions at main spring staging site). Negative density dependence was operating at a population size above c. 200 000 individuals, but the levelling off of the population could be explained by faltering lemming cycles alone. 5. Lemmings have long been known to affect population productivity of Arctic-nesting migratory birds and, more recently, possibly population dynamics of resident bird species, but this is the first evidence for effects of lemming abundance on population size of a migratory bird species. Why lemming cycles are faltering in the last two decades is unclear, but this may be associated with changes in winter climate at Taimyr Peninsula (Siberia).

Many migratory herbivores seem to follow the flush of plant growth during migration in order to acquire the most nutrient-rich plants. This has also been hypothesized for arctic-breeding geese, but so far no test of this so-called green wave hypothesis has been performed at the individual level. During four years, a total of 30 greater white-fronted geese Anser albifrons albifrons was tracked using GPS transmitters, of which 13 yielded complete spring migration tracks. From those birds we defined stopover sites and related the date of arrival at each of these stopovers to temperature sum (growing degree days, GDD), snow cover, accumulated photoperiod and latitude. We found that geese arrived at spring stopovers close to the peak in GDD jerk; the ‘jerk’ is the third derivative, or the rate of change in acceleration, and GDD jerk maxima therefore represent the highest acceleration of daily temperature per site. Day of snow melt also correlated well with the observed arrival of the geese. Factors not closely related to onset of spring, i.e. accumulated photoperiod and latitude, yielded poorer fits. A comparison with published data revealed that the GDD jerk occurs 1–2 weeks earlier than the onset of spring derived from NDVI, and probably represents the very start of spring growth. Our data therefore suggest that white-fronted geese track the front of the green wave in spring

Animals foraging in groups may benefit from a faster detection of food and predators, but competition by conspecifics may reduce intake rate. Competition may also alter the foraging behaviour of individuals, which can be influenced by dominance status and the way food is distributed over the environment. Many studies measuring the effects of competition and dominance status have been conducted on a uniform or highly clumped food distribution, while in reality prey distributions are often in-between these two extremes. The few studies that used a more natural food distribution only detected subtle effects of interference and dominance. We therefore conducted an experiment on a natural food distribution with focal mallards Anas platyrhynchos foraging alone and in a group of three, having a dominant, intermediate or subordinate dominance status. In this way, the foraging behaviour of the same individual in different treatments could be compared, and the effect of dominance was tested independently of individual identity. The experiment was balanced using a 4 × 4 Latin square design, with four focal and six non-focal birds. Individuals in a group achieved a similar intake rate (i.e. number of consumed seeds divided by trial length) as when foraging alone, because of an increase in the proportion of time feeding (albeit not significant for subordinate birds). Patch residence time and the number of different patches visited did not differ when birds were foraging alone or in a group. Besides some agonistic interactions, no differences in foraging behaviour between dominant, intermediate and subordinate birds were measured in group trials. Possibly group-foraging birds increased their feeding time because there was less need for vigilance or because they increased foraging intensity to compensate for competition. This study underlines that a higher competitor density does not necessarily lead to a lower intake rate, irrespective of dominance status.

1. Habitat use can influence individual performance in a wide range of animals, either immediately or through carry-over effects in subsequent seasons. Given that many animal species also show consistent individual differences in reproductive success, it seems plausible that individuals may have consistent patterns of habitat use representing individual specializations, with concomitant fitness consequences. 2. Stable-carbon isotope ratios from a range of tissues were used to discern individual consistency in habitat use along a terrestrial–aquatic gradient in a long-distance migrant, the Bewick’s swan (Cygnus columbianus bewickii). These individual specialisations represented

Animal distribution is usually predicted from the spatial variation in food biomass, whereas foraging theory commonly uses net energy intake rate as the currency to be maximized. We tested whether net energy intake rate better predicted the distribution and abundance of tundra swans than food biomass. In a shallow lake, we mapped the density of sago pondweed tubers during 2 years, and calculated the foraging benefits and costs to tundra swans. Swan residence was expressed in bird-days, i.e. the sum of daily counts. We used four measures of increasing complexity to predict bird-days per inlet: total food biomass (B), total food biomass above giving-up density (B+), total accessible food biomass above giving-up density (aB+), and total achievable net energy intake rate above giving-up energy intake rate (NEI+). Considering both years, observed bird-days of inlets correlated only with NEI+, and not with B, B+, or aB+. In both years, our predictions of bird-days based on the NEI+model better matched observed relationships than the predictions of the other three models. Our case study suggests that in heterogeneous wetlands, correcting for givingup density, food accessibility and foraging costs may be necessary in order to predict bird distribution and abundance.

Individuals foraging in groups constantly need to make decisions, such as when to leave a group, when to join a group, and when to move collectively to another feeding site. In recent years, it has become evident that personality may affect these foraging decisions, but studies where individuals are experimentally forced into different roles are still absent. Here, we forced individual barnacle geese, Branta leucopsis, differing in boldness scores, either in a joining or in a leaving role in a feeding context. We placed a food patch at the far end of a test arena and measured the arrival latency and number of visits of individuals to the patch either in the presence of a companion that was confined near the food patch (“joining context”) or in the presence of a companion that was confined away from the food patch (“leaving context”). We also ran trials without a companion (“nonsocial context”). Bolder individuals arrived more quickly than shyer individuals in the “leaving” context, but there was no effect of boldness in the “joining” context, suggesting that boldness differences are important in explaining variation in leaving behavior but not in joining behavior. The difference in arrival latency between the “joining” and non-social context increased with decreasing boldness score, suggesting that shyer individuals are more responsive to the presence of other individuals (i.e., social facilitation). These results indicate that individual differences in boldness play a role in patch choice decisions of group-living animals, such as when to leave a flock and when to join others at a patch.

Measuring the food left in experimental trays when study organisms cease feeding on them [so-called giving-up densities (GUDs)] is an accepted technique for assessing predation risk and disturbance. However, in natural settings, accessibility and energetic harvest costs may vary spatially, and GUDs may be confounded. In this study, we assessed whether at a heterogeneous site, non-experimental GUDs could reveal the effect of disturbance. We measured initial and GUDs of tubers of Fennel Pondweed Potamogeton pectinatus, which form here the exclusive food source of Bewick's Swans Cygnus columbianus bewickii during their migratory stopover. We calculated giving-up net energy intake rates (GUNs) by correcting for biomass accessibility and foraging costs. The study area was a shallow lake consisting of nine creeks, three of which were open to the public (i.e. disturbed). GUDs in creeks open or closed to the public were not significantly different. In contrast, GUNs were generally higher in creeks open to the public, after correcting for initial net energy intake rate. The results suggest that natural GUDs may not reflect the effects of disturbance in heterogeneous habitats. When environmental differences are large within a site, GUNs may be a useful alternative as a behavioural indicator.

We agree with Jansen et al. that a composite movement model provides a better statistical description of mussel movement than any simple movement strategy. This does not undermine the take-home message of our paper, which addresses the feedback between individual movement patterns and spatial complexity. Simple movement strategies provide more insight in the eco-evolutionary analysis and are therefore our model of choice.

Deeper burial of bulbs and tubers has been suggested as an escape against below-ground herbivory by vertebrates, but experimental evidence is lacking. As deep propagule burial can incur high costs of emergence after dormancy, burial depth may represent a trade-off between sprouting survival and herbivore avoidance. We tested whether burial depth of subterraneous tubers is a flexible trait in fennel pondweed (Potamogeton pectinatus), facing tuber predation by Bewick's swans (Cygnus columbianus bewickii) in shallow lakes in winter. In a four-year experiment involving eight exclosures, winter herbivory by swans and all vertebrate summer herbivory were excluded in a full-factorial design; we hence controlled for aboveground vertebrate herbivory in summer, possibly influencing tuber depth. Tuber depth was measured each September before swan arrival and each March before tuber sprouting. In accordance with our hypothesis, tuber depth in September decreased after excluding Bewick's swans in comparison to control plots. The summer exclosure showed an increase in tuber biomass and the number of shallow tubers, but not a significant effect on the mean burial depth of tuber mass. Our results suggest that a clonal plant like P. pectinatus can tune the tuber burial depth to predation pressure, either by phenotypic plasticity or genotype sorting, hence exhibiting flexible avoidance by escape. We suggest that a flexible propagule burial depth can be an effective herbivore avoidance strategy, which might be more widespread among tuber forming plant species than previously thought.

Aquatically foraging Bewick’s Swans Cygnus bewickii have been repeatedly reported to be accompanied by diving ducks, but the exact nature of this relationship is unclear. Based on field observations, we found a strong correlation between the number of foraging swans and the number of foraging Common Pochards Aythya ferina and Tufted Ducks Aythya fuligula at our study site (Lauwersmeer, the Netherlands), with the median ratio being close to one diving duck per swan. To determine whether the association was kleptoparasitic or commensal, we measured in a series of experimental trials the instantaneous intake rates of Bewick’s Swans and Common Pochards when foraging alone and when foraging together. On average, pochards in company with a swan attained twice the instantaneous intake rate as compared with foraging alone. This was mainly due to gleaning food items that floated away as a result of the swan’s trampling activity, and was not due to stealing: in fact, food intake rates of swans were not affected by the presence of a pochard.

In this paper, we develop a model to evaluate the capacity of accommodation areas for overwintering waterfowl, at a large spatial scale. Each day geese are distributed over roosting sites. Based on the energy minimization principle, the birds daily decide which surrounding fields to exploit within the reserve boundaries. Energy expenditure depends on distance to the roost and weather conditions. Food intake rate is determined by functional responses, and declines with consumption. A shortage occurs when birds cannot fulfil their daily energy requirement. Most foraging takes place on pasture, with complementary feeding for some of the species on cereals and harvest remains. We applied the model to five waterfowl species overwintering in the Netherlands. From a comparison with field data, the model appears to produce realistic grazing pressures on pasture, especially for geese, and a realistic decline in sward height, but the use of arable fields is less in agreement with observations. For current goose and wigeon numbers, hardly any shortages are expected, but extrapolating the population increase observed during the last decade, considerable shortages are expected in the near future (2015). However, we find that several uncertainties may contribute to more severe shortages: a probabilistic (and therefore less optimal) choice of foraging location, a shorter maximum distance to the roost, and a lower effective availability of resources due to disturbances and other edge effects. Between species we find both competition and facilitation. Both type of interactions, as well as the spatial pattern of resource exploitation, are explained from functional responses and energetic costs of the species.

In this study we investigated the effect of summer bird herbivory on the belowground tuber formation of fennel pondweed (Potamogeton pectinatus L.). Cumulative grazing pressure of four waterfowl species (mute swans, mallards, gadwalls and coots) in the summer was calculated based on timing of grazing and body mass of the grazers. The resulting grazing pressures were significantly negatively correlated with mean autumn tuber biomass in three of the four years of study. Moreover, summer grazing pressures explained more of the variance in tuber densities than water depth, sediment particle size distribution or any interactions of these variables did in the same three years. We propose that herbivory early in the summer has the most substantial impact on the clonal reproduction of macrophytes. Herbivores with a high consumption rate and early congregation for moulting may be the key waterfowl species in diminishing propagule biomass. Hence, they may present pre-emptive, time-staggered competition to consumers of the belowground biomass in autumn, such as migratory swans and diving ducks.

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Ecological theory predicts that animal movement is shaped by its efficiency of resource acquisition. Focusing solely on efficiency, however, ignores the fact that animal activity can affect resource availability and distribution. Here, we show that feedback between individual behavior and environmental complexity can explain movement strategies in mussels. Specifically, experiments show that mussels use a Lévy walk during the formation of spatially patterned beds, and models reveal that this Lévy movement accelerates pattern formation. The emergent patterning in mussel beds, in turn, improves individual fitness. These results suggest that Lévy walks evolved as a result of the selective advantage conferred by autonomously generated, emergent spatial patterns in mussel beds. Our results emphasize that an interaction between individual selection and habitat complexity shapes animal movement in natural systems.

Previous studies of interference competition have shown an asymmetric effect on intake rate of foragers on clumped resources, with only subordinate individuals suffering. However, the food distributions in these studies were uniform or highly clumped, whereas in many field situations, food aggregation is intermediate. Here we investigated whether food distribution (i.e., uniform, slightly clumped, and highly clumped) affects the behavioral response of mallards foraging alone or competing with another. Although the amount of food was the same in all distributions, the mallards reached higher intake rates, visited fewer patches, and showed longer average feeding times in the highly clumped distribution. Competing mallards had lower intake rates on the slightly clumped than on the uniform or highly clumped food distributions. Subordinates generally visited more patches and had shorter feeding times per patch, but their intake rates were not significantly lower than those of dominants. Therefore, we propose that subordinates do not necessarily suffer from interference competition in terms of intake rate, but do suffer higher search costs. In addition, although dominants had significantly higher average feeding times on the best quality patches of the highly clumped food distribution, such an effect was not found in the slightly clumped distribution. These findings indicate that in environments where food is aggregated to a lesser extent, monopolization is not the best strategy for dominants. Our results suggest that interference experiments should use food distributions that resemble the natural situation animals are faced with in the field.

At least two distinct trade-offs are thought to facilitate higher diversity in productive plant communities under herbivory. Higher investment in defence and enhanced colonization potential may both correlate with decreased competitive ability in plants. Herbivory may thus promote coexistence of plant species exhibiting divergent life history strategies. How different seasonally tied herbivore assemblages simultaneously affect plant community composition and diversity is, however, largely unknown. Two contrasting types of herbivory can be distinguished in the aquatic vegetation of the shallow lake Lauwersmeer. In summer, predominantly above-ground tissues are eaten, whereas in winter, waterfowl forage on below-ground plant propagules. In a 4-year exclosure study we experimentally separated above-ground herbivory by waterfowl and large fish in summer from below-ground herbivory by Bewick’s swans in winter. We measured the individual and combined effects of both herbivory periods on the composition of the three-species aquatic plant community. Herbivory effect sizes varied considerably from year to year. In 2 years herbivore exclusion in summer reinforced dominance of Potamogeton pectinatus with a concomitant decrease in Potamogeton pusillus, whereas no strong, unequivocal effect was observed in the other 2 years. Winter exclusion, on the other hand, had a negative effect on Zannichellia palustris, but the effect size differed considerably between years. We suggest that the colonization ability of Z. palustris may have enabled this species to be more abundant after reduction of P. pectinatus tuber densities by swans. Evenness decreased due to herbivore exclusion in summer. We conclude that seasonally tied above- and below-ground herbivory may each stimulate different components of a macrophyte community as they each favoured a different subordinate plant species.

It has been suggested that herbivorous waterfowl may be important in shaping aquatic plant communities in shallow wetlands. As such, a shift from canopy forming pondweeds to bottom-dwelling charophytes in a formerly turbid pondweed dominated lake has been partly attributed to waterfowl herbivory. Here we study the separate and combined effects of both belowground herbivory in spring by whooper swans and Bewick ‘s swans, and grazing in summer by waterfowl and fish on the community composition in a shallow Baltic estuary during one year. The macrophyte community was dominated by charophytes (mainly Chara aspera) with Potamogeton pectinatus and Najas marina present as subdominants. Other species were rare. Both spring and summer herbivory had no effect on total plant biomass. However, P. pectinatus was more abundant in plots that were closed to spring and summer herbivores. N. marina was more abundant in grazed plots, whereas Chara spp. biomass remained unaffected. Probably belowground propagules of both C. aspera and P. pectinatus were consumed by swans but since C. aspera bulbils were numerous it may have compensated for the losses. P. pectinatus may not have fully recovered from foraging on tubers and aboveground biomass. Our results are in line with other studies in Chara dominated lakes, which found no effect of grazing on summer aboveground Chara biomass, whereas several studies report strong effects of herbivory in lakes dominated by P. pectinatus.

During periods of high energy demand an animal may be constrained by a physiological maximum to its energy intake rate. Predictions by allometric equations describing this maximum for endotherms were significantly surpassed during a few recent laboratory experiments on birds and mammals, being given access to food 24 h day−1. How relevant this is in the field remains to be assessed. We predicted that Bewick's swans Cygnus columbianus bewickii might surpass this maximum during stopover on their migration. We determined intake rate by measuring initial and final biomass density, and dividing the biomass difference by the feeding time required to reach this difference. This feeding time was given by the functional response. After conversion to daily energy intake rates, these exceeded the previously assumed maximum on two of the three stopover sites studied. The exception was a stopover site where daily foraging time was limited by the tidal cycle. Our study confirms that intake rates may exceed the formerly generally supposed maximum under natural conditions when foraging is possible day and night.

Animals foraging in groups can either search for food themselves (producing) or search for the food discoveries of other individuals (scrounging). Tactic use in producer–scrounger games is partly flexible but individuals tend to show consistency in tactic use under different conditions suggesting that personality might play a role in tactic use in producer–scrounger games. Here we studied the use of producing and scrounging tactics by bold and shy barnacle geese (Branta leucopsis), where boldness is a personality trait known to be repeatable over time in this species. We defined individuals as bold, shy or intermediate based on two novel object tests. We scored the frequency of finding food patches (the outcome of investing in producing) and joining patches (the outcome of investing in scrounging) by bold and shy individuals and their feeding time. Shy individuals had a higher frequency of joining than bold individuals, demonstrating for the first time that personality is associated with tactic use in a producer–scrounger game. Bold individuals tended to spend more time feeding than shy individuals. Our results highlight the importance of including individual behavioural variation in models of producer–scrounger games.

Interference can be defined as the reduction of intake rate caused by the presence of congeneric individuals. However, surrounding congeneric individuals may also accelerate food depletion. Therefore, it is difficult to quantify interference (contest) and exploitative (scramble) competition separately, particularly in the field. In an individual-based model, where food was immediately replenished in patches after a foraging bout, we isolated the effect of interference competition. In this way, we investigated how interference shapes the intake rates of flocking Bewick’s swans, Cygnus bewickii, consuming tubers of fennel pondweed, Potamogeton pectinatus. The model predicted that intake rate declines with increasing swan density, and that the rate of decline is greater in subordinate birds than in dominant birds. Subordinate birds suffered a large reduction in intake rate at high densities because they spent a large proportion of the time avoiding more dominant individuals. A major decline in intake rate of subordinates occurred close to the maximum swan densities observed in the field. We expected subordinates to leave high-density flocks and look for another foraging location: interference competition seemed to regulate the maximum swan flock density in the field. Hence, the mean population intake rate at realized densities was only slightly lower than in the absence of interference. As interference occurred mainly as avoidance behaviour, which is difficult to observe in the field, it might commonly remain cryptic for the observer. Our results may explain other field studies where interference competition seemed to be of lesser importance.

Submerged freshwater macrophytes decline with increasing eutrophication. This has consequences for ecosystem processes in shallow lakes and ponds as macrophytes can reduce algal blooms under eutrophic conditions. We hypothesize that the productivity of submerged vegetation, biomass change under eutrophication and the suppression of algal blooms may be affected by macrophyte community composition. To test our hypothesis, we established three macrophyte community types in 36 fishless experimental ponds: one dominated by the oligotrophic species Chara globularis, one dominated by the eutrophic species Potamogeton pectinatus and a diverse vegetation which became co-dominated by Elodea nuttallii and C. globularis, and we fertilized half of the ponds. The macrophyte communities produced different amounts of biomass and they responded differently to fertilization. The community dominated by Potamogeton produced the lowest overall biomass, but was not affected by nutrient addition. The communities dominated by Chara and co-dominated by Elodea and Chara produced more than four-fold the amount of biomass produced in Potamogeton communities under oligotrophic conditions, but were strongly negatively affected by nutrient addition. Phytoplankton abundance did not differ significantly among the plant community types, but showed large variation within community types. There was a significant negative relationship between spring macrophyte biomass and the probability of summer algal blooms. The occurrence of algal blooms coincided with low daphnid densities and high pH (>10). We conclude that the macrophyte community composition, characterized by the dominant species, strongly affected the amount of biomass production as well as the short-term response of the vegetation to nutrient enrichment. Macrophyte community composition had no direct effect on algal blooms, but can affect the occurrence of algal blooms indirectly as these occurred only in ponds with low (

1. Bottom-dwelling charophytes have been observed to replace canopy-forming pondweeds within a few years in de-eutrophied shallow lakes. Competition for bicarbonate (HCO3−) may explain this shift in vegetation dominance but inhibition of pondweeds by Chara spp. through direct competition has not been shown experimentally. 2. We tested whether charophytes inhibited growth of fennel pondweed (Potamogeton pectinatus) in the absence of belowground competition by growing plants in pots in mesocosms following a replacement series experimental design. To further understand the role of bicarbonate, we studied main and interactive effects of Chara, light and bicarbonate on P. pectinatus growth in a laboratory study. 3. Early in the mesocosm experiment, high charophyte densities had a negative effect on P. pectinatus cover at a time when bicarbonate levels were low. However, bicarbonate levels eventually converged to low levels in all treatments. At final harvest, both species exhibited lower biomasses at higher densities of conspecific pots, indicating that ultimately intraspecific competition was limiting. In a laboratory study, Chara inhibited P. pectinatus the most under a combination of high light and high bicarbonate concentrations, suggesting that Chara may negatively affect P. pectinatus by acting as a general nutrient sink. 4. Our results suggest that Chara growth can reduce bicarbonate levels, delaying but not preventing a P. pectinatus growth pulse. Given the recorded inhibition under ample bicarbonate supply, Chara’s ability to act as nutrient sink may contribute to the decline of P. pectinatus under Chara recovery in shallow lakes.

The use of social information is known to affect various important aspects of an individual’s ecology, such as foraging, dispersal and space use and is generally assumed to be entirely flexible and context dependent. However, the potential link between personality differences and social information use has received little attention. In this study, we studied whether use of social information was related to personality, using barnacle geese, Branta leucopsis, where boldness is a personality trait known to be consistent over time. We found that the use of social information decreased with increasing boldness score of the individuals. Individuals had lower feeding times when they did not follow the social information and this effect was unrelated to boldness score. When manipulating social information, thereby making it incorrect, individuals irrespective of their boldness score, learned that it was incorrect and ignored it. Our results show that social information use depends on the personality type of an individual, which calls for incorporation of these personality-related differences in studies of spatial distribution of animals in which social information use plays a role.

Effective conservation of important bird areas requires insight in the number of birds an area can support, and how this carrying capacity changes with habitat modifications. When food depletion is the dominant mechanism of competition, it should in principle be possible to calculate the total time foragers can spend per patch from their functional response (intake rate as a function of food density). However, in the field there are likely to be factors modulating the functional response. In this study previously published results of experiments on captive Bewick's swans were used to obtain functional responses of swans digging for tubers of Fennel pondweed on different foraging substrates: sandy and clayey sediment, and in shallow and deep water. In a field study, four 250×250 m sections belonging to different types (sandy–shallow, clayey–shallow, sandy–deep and clayey–deep) were delineated. Here tubers were sampled with sediment corers in three years, both before and after swan exploitation in autumn, and swans were observed and mapped from a hide in two of these years. Giving-up tuber biomass densities varied among sections. Substitution of these giving-up densities in the derived patch-type-specific functional responses yielded the quitting net energy intake rates in the four sections. As expected from the marginal value theorem, the quitting net energy intake rates did not vary among sections. Moreover, the observed foraging pressure (total foraging time per area) per patch type was in quantitative agreement with the integrated functional responses. These results suggest that in spatially heterogeneous environments, patch exploitation by foragers can be predicted from their functional responses after accounting for foraging substrate.

The degree to which vertebrate herbivores exploitatively compete for the same food plant may depend on the level of compensatory plant growth. Such compensation is higher when there is reduced density-dependent competition in plants after herbivore damage. Whether there is relief from competition may largely be determined by the life-history stage of plants under herbivory. Such stage-specific compensation may apply to seasonal herbivory on the clonal aquatic plant sago pondweed (Potamogeton pectinatus L.). It winters in sediments of shallow lakes as tubers that are foraged upon by Bewick's Swans (Cygnus columbianus bewickii Yarrell), whereas aboveground biomass in summer is mostly consumed by ducks, coots, and Mute Swans. Here, tuber predation may be compensated due to diminished negative density dependence in the next growth season. However, we expected lower compensation to summer herbivory by waterfowl and fish as density of aboveground biomass in summer is closely related to photosynthetic carbon fixation. In a factorial exclosure study we simultaneously investigated (1) the effect of summer herbivory on aboveground biomass and autumn tuber biomass and (2) the effect of tuber predation in autumn on aboveground biomass and tuber biomass a year later. Summer herbivory strongly influenced belowground tuber biomass in autumn, limiting food availability to Bewick's Swans. In contrast, tuber predation in autumn by Bewick's Swans had a limited and variable effect on P. pectinatus biomass in the following growth season. Whereas relief from negative density dependence largely eliminates effects of belowground herbivory by swans, aboveground herbivory in summer limits both above- and belowground plant biomass. Hence, there was an asymmetry in exploitative competition, with herbivores in summer reducing food availability for belowground herbivores in autumn, but not the other way around.

Decisions taken during migration can have a large effect on the fitness of birds. Migration must be accurately timed with food availability to allow efficient fueling but is also constrained by the optimal arrival date at the breeding site. The decision of when to leave a site can be driven by energetics (sufficient body stores to fuel flight), time-related cues (internal clock under photoperiodic control), or external cues (temperature, food resources). An individual based model (IBM) that allows a mechanistic description of a range of departure decision rules was applied to the spring migration of pink-footed geese (Anser brachyrhynchus) from wintering grounds in Denmark to breeding grounds on Svalbard via 2 Norwegian staging sites. By comparing predicted with observed departure dates, we tested 7 decision rules. The most accurate predictions were obtained from a decision rule based on a combination of cues including the amount of body stores, date, and plant phenology. Decision rules changed over the course of migration with the external cue decreasing in importance and the time-related cue increasing in importance for sites closer to breeding grounds. These results are in accordance with descriptions of goose migration, following the "green-wave": Geese track the onset of plant growth as it moves northward in spring, with an uncoupling toward the end of the migration if time is running out. We demonstrate the potential of IBMs to study the possible mechanisms underlying stopover ecology in migratory birds and to serve as tools to predict consequences of environmental change.

Deep burial in the sediment of tubers of fennel pondweed (Potamogeton pectinatus) has been explained in terms of avoidance by escape against consumption by Bewick's swans (Cygnus columbianus bewickii) in autumn. We therefore expected changes in foraging pressure to ultimately result in a change in the tuber distribution across sediment depth. A trade-off underlies this idea: deep tubers are less accessible to swans but must be larger to meet the higher energy demands of sprouting in spring. To test this prediction, we compared tuber burial depth over a gradient of foraging pressure both across space and across time. Tuber samples were obtained after aboveground plant senescence but before arrival of Bewick's swans. First, we compared the current tuber bank depth profile in a shallow lake with high foraging pressure, the Lauwersmeer, with that in two wetlands with moderate and low foraging pressure. Second, we compared the current tuber burial in the Lauwersmeer with that in the early 1980s when exploitation by swans had just started there. In accordance with our hypothesis, we found significantly deeper burial of tubers under high consumption risk compared to low consumption risk, both when comparing sites and comparing time periods. Since tubers in effect only survive to the next spring, the observed differences in burial depth among sites and over time cannot be a direct result of tuber losses due to consumption by swans. Rather, these observations suggest adaptive responses in tuber burial related to foraging pressure from Bewick's swans in the recent past. We thus propose that fennel pondweed exhibits flexible avoidance by escape, of a kind rarely described for plants, where both phenotypic plasticity and genotype sorting may contribute to the observed differences in tuber burial.

For many animals, notably herbivores, plants are often an inadequate food source given the low content of protein and high content of C-rich material. This conception is mainly based on studies on ectotherms. The validity of this conception for endotherms is unclear given their much higher carbon requirements for maintenance energy metabolism than ectotherms. Applying stoichiometric principles, we hypothesized that endotherms can cope with diets with much higher (metabolizable) carbon to nitrogen ratios than ectotherms. Using empirical data on birds, eutherian mammals, marsupials and reptiles, we compiled and compared measurements and allometric equations for energy metabolism as well as nitrogen requirements. Our analysis supports our hypothesis that plants, and especially their leaves, are generally sufficiently rich in nitrogen to fulfil protein demands in endotherms, at least during maintenance conditions, but less so in ectotherms. This has important implications with respect to community functioning and the evolution of endothermy.

Organisms usually benefit from heterogeneous conditions, but, by doing so, may reduce the degree of heterogeneity. The question therefore arises how heterogeneity is maintained. We investigated within-year spatiotemporal patterns in a monospecific stand of a submerged plant (fennel pondweed, Potamogeton pectinatus), with the novelty that we distinguished between different forms of heterogeneity: spatial variance (the frequency distribution of densities) and spatial pattern (the spatial distribution of densities). We repeatedly measured plant biomass that was affected by swan predation, winter mortality, and summer regrowth. Spatial variance was enhanced mostly by swan foraging, despite the fact that swans appear to exploit patches to the same threshold level. Spatial pattern, which had vanished after swan foraging, reestablished due to spatial pattern in winter mortality and was further enhanced by plant regrowth. We found that variance and pattern each have their own temporal dynamics and are maintained by different biological processes. We therefore advocate that it is pivotal to distinguish between variance and pattern in the study of spatial heterogeneity.

It would be valuable to be able to infer the habitat quality of underwater-feeding birds and mammals from their diving or head-dipping behavior, especially when underwater sampling is difficult. In air-breathing animals that obtain their food by diving from the water surface to the bottom (or any specific depth), the underwater feeding time is expected to be a function of travel time and gain rate. Hence, for a given travel time, the underwater feeding time should reflect the (initial) quality of the patch where the animals forage. We applied this approach to estimate habitat quality of tundra swans digging for starch-rich belowground propagules of aquatic macrophytes on migratory stopover sites. Swans do not travel underwater but instead trample with their heads above water before dipping their heads, and the optimal dive model was adjusted for this. At an autumn staging site, habitat quality estimated from head-dipping time was not significantly different from locally measured habitat quality, but the uncertainty around the estimated quality was large. Significantly longer head-dipping time was recorded at a spring staging site, despite a shorter trampling time. This long underwater feeding time indicated that the habitat quality was lower, while the short trampling time was explained by more shallow burial of belowground propagules at the spring staging site than at the autumn staging site. Both features were corroborated with biomass data. It should be noted, however, that the precision of the indirect method of habitat assessment varies with habitat quality, being greater for low quality habitat, but probably still not very precise. Further research is needed to confirm the inferred low habitat quality of the spring staging site and to clarify how tundra swans complete their vernal migration.

The role of aquatic macrophytes in stimulating biodiversity and maintaining clear waters is currently undisputed. The management of (eutrophic) shallow waters is therefore often directed at (re-)establishing macrophyte domination. In contrast, the role of water birds has long been considered of minor importance for the functioning of fresh water ecosystems. Indeed, in terms of biomass and production, water birds constitute only a minor part of these systems. However, water birds may graze heavily on water plants under certain circumstances, and the question arises whether herbivorous water birds have an important indirect effect on shallow fresh water systems. Mainly illustrated with the interaction between Bewick’s Swans and Fennel Pondweed, we present data on the role that water plants may play in the life of water birds and how water birds may impact water plants’ fitness in terms of survival, production, dispersal and competitive ability. It appears that water plants may be crucial for water birds during periods of high-energy requirements, such as migration. Despite the plants’ costs associated with water bird grazing, the interaction between water birds and water plants varies in nature from an apparent predator–prey relationship to a mutually beneficial interaction depending on the context and the perspective. For the case of the Bewick’s Swan–Fennel Pondweed interaction, regular bird grazing is sustainable and may actually favour the plant’s dispersal. Thus, Bewick’s Swans themselves may in fact play a crucial role in establishing and maintaining the Fennel Pondweed rich staging sites between the swans’ wintering and breeding grounds, which are vital for the swans’ successful migration.

1. Heterogeneity in food abundance allows a forager to concentrate foraging effort in patches that are rich in food. This might be problematic when food is cryptic, as the content of patches is unknown prior to foraging. In such case knowledge about the spatial pattern in the distribution of food might be beneficial as this enables a forager to estimate the content of surrounding patches. A forager can benefit from this pre-harvest information about the food distribution by regulating time in patches and/or movement between patches. 2. We conducted an experiment with mallard Anas platyrhynchos foraging in environments with random, regular, and clumped spatial configurations of full and empty patches. An assessment model was used to predict the time in patches for different spatial distributions, in which a mallard is predicted to remain in a patch until its potential intake rate drops to the average intake rate that can be achieved in the environment. A movement model was used to predict lengths of interpatch movements for 3. Consistent with predictions, in the clumped distribution mallard spent less time in an empty patch when the previously visited neighbouring patch had been empty than when it had been full. This effect was not observed for the random distribution. This shows that mallard use pre-harvest information on spatial pattern to improve patch assessment. Patch assessment could not be evaluated for the regular distribution. 4. Movements that started in an empty patch were longer than movements that started in a full patch. Contrary to model predictions this effect was observed for all spatial distributions, rather than for the clumped distribution only. In this experiment mallard did not regulate movement in relation to pattern. 5. An explanation for the result that pre-harvest information on spatial pattern affected patch assessment rather than movement is that mallard move to the nearest patch where the expected intake rate is higher than the critical value, rather than to the patch where the highest intake rate is expected.

Large migratory birds may bring along stores in order to survive adverse conditions and produce a clutch upon arrival (‘capital breeders’) or they may acquire all the necessary resources on the breeding grounds (‘income breeders’). Whether birds are capital- or income-breeders may depend on the distance between the last stopover site and the breeding grounds and the length of the summer season. The degree of capital-breeding may therefore differ among flyway populations of the same species. I used migration speed as a proxy for the degree of capital-breeding, and compared the observed migration speeds of satellite-tracked Tundra Swans Cygnus columbianus from all four main flyway populations with those predicted by an allometric model for 6-kg waterfowl. Average overall spring migration speed of Tundra Swans was 52.2 km/d, close to that predicted under a capital-breeding strategy (i.e. carrying the stores for the whole clutch and allowing for 8 days of starvation). This migration speed was in accordance with the speed at which the ice retreats in spring. However, the energy stores for clutch formation and survival on the breeding grounds may be primarily put on at the last stopover. Tundra Swans (‘Whistling Swans’) in the Nearctic, and especially in the Western Nearctic, make use of the rapid advancement of spring on their last leg and accelerate their migration speed accordingly, and so probably rely on income-breeding. In the Western Palearctic, Tundra Swans (‘Bewick’s Swans’) are faced with a slow advancement of spring on their last track, and they may fare better by building up stores on the last stopover and bring these to the breeding grounds. This may not be an option for swans in the Eastern Palearctic, as they are confronted with a very long last leg. The variation in migration speeds on the last leg suggests a large variation in the degree of capital-breeding among flyway populations of Tundra Swans.

We measured the patch use behaviour of Bewick's swans (Cygnus columbianus bewickii) feeding on below ground tubers of fennel pondweed (Potamogeton pectinatus). We compared the swans' attack rates, foraging costs and giving-up densities (GUDs) in natural and experimental food patches that differed in water depth. Unlike most studies that attribute habitat-specific differences in GUDs to predation risk, food quality or foraging substrate, we quantified the relative importance of energetic costs and accessibility. Accessibility is defined as the extent to which the animal's morphology restricts its harvest of all food items within a food patch. Patch use behaviours were measured at shallow (ca 0.4 m) and deep (ca 0.6 m) water depths on sandy sediments. In a laboratory foraging experiment, when harvesting food patches, the swan's attack rate (m3 s1) did not differ between depths. In deep water the energetic costs of surfacing, feeding and trampling were 1.13 to 1.21 times higher than in shallow water with a tendency to spend relatively more time trampling, the most expensive activity. Taking time allocation as measured in the field into account, foraging in deep water was 1.26 times as expensive as in shallow water. In the lake the GUD in shallow water was on average 12.9 g m2. If differences in energetic costs were the only factor determining differences in GUDs, then the deep water GUD should be 14.2 g m2. Instead, the mean GUD in deep water was 20.2 g m2, and therefore energetic costs explain just 18% of the difference in GUDs. At deep sites, 24% of tuber biomass was estimated to be out of reach, and we calculated a maximum accessible foraging depth of 0.86 m. This is close to the published 0.84 m based on body measurements. A laboratory experiment with food offered at a depth of 0.89 m confirmed that it was just out of reach. The agreement between calculated and observed maximum accessible foraging depths suggests that accessibility largely explains the remaining difference in GUDs with depth, and it confirms the existence of partial prey refuges in this system.

1. The carrying capacity of a site for migratory water birds, expressed in bird-days, can be of particular conservation value. Several attempts have been made to model this carrying capacity using ideal free distribution models such as, for instance, depletion models, in which the distribution is fully determined by exploitative competition. 2. In the tests of depletion models carried out so far, no alternative models were compared; rather, one specific model was tested. We tested whether bird-days were more in accordance with birds depleting the food resource (a1) until a critical food density which just enabled survival or (a2) until a threshold food density which renders the site as profitable as an alternative site; and birds (b1) satisfying their daily requirements or (b2) maximizing daily intake. 3. We studied Bewick's swans feeding on below-ground tubers of fennel pondweed in one part of an autumn staging site. In most years between 1995 and 2005, we measured tuber biomass densities around September, November and March, and counted swans daily during their stopover in October. 4. The best fit between observed and predicted bird-days was obtained by assuming that the swans were maximizing their daily intake and depleting the tubers until a threshold biomass density (which yielded the same energetic return as the alternative food source after accounting for a small part of the initial tuber biomass being out of reach of the swans). Also in line with daily intake maximization, the daily feeding time did not differ from 10 h day1, the value predicted for Bewick's swans b 5. Our results suggests that the applicable model to calculate carrying capacity may depend strongly on whether birds use a site to stopover or to winter, because it determines whether the birds are more likely to use a threshold or critical food density, and to behave as energy maximizers or satisficers.

Browsing may lead to an induced resistance or susceptibility of the plant to the herbivore. We tested the effect of winter browsing by Eurasian beavers (Castor fiber L., 1758) on food quality of holme willows (Salix dasyclados Wimm.) in and after the following growth season. Shrubs were pruned in February, and new shoots from these (cut) shrubs were compared with those of untreated (uncut) ones in May and November. The shoots were analysed for dry matter, nitrogen, acid detergent fibre, and total phenolics. In May, the leaves from the cut treatment had a better food quality (more water, more nitrogen, and less phenolics) than those from the uncut one. There was in part also a systemic response, with lower total phenolics in both the cut and untreated parts of pruned shrubs (uncut–cut) than in the uncut shrubs. In November, we did not find significant differences in biochemistry of bark among cut, uncut, or uncut–cut treatments. These results are in accordance with a cafeteria experiment in the field: in May the beavers preferred shoots from the cut treatment, but in November they showed no preference. The results suggest that willows invest in compensatory growth rather than a defence response early in the regrowing phase.

The efficiency in which high-density food patches are found is determined by the way foragers move between patches. In this study we explore the effect of social interactions on the foraging path, in particular the distance moved between patches. We studied Bewick’s Swans Cygnus columbianus bewickii that foraged on belowground tubers of Fennel Pondweed Potamogeton pectinatus. We accurately mapped the foraging path of individual swans and determined the distances between visited patches. 24% of inter-patch movements are associated with social interactions. When a swan retreats from a patch because it is chased away by another swan, it moves a significantly larger distance to a patch than if the movement is not associated with a social interaction. Such longer movements are thought to reduce the rate at which high-density patches are encountered, and thus the energy gain rate a swan can achieve. We observed a strong social hierarchy in which families are dominant over pairs and singletons, and pairs are dominant over singletons, which represents a producer–scrounger model. Singletons were most frequently observed to retreat from patches and are consequently thought to achieve the lowest gain rates, and also as a result exhibit the slowest speed of movement between patches. However singletons might partly compensate for more frequent retreats from patches by moving larger distances to arrive at the front edge of a flock where they encounter unexploited resources.

We tested whether Tundra Swans use information on the spatial distribution of cryptic food items (belowground Sago pondweed tubers) to shape their movement paths. In a continuous environment, swans create their own food patches by digging craters, which they exploit in several feeding bouts. Series of short (1 m). Tuber biomass densities showed a positive spatial auto-correlation at a short distance (25 g/m2) and to a more distant patch (at 7–8 m) if the food density in the current patch had been low (3 m) from a low-density patch and a short distance (

Learning animals are predicted to use a flexible patch-leaving threshold (PLT) while foraging in a depletable environment under exploitative competition. This prediction was tested in flock-feeding Bewick's swans (Cygnus columbianus bewickii) depleting hidden tubers of fennel pondweed (Potamogeton pectinatus) in a two-dimensional, continuous environment. The swans' patch residence time was measured by combining recordings of the foraging behaviour and movement paths. The tuber biomass density was measured before and after the period of exploitation, using the presumable foraging window of the swans as the scale of measurement. Swan foraging was simulated in order to predict the effects of flexible and fixed PLTs, respectively, on the patch residence time and the spatial heterogeneity of the tuber biomass density. Flexible PLTs were predicted to lead to short and decreasing patch residence times and a decrease in the coefficient of variation in tuber biomass densities, whereas the reverse was generally the case for fixed PLTs. Observed patch residence times did not decrease with time and were intermediate between those predicted for swans with flexible and fixed PLTs. Furthermore, an increase of the coefficient of variation in the tuber biomass density was observed. Given the observed giving-up biomass densities the most likely model was one with swans with a fixed rather than a flexible PLT. These results point at factors that may affect the spacing behaviour or constrain the use of a flexible PLT in swans.

Abstract If the food distribution contains spatial pattern, the food density in a particular patch provides a forager with information about nearby patches. Foragers might use this information to exploit patchily distributed resources profitably. We model the decision on how far to move to the next patch in linear environments with different spatial patterns in the food distribution (clumped, random, and regular) for foragers that differ in their degree of information. An ignorant forager is uninformed a [KEYWORDS: Area-restricted search ; Information state ; Foraging scale ; Marginal value theorem ; Optimal foraging strategies ; Prior knowledge ; Spatial heterogeneity]

The ability to respond to spatial heterogeneity in food abundance depends on the scale of the food distribution and the foraging scale of the forager. The aim of this study is to illustrate that a foraging scale exists, and that at larger scaled food distributions foragers benefit from the ability to subdivide a continuous (non-discrete) heterogeneous environment into profitable and non-profitable areas. We recorded search patterns of mallards Anas plathyrhynchos foraging in shallow water on cryptic prey items (millet seeds), distributed at different scales. A small magnet attached to the lower mandible allowed us to record in great detail the position and movements of the bill tip within a feeding tray underlain by magnet sensors. Instantaneous intake rate was determined in a subsequent experiment. We successfully determined the foraging scale (about 2×2 cm), defined as the scale above which foragers do respond (coarse scaled distribution) and below which foragers do not respond (fine scaled distribution) to spatial heterogeneity, by concentrating foraging effort within areas of high food density. A response resulted in a significantly higher intake rate, compared to a homogeneous distribution with an equal overall density. Unlike systematic search cell revisitation was common in trials, and at coarse scaled food distributions even slightly (but significantly) more frequently observed than predicted for random search. Mallards respond to food capture by restricting displacement (area restricted search) at food distributions that are considered to be clumped for the forager (large scaled coarse distributions). We argue that partitioning the environment at the foraging scale in itself could be a mechanism to concentrate foraging efforts within profitable areas, because mallard were able to respond to heterogeneity at coarse scaled food distributions even when non-clumped (i.e. without conducting area restricted search).

Climate change will alter freshwater ecosystems but specific effects will vary among regions and the type of water body. Here, we give an integrative review of the observed and predicted impacts of climate change on shallow lakes in the Netherlands and put these impacts in an international perspective. Most of these lakes are man-made and have preset water levels and poorly developed littoral zones. Relevant climatic factors for these ecosystems are temperature, ice-cover and wind. Secondary factors affected by climate include nutrient loading, residence time and water levels. We reviewed the relevant literature in order to assess the impact of climate change on these lakes. We focussed on six management objectives as bioindicators for the functioning of these ecosystems: target species, nuisance species, invading species, transparency, carrying capacity and biodiversity. We conclude that climate change will likely (i) reduce the numbers of several target species of birds; (ii) favour and stabilize cyanobacterial dominance in phytoplankton communities; (iii) cause more serious incidents of botulism among waterfowl and enhance the spreading of mosquito borne diseases; (iv) benefit invaders originating from the Ponto-Caspian region; (v) stabilize turbid, phytoplankton-dominated systems, thus counteracting restoration measures; (vi) destabilize macrophyte-dominated clear-water lakes; (vii) increase the carrying capacity of primary producers, especially phytoplankton, thus mimicking eutrophication; (viii) affect higher trophic levels as a result of enhanced primary production; (ix) have a negative impact on biodiversity which is linked to the clear water state; (x) affect biodiversity by changing the disturbance regime. Water managers can counteract these developments by reduction of nutrient loading, development of the littoral zone, compartmentalization of lakes and fisheries management. [KEYWORDS: Biodiversity ; Carrying capacity ; Invading species ; Nuisance species ; Temperature ; Transparency]

In temperate regions climate change has led to advances in plant phenology which may disrupt the synchrony between food availability and reproductive requirements of higher trophic levels. Because leaf quality generally drops with leaf maturation, for herbivorous animals a stoichiometric effect of climate change may be of greater importance than a shift in food quantity. We hypothesized that such a climate-related stoichiometric effect caused the low reproductive success of a beaver Castor fiber population that was translocated westward in Europe at the time of a rapid increase in spring temperatures. The staple food of these beavers was willow Salix spp. In 2003 willow bud burst occurred approximately one month earlier at the release site than at the site of origin. After bud burst, leaf quality, measured as nitrogen and phosphorus concentration, decreased in the course of spring in a predictable manner with accumulated degree-days. Nitrogen and phosphorus concentrations calculated from accumulated degree-days were, at a given date, around 20% lower at the release site during the 15 years after the translocation than at the site of origin during the 15 years prior to the translocation. Previous work had identified phosphorus as a likely limiting resource for beaver reproduction, and in most post-translocation years phosphorus concentrations at the release site were below the required level towards the end of gestation. In addition to an increase over time, annual reproductive rate of the translocated beavers was highest in cold springs. We conclude that it was initially low due to a combination of low food quality and factors not related to climate.

According to current theories of territoriality, an animal is expected to defend the smallest area that can provide resources for maximisation of reproduction, known as the economically defendable area. In group territorial species however, the strategies behind resource defence are likely to be more complex with corporate territoriality, cooperative breeding, delayed dispersal and intra-group competition all potentially playing a role. Here we examined group territoriality in a social herbivorous rodent, the Eurasian beaver Castor fiber. Beavers in our study do not inhabit economically defendable territories. Instead the sequence of arrival of pairs into unoccupied areas seems to play a more important role in determining the size of the territory, whereas group size is determined by past reproductive success. We argue that the settlement pattern and reproductive history have a lasting impact in the territorial system of beavers due to a combination of the low adult mortality, high dispersal costs, and avoidance of resource depletion. [KEYWORDS: Prospective resource defence ; Resource depletion ; Territory size ; Cooperative breeding ; Delayed dispersal ; Territorial inheritance ; Habitat quality]

1. The consequences of plastic responses of the avian digestive tract for the potential of birds to disperse other organisms remain largely uninvestigated. 2. To explore how a seasonal diet switch in Mallard (Anas platyrhynchos L.) influences their potential to disperse plants and invertebrates, we recorded the retention time of markers, following exposure to two diets of contrasting digestibility (trout chow vs seeds). 3. We then recorded the retrieval and germination of Fennel Pondweed (Potamogeton pectinatus L.) seeds and Brine Shrimp (Artemia franciscana Kellogg) cysts ingested by the same birds. 4. Gut passage rates of markers were increasingly longer in birds on the seed-based, high-fibre diet and shorter in birds on the animal-based, low-fibre one. 5. Propagule digestibility, and thus survival to gut passage, differed between diet groups, with more seeds and fewer cysts retrieved from ducks on the animal-based diet. Germination decreased with retention time, but was not affected by diet. 6. Differences in passage rates of markers but not of seeds and cysts suggest no change in dispersal distances of plants and invertebrates between seasons, while differences in digestibility would affect the numbers of propagules dispersed. [KEYWORDS: Artemia franciscana ; diet switch ; endozoochorous dispersal ; Potamogeton pectinatus]

1. The general notion is that negative effects of vertebrate herbivores on water plants, which play a key role in freshwaters, prevail, and that positive feedbacks of herbivores on plants are insignificant. 2. The most likely systems to find such positive feedbacks are those in which herbivores exert strong feeding pressures on plants during part of the year. Previous theoretical work has suggested that compensatory production occurs when migratory Bewick's swans forage on tubers of fennel pondweed. As a corollary, the swans can exploit the tubers down to a level that maximises their tuber yield. 3. In order to test these hypotheses, I measured pondweed tuber biomass on three occasions per year (just before and after foraging, and just before tuber sprouting) in three consecutive years. The 17 sampling sites in the Lauwersmeer (the Netherlands) were classified according to their silt content and water depth. Within four silt-depth classes, I predicted for each year tuber biomass production and, from that, the optimum foraging threshold that would result in the maximum tuber biomass yield 4. Water depth did not affect tuber production, and silt content only did in one of the 3 years. In accordance with overcompensation predictions, tuber production was higher at plots with moderate foraging pressures than at plots with little or no grazing. However, the winter and summer conditions following the swan foraging had large unpredictable effects on tuber mortality and production. 5. These results indicate that overcompensation by fennel pondweed occurs and that Bewick's swans are generally able to profit from it, albeit without fine-tuning of the foraging threshold to the yield. [KEYWORDS: compensatory production ; foraging threshold ; intraspecific competition ; maximum yield ; sequential model]

Because energy reserves limit flight range, wind assistance may be of crucial importance for migratory birds. We tracked eight Bewicks swans Cygnus columbianus bewickii, using 95-g satellite transmitters with altimeters and activity sensors, during their spring migration from Denmark to northern Russia in 1996. During the 82 occasions where a swans location was recorded in flight, average flight altitude was 165 m a.s.l. with a maximum of 759 m a.s.l., despite winds often being more favourable at higher altitudes. We also counted Bewicks swans departing from the Gulf of Finland and subsequently passing an observatory in the next major stop-over area 800 km further north in the White Sea, northern Russia, during the springs of 1994, 1995 and 1996. A comparison of these counts with wind data provided evidence for Bewicks swans using favourable changes in wind conditions to embark on migration. Changes in the numbers of birds arriving in the White Sea correlated best with favourable changes in winds in the Gulf of Finland 1 day earlier. Again, migratory volume showed a correlation with winds at low altitudes only, despite wind conditions for the swans being more favourable at high altitudes. We conclude that the relatively large Bewicks swan tends to gear its migration to wind conditions at low altitude only. We argue that Bewicks swans do not climb to high altitudes because of mechanical and physiological limitations with respect to the generation of power for flight and to avoid rapid dehydration. [KEYWORDS: Bewicks swan ; Cygnus columbianus bewickii ; Migration ; Flight altitude ;Tailwind vector ; Satellite transmitters]

Many birds feed on submerged macrophytes during a temporally discrete period every year, for instance during migratory stopover or at the wintering grounds. Hence, seasonal herbivory is a common feature of the life cycle in many aquatic macrophytes. We are interested in the effect of Bewick's swans (Cygnus columbianus bewickii) feeding on the tubers of fennel pondweed (Potamogeton pectinatus) in the Netherlands every autumn. For that purpose, we developed a sequential macrophyte population model, including seasons of tuber production, herbivory and winter mortality as distinct and unambiguously defined events. The model is characterised and parameterised with both field and laboratory data. Tuber consumption inevitably decreases the density of ramets sprouting next spring, but it may actually increase the density of tubers produced in the following autumn. Hence, we can only understand the effect of grazing on the fennel pondweed population by recognising the seasonal structure of density-dependence. The mean density of fennel pondweed and the yield taken by swans are dependent on the foraging threshold below which no grazing takes place. Furthermore, the consumption has a stabilising effect for a wide range of parameter values. [KEYWORDS: aquatic macrophytes, Cygnus columbianus, functional response, Potamogeton pectinatus, sequential density-dependence]

We review whether migratory Anatidae, i.e., swans, geese and ducks, could be acting as vectors for dispersal of Zostera, Ruppia and Potamogeton propagules by endozoochory (carrying seeds in their guts). We list six prerequisites that must all be fulfilled, if successful dispersal should occur. Several Anatidae species feed on these macrophytes, and undertake rapid long-distance movements, making dispersal possible. We identify four problems, which in combination leads us to conclude that long-distance dispersal events are likely to be rare. (i) Most long-distance movements are out of phase with the reproductive efforts of the plants, and if birds arrive at sites when plants still bear seeds, they are likely to depart well after seed stocks have been depleted. (ii) Seed transport by birds will usually be uni-directional, from north to south on autumn migrations. (iii) Most of the gut contents of migratory birds are likely to have been discarded within 300 km of departure. (iv) In many cases, birds will arrive in habitats seriously different from those they departed, i.e., any seeds carried along will have low chances of surviving in their new site. We suggest that northbound dispersal by endozoochory can only occur during spring if waterbirds feed on seeds that have not been depleted and remained frozen down or buried in sediments, or during moult- or post-moult migrations. Moult migration takes place in summer in phase with the reproductive efforts of the plants. Also epizoochorous dispersal (external attachment) is subject to restrictions i, ii and iv. [KEYWORDS: Anas; Branta; Cygnus; Potamogeton; Ruppia; Zostera marina]

1.In a system where depletion drives a habitat shift, the hypothesis was tested that animals switch habitat as soon as the average daily net energy intake (or gain) drops below that attainable in the alternative habitat. 2. The study was performed in the Lauwersmeer area. Upon arrival during the autumn migration, Bewick's swans first feed on below-ground tubers of fennel pondweed on the lake, but subsequently switched to feeding on harvest remains in sugar beet fields. 3. The daily energy intake was estimated by multiplying the average time spent foraging per day with the instantaneous energy intake rate while foraging. In the case of pondweed feeding, the latter was estimated from the functional response and the depletion of tuber biomass. In the case of beet feeding, it was estimated from dropping production rate. Gross energy intake was converted to metabolizable energy intake using the assimilation as determined in digestion trials. The daily energy expend 4. The daily gain of pondweed feeding at the median date of the habitat switch (i.e. when 50% of the swans had switched) was compared with that of beet feeding. The daily gain of beet feeding was calculated for two strategies depending on the night activity on the lake: additional pondweed feeding (mixed feeding) or sleeping (pure beet feeding). 5. The majority of the swans switched when the daily gain they could achieve by staying on the pondweed bed fell just below the average daily gain of pure beet feeders. However, mixed feeders would attain an average daily gain considerably above that of pondweed feeders. A sensitivity analysis showed that this result was robust. 6. We therefore reject the hypothesis that the habitat switch by swans can be explained by simple long-term energy rate maximization. State-dependency, predation risk, and protein requirements are put forward as explanations for the delay in habitat switch.

Previous research has found that efficiency, or, more precisely, the foraging gain ratio (FGR), is a valid currency in foraging theory when (1) there is a limit to the energy that can be assimilated by the forager and (2) a forager is trying to meet an energy requirement. The FGR is b/ (c — cr), where b is the rate of metabolizable energy intake, and c and cr are the rates of energy expenditure while foraging and resting, respectively. Here I show that, when energy expenditure has a cost besides energy, animals should also choose the option with the highest FGR when they are aiming at a given positive daily gain. The next question is which gain they should aim for? Researchers have shown that observed intake levels of growing ruminants are close to the levels predicted by maximization of the efficiency of oxygen utilization. This currency can be approximated by (B — C + Cr) / C, where B is the daily metabolizable energy intake, and C and Cr are the total and basal daily energy expenditures, respectively. By simulating growth at different intake levels, I found that mass-specific oxygen consumption rate is indeed minimal at the observed intake levels. This is the first study in which these efficiency measures (FGR and the efficiency of oxygen utilization) are combined. [KEYWORDS: energy balance, feed intake regulation, foraging gain ratio, growth, optimal foraging theory, oxygen utilization]

Some migratory birds refuel at stopover sites that they by-pass on the return trip. In theory, this skipping behaviour is only expected in time-selected migrants when the overflown site is of a lower quality than the departure site. We provide empirical evidence that quality differences in stopover sites are the cause for skipping in Bewick's Swans Cygnus bewickii tracked by satellite telemetry. Two and five complete tracks were recorded in spring and autumn, respectively, showing that the White Sea was visited for c. 2 weeks in spring, but by-passed (or visited for a few days at the most) in autumn. Skipping of the White Sea in autumn was predicted by a dynamic programming model which was based on calculated gain rates during stopover in the Pechora Delta and the White Sea. This prediction was not sensitive to plausible variations in gain rates. Relative to the Pechora Delta the White Sea is a poor site because a large tidal amplitude precludes foraging on the beds of the submerged macrophyte Fennel Pondweed Potamogeton pectinatus during high tide. The dynamic programming model predicted a fast autumn migration. However, the phenology of autumn arrival dates of Bewick's Swans on the wintering grounds revealed that only in three out of ten years a significant number of birds was able to reach the wintering grounds without refuelling. In the other years, unfavourable wind conditions along the Russian/Baltic part of the route prevented such non-stop migration [KEYWORDS: Cygnus bewickii; fuelling rates; dynamic programming; optimal migration; satellite tracking; wind assistance]

1. Tundra swans forage on below-ground pondweed tubers that are heterogeneously distributed in space. The swans have no visual cues to delineate patches. It was tested whether swans employ an area-restricted search tactic. Theory predicts that swans should alternate between an intensive (low-speed, sinuous) search mode in high tuber density areas and an extensive (high-speed, directed) search mode between these areas. 2. A quantitative analysis of movement paths recorded over short time frames (15 min) revealed that the sequential step lengths were strongly autocorrelated. After partitioning the data in low-speed paths and high-speed paths, this autocorrelation was very much reduced. 3. Movement paths with low speed were non-directional and could well be described as random walks. In contrast, high-speed paths were directed forward, and were better described as correlated (i.e. directional) random walks. 4. Movement paths recorded over longer time frames (1-4 h) provided empirical evidence that an alternation of low-speed, sinuous and high-speed, directed searches occurred. 5. There was a spatial autocorrelation in tuber biomass density, being significantly positively correlated until c. 10 m distance. The scale of the food clump size and step length of high-speed paths matched, suggesting that they were causally linked. 6. Computer simulations confirmed that swans using the observed search tactic achieved a higher energy gain than swans using only an intensive search mode, provided that the tuber biomass density occurred in clumps. They also achieved a higher gain than swans that alternated between intensive and extensive search mode, but always moved in a random direction.

We tested whether the spatial variation in resource depletion by Tundra Swans (Cygnus columbianus) foraging on belowground tubers of sage pondweed (Potnmogeton pectinatus) was caused by differences in net energy intake rates. The variation in giving up densities within the confines of one lake was nearly eightfold, the giving-up density being positively related to water depth and, to a lesser extent, the silt content of the sediment. The swans' preference (measured as cumulative foraging pressure) was negatively related to these variables. We adjusted a model developed for diving birds to predict changes in the time allocation of foraging swans with changes in power requirements and harvest rate. First, we compared the behavior of free-living swans foraging in shallow and deep water where they feed by head-dipping and up-ending, respectively. Up-ending swans had 1.3-2.1 times longer feeding times than,head-dipping swans. This was contrary to our expectation, since the model predicted a decrease in feeding time with an increase in feeding power. However up-ending swans also had 1.9 times longer trampling times than head-dipping swans. The model predicted a strong positive correlation between trampling time and feeding time, and the longer trampling times may thus have masked any effect of an increase in feeding power. Heart rate measurements showed that trampling was the most energetically costly part of foraging. However, because the feeding time and trampling time changed concurrently, the rate of energy expenditure was only slightly higher in deep water (1.03-1.06 times). This is a conservative estimate since it does not take into account that the feeding costs of up-ending are possibly higher than that of head-dipping. Second, we compared captive swans foraging on sandy and clayey sediments. We found that the harvest rate on clayey sediment was only 0.6 times that on sandy sediment and that the power requirements for foraging were 1.2-1.4 times greater. Our results are in qualitative agreement with the hypothesis that the large spatial variation in giving-up densities was caused by differences in net rates of energy intake. This potentially has important implications for the prey dynamics, because plant regrowth has been shown to be related to the same habitat factors (water depth and sediment type.) [KEYWORDS: Bewick's Swan, Cygnus columbianus bewickii, energy expenditure, fennel pondweed, giving-up density, Lauwersmeer, The Netherlands, optimal foraging, patch use, plant-herbivore interaction, Potamogeton pectinatus, sago pondweed, Tundra Swan]

We searched for a major stopover site of Bewick's Swans Cygnus columbianus bewickii in the White Sea following the suggestion that one should exist on the stretch between Estonia and the breeding grounds (1750 km). We discovered 733 Swans in Dvina Bay during a late aerial survey in spring 1993. Subsequently, ground-based research was carried out in May 1994, 1995 and 1996 in the Dry Sea, a tidal, shallow bay with fresh to brackish water just north of the Dvina Delta. The total number of passing Bewick's Swans was estimated at 10 974 (1994), 9593 (1995) and 17 972 (1996) (32-60% of the flyway population). Estimated peak numbers staging were 1500-2000 (9 May 1994), 4937 (17 May 1995) and 4457 (24 May 1996) (> 5-16% of the flyway population). The Swans foraged almost exclusively on submerged water plants apart from some supplemental feeding on emerged food plants around high tide. Stoneworts Chara spp. were an important food in the late spring of 1996, because they grew in places where bog streams quickly melted the ice. At this latitude (65 degreesN) food alternatives to the submerged macrophytes are rare in spring, but we cannot rule out the possibility that the Swans forage on grass rhizomes on inundated pastures. One bird tracked by satellite staged 15 days in Dvina Bay, of which four days were spent in the Dry Sea, in accordance with other indications that the Dry Sea is part of a larger stopover site within Dvina Bay. Recent evidence shows that the Swans largely skip the White Sea during autumn migration. However, in spring the birds probably need this stopover to be able to carry reserves to the breeding grounds. At present, the preservation of the submerged vegetation in Dvina Bay seems to be crucial to the conservation of this Bewick's Swan population.

Due to over-hunting c. 1200 Eurasian beavers Castor fiber survived in eight relict populations in Europe and Asia at the beginning of the 20th century. Following hunting restrictions and translocation programmes in IS countries, the Eurasian beaver became re-established over much of its former range, and presently numbers c. 430 000. The translocated populations often consist of a mixture of geographical forms. Preservation of the original, unmixed populations has therefore top priority. all five in Europe have reached the assumed minimum viable population size of c. 1880 animals each, but the three in Asia are still endangered. Their protection should be carried out at the level of river catchments. Nowadays the main threats for beavers are habitat destruction and introduced North American beavers Castor canadensis. On the other hand, gr owing beaver populations cause increasing conflicts with man, and population and/or damage control may therefore be required. In view of these two very different problems, we conclude that the conservation of beavers is best served by preservation and restoration of riparian woods with intact natural water regimes. [KEYWORDS: conservation biology; reintroduction; translocation; population control and damage control European beaver; population; canadensis; forest]

Current models of bird migration consider the optimal decision making of individuals irrespective of the behaviour of conspecifics. We observed the foraging behaviour of Bewick's Swans Cygnus columbianus during their stop-over in the Dvina Bay in the White Sea in spring 1995 and 1996. Here the swans could only feed on their main food, tubers of fennel pondweed Potamogeton pectinatus. during low tide. In the course of the staging period, the swans tended to forage at progressively lower water levels, indicating that they depleted their main food and exploited increasingly deeper parts of the tuber-bank. This reduced the swans' main foraging period from 6.0 h/tide on 20 May to 3.3 h/tide on 28 May 1996. We calculated that this must have greatly decreased the rate of refuelling during the staging period. In accordance with this, swans arriving early stayed shorter than those arriving late. It seems of paramount importance to the swans to arrive as early as possible at the stop-over site. We conclude that when food depletion occurs, a game theoretical approach is needed to improve our understanding of the birds' adaptive behaviour during their migration. [KEYWORDS: BIRDS, PERFORMANCE, FLIGHT]

We tested the hypothesis that a main function of territory marking in Eurasian beaver (Castor fiber) is defense of the territory. The results showed that: (1) beaver colonies with close neighbors scent-mark more often than isolated ones; (2) the number of scent markings increased significantly with the number of neighboring territories and individuals, the mean distance to all other territories, duration of territory occupancy and length of wooded banks within the territory; (3) the number of scent markings by a colony was not related to the number of animals in the colony, the distance to nearest neighbors' main resting-site, or age of beaver; (4) the number of scent mounds or scent markings in territories with breeding and nonbreeding beaver did not differ; (5) there was no significant difference in the number of scent markings during the season (16 April-31 August); and (6) some of the scent mounds were concentrated at feeding sites, resting sites, and near trails, but most had no apparent relationship to any of these sites. Therefore we concluded that scent marking apparently plays an important role in territory defense of the Eurasian braver. [KEYWORDS: beaver; Castor fiber; scent communication; scent mound; scent marking; radiotelemetry; territorial behavior; Netherlands]

Between 1988 and 1994, 58 beavers were translocated from the Elbe region (Germany) to the Netherlands. In 43 animals, radio-transmitters were implanted with a pulse interval which was dependent on body temperature; subsequently, 22 of the released animals were found dead and the cause of death was investigated. Infectious diseases were the most important cause of death (50%), in particular yersiniosis and leptospirosis. Yersiniosis is relatively rare in the source population along the Elbe, and leptospirosis has not been reported before in beavers. We hypothesize that the immune system of the translocated animals was weakened by stress at a time of exposure to new pathogens. In future translocations of beavers, we recommend to reduce the stress, improve the hygiene, and to vaccinate against Yersinia pseudotuberculosis and leptospirosis prior to the release.

We monitored survival, reproduction and emigration of a translocated beaver Castor fiber population in the Netherlands for five years and used a stochastic model to assess its viability. Between 1988 and 1991, 42 beavers were released in the Biesbosch National Park. The mortality was initially high but gradually fell to normal rates. However, the breeding success was low, and we hypothesized that this was either a temporary phenomenon (the translocation hypothesis) or a permanent feature (the poor habitat hypothesis). According to the computer simulations, the isolated population was viable under the first but not under the second hypothesis. In the latter case, the prospects generally improved by the foundation of another population in the Gelderse Poort (100 km from the Biesbosch), However, this second habitat should be optimal for beavers in order to reduce the extinction probability of the Biesbosch population to below 10% in 100 years,' the loss of genetic variability (1-2% per generation) was just above the applied tolerable risk (1%), but the effects of inbreeding are unknown in beavers. We conclude that the beaver population in the Biesbosch is not viable unless the reproductive success increases, either in the Biesbosch itself or in a nearby population. We recommend applying such viability analyses to evaluate the likely success of any translocation. [KEYWORDS: population viability analysis; reintroduction;translocation; simulation model; Castor fiber Species conservation; extinction]

Linear programming has been remarkably successful in predicting the diet choice of generalist herbivores. We used this technique to test the diet choice of free-living beavers (Castor fiber) in the Biesbosch (The Netherlands) under different Foraging goals, i.e. maximization of intake of energy, nitrogen, phosphorus or sodium, or minimization of feeding time. Three food types were distinguished, Le, woody food, herbs and roots of monocots. We assessed forage quality by measuring the dry matter, energy and mineral contents of the food plants as well as food intake rates, digestibility and metabolisability in captive beavers. Actual diet was in accordance with the predicted food choice in the summer when the beavers were minimizing feeding time by mainly eating woody food. However, in the winter and spring, the beavers were predicted to feed upon non-woody food, whereas they (again) nearly exclusively are woody food. The major reasons for this discrepancy might be: (1) the foraging constraints were inappropriate, (2) the foraging goals were inadequately defined, or (3) the beavers were not foraging optimally. We sugg [KEYWORDS: optimal foraging, nutrition, re-introduction, Castor fiber, The Netherlands]