Swans Cygnus Columbianus Bewickii Research Articles

Using satellite tracking to assess the use of protected areas and alternative roosts by Whooper and Bewick's Swans

Protected areas are one of the major tools used in the conservation of biodiversity, but animals are unlikely always to remain within these human‐made boundaries. Understanding when and why species choose to leave protected areas can help us to improve the effectiveness of these management tools. Here, we investigate the use of protected and non‐protected areas by two migratory species undergoing rapid wintering population changes in northwest Europe: Whooper Swans Cygnus cygnus and Bewick's Swans Cygnus columbianus bewickii. Global positioning system tags were fitted to 15 Whooper Swans in winter 2008/09 and to 18 Bewick's Swans from winter 2013/14 to 2014/15 at the Ouse Washes Special Protection Area (an internationally important roost for wintering waterbirds) and on adjacent fields in southeast England. Here, swans feed on farmland during the day but return to designated reserves to roost at night, where they receive protection from predators and disturbance within managed roost habitats. When swans roost elsewhere at alternative sites, they may face more adverse conditions, and so understanding the extent and causes of the use of alternative roosts is important for swan conservation efforts. The alternative roosting proportion, defined as the proportion of nights spent outside protected reserves, was 0.237 for Bewick's Swans and challenging to quantify accurately for Whooper Swans. A generalized additive mixed model to model repeated measurements on individuals showed that the proportion of time that Bewick's Swans spent at alternative roosts correlated positively with river level and negatively with temperature. Competition and foraging flight distances are thought to drive these relationships, as swans seek access both to roost space and to nearby feeding habitats. Our findings improve our understanding of the environmental conditions under which migratory waterbirds may choose to roost outside protected areas.

Conservation of Bewick’s swans (Cygnus columbianus bewickii): Insights from the identification of critical stopover sites and migration corridors

Migratory birds face diverse threats during migration. Critical stopover sites (CSSs) are essential refueling and resting sites for migratory birds that ensure their complete migration and survival. Therefore, identifying bird migration patterns, routes, and critical habitats is vital for conservation. From 2018–2022, we deployed satellite tracking devices on 30 Bewick’s swans (Cygnus columbianus bewickii) wintering in China to determine their migration routes. Using a dynamic Brownian bridge movement model, we identified migration corridors, core movement areas, and CSSs for Bewick’s swans. Combining protected area databases and human settlement types, we further assessed the swans’ conservation status and human impacts on CSSs. The results showed that Bewick’s swans migrated north from their wintering grounds using one of three routes (west, middle, and east), passing through Mongolia to reach the Russian Arctic (breeding grounds) in spring. They followed similar routes during autumn to return to wintering grounds. We found a new middle route within the East Asian–Australasian Flyway (EAAF) and several northward expanded wintering sites. Our study revealed similarities and differences in the spring and autumn migrations, with longer stopover durations in spring due to migration strategies and ice conditions. Moreover, our findings identified the Inner Mongolia region, the Songnen Plain, the Bohai Rim of China, and the main streams of the Lena River and the Ob River of Russia as CSSs for Bewick’s swans. However, the conservation status of the CSSs was relatively low; and the situation was more severe in spring than in autumn, only 4.3% of the total area was protected, likely due to the distribution of farmland and urban areas. Specifically, 14.3% of the CSSs in China were in urban areas during spring, while in Mongolia and Russia this figure was less than 1%. Therefore, it is necessary to balance waterbird conservation with sustainable agriculture and urban development. This research contributes to our understanding of the migratory ecology of Bewick’s swans wintering in China. The identified migration corridors and CSSs are crucial for the future conservation of swans along the EAAF.

Sensitivity analysis of collision risk at wind turbines based on flight altitude of migratory waterbirds

Abstract A rapid increase in wind power generation has led to bird collisions becoming a serious problem worldwide. Developing useful sensitivity maps to select low‐risk sites for birds is an urgent issue. For migratory birds, such as geese and swans, that visit different habitats throughout their life cycle, it is important to conduct risk assessments that take into account their behavioural characteristics in each habitat. Geese and swans fly and migrate at varying altitudes (above the ground) ranging from 10 to hundreds of metres. Accurate predictions of avian flight altitudes are essential in assessing the risks of collisions with human‐made structures. We first obtained location data for four species of geese and swans to identify their spring migratory routes within Japan (Bean Goose Anser fabalis and Anser serrirostris, Greater White‐fronted Goose Anser albifrons, Tundra Swan Cygnus columbianus bewickii and Whooper Swan Cygnus cygnus). As all four species used the same roosts and overlapping foraging areas from winter to spring, a single migratory route was defined by integrating the location data of the four species. Flight trajectories were tracked using an ornithodolite. The median flight height for these four species in all landscape types was 150 m or less. Then a LASSO regression model was created with flight altitude obtained as the response variable and topographic and landscape factors as explanatory variables. Trends in flight altitude with environmental differences were similar for the four species, indicating that topographical factors strongly influence flight altitude. Finally, a statistical model was used to predict flight altitudes along migration routes. The sensitivity maps we generated showed that for all four species, most flight heights during spring were within the wind turbine range, suggesting that their risk of collision with wind turbines was greater along their migratory route. Sensitivity maps that accurately reflect avian flight characteristics help provide useful information when considering the location of further wind turbine construction.

Predicting intention to hunt protected wildlife: a case study of Bewick's swan in the European Russian Arctic

AbstractIllegal killing of wildlife is a major conservation issue that, to be addressed effectively, requires insight into the drivers of human behaviour. Here we adapt an established socio-psychological model, the theory of planned behaviour, to explore reasons for hunting the Endangered Bewick's swan Cygnus columbianus bewickii in the European Russian Arctic, using responses from hunters to a questionnaire survey. Wider ecological, legal, recreational and economic motivations were also explored. Of 236 hunters who participated overall, 14% harboured intentions to hunt Bewick's swan. Behavioural intention was predicted by all components of the theory of planned behaviour, specifically: hunters' attitude towards the behaviour, perceived behavioural control (i.e. perceived capability of being able to perform the behaviour) and their subjective norms (perception of social expectations). The inclusion of attitude towards protective laws and descriptive norm (perception of whether other people perform the behaviour) increased the model's predictive power. Understanding attitudes towards protective laws can help guide the design of conservation measures that reduce non-compliance. We conclude that conservation interventions should target the socio-psychological conditions that influence hunters' attitudes, social norms and perceived behavioural control. These may include activities that build trust, encourage support for conservation, generate social pressure against poaching, use motivations to prompt change and strengthen peoples' confidence to act. This approach could be applied to inform the effective design, prioritization and targeting of interventions that improve compliance and reduce the illegal killing of wildlife.

Behavioural and energetic consequences of competition among three overwintering swan (Cygnus spp.) species

BackgroundWinter numbers of the northwest European population of Bewick’s Swans (Cygnus columbianus bewickii) declined recently by c. 40%. During the same period, numbers of two sympatric and ecologically-similar congeners, the Mute Swan (Cygnus olor) and Whooper Swan (Cygnus cygnus) showed increases or stability. It has been suggested that these opposing population trends could have a causal relationship, as Mute and Whooper Swans are larger and competitively dominant to Bewick’s Swans in foraging situations. If so, effects of competition of Mute and Whooper Swans on Bewick’s Swans should be detectable as measurable impacts on behaviour and energetics.MethodsHere, we studied the diurnal behaviour and energetics of 1083 focal adults and first-winter juveniles (“cygnets”) of the three swan species on their winter grounds in eastern England. We analysed video recordings to derive time-activity budgets and these, together with estimates of energy gain and expenditure, were analysed to determine whether individual Bewick’s Swans altered the time spent on key behaviours when sharing feeding habitat with other swan species, and any consequences for their energy expenditure and net energy gain.ResultsAll three swan species spent a small proportion of their total time (0.011) on aggressive interactions, and these were predominantly intraspecific (≥ 0.714). Mixed-effects models indicated that sharing feeding habitat with higher densities of Mute and Whooper Swans increased the likelihood of engaging in aggression for cygnet Bewick’s Swans, but not for adults. Higher levels of interspecific competition decreased the time spent by Bewick’s Swan cygnets on foraging, whilst adults showed the opposite pattern. When among low densities of conspecifics (< c. 200 individuals/km2), individual Bewick’s Swans spent more time on vigilance in the presence of higher densities of Mute and Whooper Swans, whilst individuals within higher density Bewick’s Swan flocks showed the opposite pattern. Crucially, we found no evidence that greater numbers of interspecific competitors affected the net energy gain of either adult or cygnet Bewick’s Swans.ConclusionsWe found no evidence that Bewick’s Swan net energy gain was affected by sharing agricultural feeding habitat with larger congeners during winter. This was despite some impacts on the aggression, foraging and vigilance behaviours of Bewick’s Swans, especially among cygnets. It is unlikely therefore that competition between Bewick’s Swans and either Mute or Whooper Swans at arable sites in winter has contributed to the observed decline in Bewick’s Swan numbers. Further research is needed, however, to test for competition in other parts of the flyway, including migratory stopover sites and breeding areas.

Predicting avian herbivore responses to changing food availability and competition

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.

Calibrating tri-axial accelerometers for remote behavioural observations in Bewick's swans

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.

Apparent breeding success drives long‐term population dynamics of a migratory swan

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 swanCygnus 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.

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

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

Colin James Pennycuick (1933–2019)

Colin James Pennycuick (1933–2019)

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

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

The Influence of Food Density, Flock Size, and Disturbance on the Functional Response of Bewick's Swans (Cygnus columbianus bewickii) in Wintering Habitats.

Simple SummaryChanges in environmental conditions cause animals to adjust their behavioral strategies to survive. We investigated foraging behavior in different habitats of wintering Bewick’s swans. We found that the observed feeding rate was not affected by food density but showed a negative relationship with flock size and disturbance time. Handling time had a negative relationship with food density and flock size, but a positive relationship with disturbance. Searching rate was negatively correlated with food density, flock size, and disturbance time. This provides insight into how wintering waterbirds adapt their foraging behavior in complex environments.Perceiving how animals adjust their feeding rate under a variety of environmental conditions and understanding the tradeoffs in their foraging strategies are necessary for conservation. The Holling functional response, which describes the relationship of feeding rate and food density to searching rate and handling time, has been applied to a range of waterbirds, especially with regard to Type II functional responses that describe an increasing feeding rate with food density but at a decelerating rate as the curve approaches the asymptote. However, feeding behavior components (feeding rate, searching rate, and handling time) are influenced by factors besides prey density, such as vigilance and flock size. In this study, we aim to elucidate how Bewick’s swans (Cygnus columbianus bewickii) adopt flexible foraging strategies and vary their feeding behavior components in response to disturbance, flock size, and food density. We collected focal sampling data on the foraging behavior of swans that foraged rice grains, foxnut seeds, and tubers in paddy field, foxnut pond, and lake habitats, respectively, in Shengjin and Huangpi lakes during winter from 2016 to 2018. The observed feeding rate was not correlated with food density and displayed a positive relationship with searching rate but negative relationships with handling time, flock size, overall vigilance time, and disturbance time. Handling time was negatively correlated with food density and flock size, yet it increased with disturbance, overall vigilance time, and normal vigilance time. Searching rate was negatively correlated with food density, flock size, and disturbance time. Feeding rate was affected by the combined effects of handling time and searching rate, as well as food density and searching rate. The shape of the observed functional response could not be fitted to Holling’s disc equation. However, the disc equation of the predicted feeding rate of wintering swans was found to be driven by food density. This provides insight into how wintering waterbirds adopt appropriate foraging strategies in response to complicated environmental factors, which has implications for wildlife conservation and habitat management.

Seasonal variation in energy gain explains patterns of resource use by avian herbivores in an agricultural landscape: Insights from a mechanistic model

Our understanding of how energy shapes animal behavioural decisions has been limited by the difficulty of measuring directly the energy gain and expenditure in free-living animals. Mechanistic models that simulate energy gain and expenditure from estimable parameters can overcome these limitations and hence could help scientists to gain a predictive understanding of animal behaviour. Such models could be used to test mechanistic explanations of observed patterns of resource use within a landscape, such as behavioural decisions to switch among food resources. Here, we developed mechanistic models of the instantaneous and daily rates of net energy gain for two species of migratory swans, the Bewick’s swan (Cygnus columbianus bewickii) and whooper swan (Cygnus cygnus), that feed on root and cereal crops within an agricultural landscape in eastern England. Field data show that both species shift from using predominantly root crops (e.g. sugar beet and potatoes) in early winter to using mostly cereals (e.g. wheat) in late winter. Our models correspondingly predicted that swans could achieve the greatest rates of net energy gain on root crops in early winter and on cereal crops in late winter. The change from root crops to cereal crops providing the greatest net rates of energy gain was predicted to occur at the same time as the birds’ switch from feeding predominantly on root crops to predominantly cereal crops (between December and January). We used Monte Carlo simulations to account for variance in model parameters on predictions of energy gain and profitability. A sensitivity analysis indicated that predictions of net energy gain were most sensitive to variance in the intake rate and food quantity parameters. The agreement between our model estimates of energy gain and the observed shifts in resource use observed among the overwintering swans suggests that maximising net rates of energy gain is an important resource selection strategy among overwintering birds. A mechanistic understanding of where and when birds will use food resources can inform the conservation management of key feeding areas for species of conservation concern, as well as the deployment of crop protection strategies.

Conservation implications of misidentification and killing of protected species

Killing protected species mistaken for morphologically similar quarry species, or species with weaker protection, can hinder their conservation. Despite policy aims to reduce threats from illegal killing, information is lacking on susceptible species, conservation impacts and the identification accuracy of hunters. We examined the ability of hunters (n = 232) in Arctic Russia to identify the endangered Northwest European Bewick's swan Cygnus columbianus bewickii using photographs. Only 14% (n = 33) identified this species correctly and distinguished it from sympatric and congeneric whooper swans C. cygnus and mute swans C. olor, with 15% of individuals admitting to accidentally hunting a Bewick's swan in the previous 3 years. We conclude that there is a risk of Bewick's swans being shot accidentally when mistaken for similar species with less legal protection. Improving hunters' skills in discerning protected from legitimate quarry species is likely to be an effective tool for conservation of morphologically similar species.

Are long-term trends in Bewick’s Swan Cygnus columbianus bewickii numbers driven by changes in winter food resources?

SummaryThe north-west European population of Bewick’s SwanCygnus columbianus bewickiideclined by 38% between 1995 and 2010 and is listed as ‘Endangered’ on the European Red List of birds. Here, we combined information on food resources within the landscape with long-term data on swan numbers, habitat use, behaviour and two complementary measures of body condition, to examine whether changes in food type and availability have influenced the Bewick’s Swan’s use of their main wintering site in the UK, the Ouse Washes and surrounding fens. Maximum number of Bewick’s Swans rose from 620 in winter 1958/59 to a high of 7,491 in winter 2004/05, before falling to 1,073 birds in winter 2013/14. Between winters 1958/59 and 2014/15 the Ouse Washes supported between 0.5 and 37.9 % of the total population wintering in north-west Europe (mean ± 95 % CI = 18.1 ± 2.4 %). Swans fed on agricultural crops, shifting from post-harvest remains of root crops (e.g. sugar beet and potatoes) in November and December to winter-sown cereals (e.g. wheat) in January and February. Inter-annual variation in the area cultivated for these crops did not result in changes in the peak numbers of swans occurring on the Ouse Washes. Behavioural and body condition data indicated that food supplies on the Ouse Washes and surrounding fens remain adequate to allow the birds to gain and maintain good body condition throughout winter with no increase in foraging effort. Our findings suggest that the recent decline in numbers of Bewick’s Swans at this internationally important site was not linked to inadequate food resources.

Has winter body condition varied with population size in a long-distance migrant, the Bewick’s Swan (Cygnus columbianus bewickii)?

Assessments of body condition can provide useful information on changes in the state of individuals within a population, which may in turn help to inform conservation efforts. For example, decreases in body condition over time can indicate reduced food resources. Mass and skull length measures recorded for 195 adult and 467 first winter (cygnets) Bewick’s Swans (Cygnus columbianus bewickii) at wintering sites in the UK between winters 1966/1967 and 2017/2018 therefore were analysed to determine whether a ca. 40% decline in numbers in the Northwest European Bewick’s Swan population between 1995 and 2010 corresponded with poorer body condition from the mid-1990s onwards. Parents and siblings were known for all individuals, allowing us to account for shared genetic factors and rearing environment in our analysis. We used linear mixed-effects models and an information-theoretic approach to test different models of temporal variation in scaled body mass index (SBMI). Within our study population, although SBMI values varied both within and between years, we found no evidence of any directional trends in body condition. Of our competing time models of swan SBMI, a model in which age-specific body condition was constant over time received the greatest support in the data. Body condition was greater for adults than cygnets, but did not vary between sexes or wintering sites. Our findings suggest no connection between the recent declines in population size and body condition. Population decline is therefore unlikely to be caused by inadequate food supplies.

Apparent survival of an Arctic‐breeding migratory bird over 44 years of fluctuating population size

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 SwanCygnus columbianus bewickii, which declined fromc. 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 inRMarkto 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%CI0.738–0.805; neck‐collar = 0.725, 95%CI0.681–0.764). 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.

Bill colour pattern in Bewick’s swan: information on sex and body size displayed on face?

Ornamental traits in birds (plumage/bare part colours) have been shown to provide information on diverse aspects of individual quality and behaviour. In bird species with achromatic plumages (uniformly white, grey or black), signals encoded in bare parts are the major mode of visual communication. In this study we analysed the influence of sex, age, body condition and selected biometric traits on the bill colour pattern in Bewick’s swans Cygnus columbianus bewickii. The results indicate that sex and body size (weight and tarsus length) are associated with the bill colour pattern in Bewick’s swans. Individuals with a larger black area on the bill were heavier and had a longer tarsus. Moreover, females had more black on their bills than males did. Age, head length and body condition index were not significantly related to bill colour pattern. The present work indicates that Bewick’s swans may display information on their sex and size by means of their bill colour pattern. These findings also suggest the possibility that swans can remotely quantify one another’s traits from the appearance of their bill colouration. Proper assessment of an opponent or of mate quality during face-to-face signalling may help birds to avoid unnecessary interaction or to maintain bonds between mates and family groups.

Inter‐annual variability and long‐term trends in breeding success in a declining population of migratory swans

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 swanCygnus columbianus bewickii, which is currently undergoing a population decline: 1) the percentage of young within the wintering population and 2) mean brood size. We used an information‐theoretic approach to test how these two measures of productivity were influenced over a 26 yr 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.

Lower foraging efficiency of offspring constrains use of optimal habitat in birds with extended parental care

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.