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Body condition is an important concept in behaviour, evolution and conservation, commonly used as a proxy of an individual's performance, for example in the assessment of environmental impacts. Although body condition potentially encompasses a wide range of health state dimensions (nutritional, immune or hormonal status), in practice most studies operationalize body condition using a single (univariate) measure, such as fat storage. One reason for excluding additional axes of variation may be that multivariate descriptors of body condition impose statistical and analytical challenges. Structural equation modelling (SEM) is used in many fields to study questions relating multidimensional concepts, and we here explain how SEM is a useful analytical tool to describe the multivariate nature of body condition. In this 'Research Methods Guide' paper, we show how SEM can be used to resolve different challenges in analysing the multivariate nature of body condition, such as (a) variable reduction and conceptualization, (b) specifying the relationship of condition to performance metrics, (c) comparing competing causal hypothesis and (d) including many pathways in a single model to avoid stepwise modelling approaches. We illustrated the use of SEM on a real-world case study and provided R-code of worked examples as a learning tool. We compared the predictive power of SEM with conventional statistical approaches that integrate multiple variables into one condition variable: multiple regression and principal component analyses. We show that model performance on our dataset is higher when using SEM and led to more accurate and precise estimates compared to conventional approaches. We encourage researchers to consider SEM as a flexible framework to describe the multivariate nature of body condition and thus understand how it affects biological processes, thereby improving the value of body condition proxies for predicting organismal performance. Finally, we highlight that it can be useful for other multidimensional ecological concepts as well, such as immunocompetence, oxidative stress and environmental conditions.

Individual variation in disturbance vulnerability (i.e. the likelihood that disturbance negatively affects an individual's fitness) can affect how disturbance impacts animal populations, as even at low disturbance levels some individuals could be severely affected and die. Individual variation in vulnerability can arise due to different responses to disturbance. We propose a new hypothesis that even when individuals respond similarly to disturbance, time-limited individuals are more at risk that their condition deteriorates since they have limited ability to extend their foraging time to compensate for disturbance. We investigate this ‘time-limitation hypothesis' both empirically and mathematically by studying how individuals that differ in their average foraging time (presumably due to differences in foraging efficiency) are affected by disturbance. We used tracking data of 22 wintering Eurasian oystercatchers Haematopus ostralegus to compare time budgets between disturbed and undisturbed tidal periods. In three tidal periods with severe disturbance by transport airplanes, more than a third of the variation in additional flight time and foraging time loss was explained by individual differences. Inefficient individuals that foraged longer in undisturbed tidal periods experienced higher costs in disturbed tidal periods, since they lost more foraging time. We next used an analytical energy balance model to study how time-limited individuals differed in their maximum disturbance thresholds. Both our theoretical model and empirical study suggest that inefficient individuals in a time-limited environment may be unable to increase their foraging time to compensate for the effects of disturbance. Consequently, the magnitude of individual variation in time budgets strongly determines what proportion of the population is at risk that their condition deteriorates due to disturbance. Our hypothesis implies that, when assessing disturbance effects on a population, it is not only important to consider individual variation in disturbance responses, but also variation in time budgets that determine the consequences of disturbance.

AIM: In many species, density‐dependent effects on reproduction are an important driver of population dynamics. However, it is rarely considered that the direction of density dependence is expected to vary over space and time depending on anti‐predator behaviour and predator community. Aggregation may allow for effective group mobbing against avian nest predators while aggregation may also attract mammalian predators, causing negative density dependence. We aim to quantify spatial variation in the effect of conspecific breeding density on nest survival in a mobbing bird species (Eurasian oystercatcher; Haematopus ostralegus) and identify whether this variation in density dependence can be explained by the predator community. LOCATION: Country‐wide (The Netherlands). METHODS: We integrated reproductive data with breeding territory maps of Eurasian oystercatchers and occupancy maps of avian and mammalian predator species across the Netherlands for a 10‐year period. RESULTS: Spatial variation in the composition of the predator community explained the effects of neighbour density, showing decreasing nest survival when both conspecific density and mammalian dominance increased. Also, heterospecific density (from breeding godwits and lapwing) has an additional effect on the oystercatcher nest survival. Strikingly, this pattern did not extend to mammal‐free island populations. MAIN CONCLUSIONS: Our study provides evidence that both the strength and sign of density dependence can vary spatially within species, implying that it is dangerous to generalize results from a single local population to large‐scale management implications and modelling exercises. The study also suggests that conservation actions that aim to attract breeding birds should be prioritized in areas with fewer mammalian predators, but this idea requires further testing on island populations.

Understanding which factors cause populations to decline begins with identifying which parts of the life cycle, and which vital rates, have changed over time. However, in a world where humans are altering the environment both rapidly and in different ways, the demographic causes of decline likely vary over time. Identifying temporal variation in demographic causes of decline is crucial to assure that conservation actions target current and not past threats. However, this has rarely been studied as it requires long time series. Here we investigate how the demography of a long-lived shorebird (the Eurasian Oystercatcher Haematopus ostralegus) has changed in the past four decades, resulting in a shift from stable dynamics to strong declines (−9% per year), and recently back to a modest decline. Since individuals of this species are likely to respond differently to environmental change, we captured individual heterogeneity through three state variables: age, breeding status, and lay date (using integral projection models). Timing of egg-laying explained significant levels of variation in reproduction, with a parabolic relationship of maximal productivity near the average lay date. Reproduction explained most variation in population growth rates, largely due to poor nest success and hatchling survival. However, the demographic causes of decline have also been in flux over the last three decades: hatchling survival was low in the 2000s but improved in the 2010s, while adult survival declined in the 2000s and remains low today. Overall, the joint action of several key demographic variables explain the decline of the oystercatcher, and improvements in a single vital rate cannot halt the decline. Conservations actions will thus need to address threats occurring at different stages of the oystercatcher's life cycle. The dynamic nature of the threat landscape is further supported by the finding that the average individual no longer has the highest performance in the population, and emphasizes how individual heterogeneity in vital rates can play an important role in modulating population growth rates. Our results indicate that understanding population decline in the current era requires disentangling demographic mechanisms, individual variability, and their changes over time.

To understand the consequences of anthropogenic and environmental changes for wildlife populations, it is important to study how individuals differ in their sensitivity to environmental change and whether this depends on individual characteristics. An individual's reproductive performance may provide an integrative, unidimensional proxy of an individual's characteristics. In this study, we define an individual's characteristics by three such reproductive states, namely successful, failed and non-breeders in the previous year. We used a 16-year dataset of individually marked breeding Eurasian Oystercatchers Haematopus ostralegus to examine the interannual fluctuations in reproductive success and survival among breeding states, and their state-dependent sensitivity to environmental conditions. Environmental conditions included available biomass of the main prey species of breeding Oystercatchers (Ragworm, Baltic Tellin and Lugworm), tidal height, which reflects one of the main causes of nest loss (flooding), and conditions that may impact the energetic requirements during incubation, such as temperature. We also included environmental variables measured in winter, including available biomass of the main winter prey species (Blue Mussel and Common Cockle) along with factors that may affect food availability and energetic requirements for homeostasis, namely bivalve weight loss, windchill, winter severity and precipitation. Breeding birds that were successful the previous year had higher survival and were more likely to remain successful, in comparison with failed or non-breeders. The effects of environmental conditions acted in the same direction on reproductive success but had opposite effects on survival among the three breeding states, especially for windchill and Blue Mussel biomass. The contrasting state-dependent effects of the environment on survival thus averaged out when examining consequences for lifetime reproductive nest success (LRnS); instead, LRnS was largely influenced by environmental conditions acting upon reproduction. Our study indicates that an individual's previous breeding state provides an integrative measure of heterogeneity in individuals' sensitivity of reproduction and survival to environmental change. Incorporating previous breeding state as a source of individual heterogeneity in population modelling may improve predictions of future population dynamics in a rapidly changing world.

Feeding specialization is a common cause of individual variation. Fitness payoffs of specialization vary with environmental conditions, but the underlying behavioral mechanisms are poorly understood. Such mechanistic knowledge, however, is crucial to reliably predict responses of heterogeneous populations to environmental change. We quantified spatiotemporal allocation of foraging behavior in wintering Eurasian oystercatchers (Haematopus ostralegus), a species in which feeding specialization can be inferred from bill shape. We combined global positioning system (GPS) and accelerometer data to quantify foraging time of 64 individuals for every tidal period in one or two winter seasons. Individuals varied widely in foraging time (3.7-6.5 h per tidal period) and individuals that spend more time foraging had lower inferred survival. Feeding specialization appeared a major determinant of individual variation in foraging time and its spatiotemporal allocation. Visually hunting worm specialists foraged more during day time and complemented intertidal foraging with grassland foraging when the exposure of intertidal flats was limited and nights were well illuminated. Shellfish specialists increased total foraging time in cold weather, whereas foraging time of worm specialists decreased as frosty grasslands became inaccessible. Our results imply that worm specialists may be most sensitive to cold snaps and daytime disturbance, whereas shellfish specialists are most sensitive to high water levels. These behavioral responses can be implemented in population models to predict the vulnerability of heterogeneous populations to environmental change and, thereby, provide a shortcut to long-term population studies that require fitness data across many years and conditions to make similar projections. Lay Summary: Populations consist of different types of individuals. Predicting how heterogeneous populations of long-lived species respond to environmental change is hard as it takes decades to quantify how fitness varies among individuals in different environmental conditions. We studied the behavior of GPS-tracked oystercatchers and showed how foraging time is differentially affected by environmental conditions among individuals that specialize in different prey. This knowledge can be implemented in models to predict the vulnerability of heterogeneous populations to environmental change.

Shorebird populations, especially those feeding on shellfish, have strongly declined in recent decades and identifying the drivers of these declines is important for conservation. Changing food stocks are thought to be a key driver of these declines and may also explain why trends have not been uniform across Europe's largest estuary. We therefore investigated how winter population trends of Eurasian oystercatchers (Haematopus ostralegus) were linked to food availability in the Dutch Wadden Sea. Our analysis incorporated two spatial scales, a smaller scale focused on roost counting areas and food available to birds in these areas and a larger spatial scale of tidal basins. A novelty in our study is that we quantify the connectivity between roosting and foraging areas, identified from GPS tracking data. This allowed us to estimate food available to roosting birds and thus how food availability may explain local population trends. At the smaller spatial scale of roost counting areas, there was no clear relationship between available food and the number of roosting oystercatchers, indicating that other factors may drive population fluctuations at finer spatial scales. At the scale of tidal basins, however, there was a significant relationship between population trends and available food, especially cockle Cerastoderma edule,. Mortality and recruitment alone could not account for the large fluctuations in bird counts, suggesting that the site choice of wintering migratory oystercatchers may primarily drive these large fluctuations. Furthermore, the relationship between oystercatcher abundance and benthic food stocks, suggests winter shorebird counts could act as ecological indicators of ecosystem health, informing about the winter status of food stocks at a spatial scale of tidal basins.

In many coastal areas, high numbers of recreationists may exceed ecological capacities. Careful monitoring of visitor flows is a first prerequisite for coastal area management. We show how AIS ship data can be translated into interpretable information on recreational boats and investigate whether AIS can provide monitoring information when compared to nature conservation policy targets. In the Wadden Sea UNESCO World Heritage Site we used nearly 9 million data points to create spatiotemporal patterns for the 2018 recreation season. We combined this with shipping lanes and bathymetry data and compared the resulting patterns with nature protection regulations. Our results show that most of the traffic is concentrated around tidal channels. We also show that exceeding speed limits is not predominant behaviour, but the effect of speeding on birds and seals might be more severe than the data suggests. We mapped favourite tidal flat moor activities, and observed where this occurs in Marine Protected Areas. We conclude that AIS analysis can provide valuable recreational boating monitoring, relevant to sensitive coastal area management in the entire Dutch Wadden Sea for the full recreational season. Broader integration of AIS with radar data and ecological data can add to the power of using AIS.

Changes in the frequency of extreme climatic events (ECEs) can have profound impacts on individual fitness by degrading habitat quality. Organisms may respond to such changes through habitat selection, favouring those areas less affected by ECEs; however, documenting habitat selection in response to ECEs is difficult in the wild due to the rarity of such events and the long-term biological data required. Sea level rise and changing weather patterns over the past decades have led to an increase in the frequency of coastal flooding events, with serious consequences for ground nesting shorebirds. Shorebirds therefore present a useful natural study system to understand habitat selection as a response to ECEs. We used a 32-year study of the Eurasian oystercatcher (Haematopus ostralegus) to investigate whether habitat selection can lead to an increase in nest elevation and minimize the impacts of coastal flooding. The mean nest elevation of H. ostralegus has increased during the last three decades. We hypothesized that this change has been driven by changes in H. ostralegus territory settlement patterns over time. We compared various possible habitat selection cues to understand what information H. ostralegus might use to inform territory settlement. There was a clear relationship between elevation and territory settlement in H. ostralegus. In early years, settlements were more likely at low elevations but in more recent years the likelihood of settlement was similar between high and low elevation areas. Territory settlement was associated with conspecific fledgling output and conspecific density. Settlement was more likely in areas of high density and areas with high fledgling output. This study shows that habitat selection can minimize the effects of increasingly frequent ECEs. However, it seems unlikely that the changes we observe will fully alleviate the consequences of anthropogenic climate change. Rates of nest elevation increase were insufficient to track current increases in maximum high tide (0.5 vs. 0.8 cm/year). Furthermore, habitat selection cues that rely on information from previous breeding seasons (e.g. conspecific fledgling output) may become ineffective as ECEs become more frequent and environmental predictability is diminished.

Background Anthropogenic disturbance can negatively affect an animal’s energy budget by evoking movement responses. Existing research focuses mainly on immediate displacement as a disturbance effect, since this can be easily observed in the field. However, effects on movement over longer timescales are poorly examined and it is largely unknown if and to what extent they reflect immediate responses. Longer-term responses could for example be larger than immediate responses if birds, after disturbance, return to the original location and thereby travel twice the immediate disturbed distance. Methods We combined GPS tracking data with observational data to quantify the effects of anthropogenic (air force and walkers) and non-anthropogenic disturbances on distances travelled by roosting Eurasian Oystercatchers (Haematopus ostralegus) during the non-breeding season. We compared immediate displacement after a disturbance with distance travelled during the entire high tide period (longer-term response), while accounting for environmental factors. Additionally, we calculated energy expenditure due to disturbance based on observed disturbance frequencies. Results Disturbance resulted in an immediate displacement response of ~ 200 m (median). Air force disturbances tended to yield larger immediate responses than walker and, especially, than non-anthropogenic disturbances. Longer-term responses and immediate responses were approximately similar, suggesting that, over longer timescales, spatial disturbance effects in the study area remain confined to immediate effects. However, disturbances were infrequent (0.17 disturbances per bird per hour) and most disturbances were of natural origin (62%). Consequently, anthropogenic disturbance of roosting oystercatchers in the study area on average costs 0.08% of the daily energy expenditure. Conclusions Our results suggest that immediate spatial responses to disturbance can be a useful proxy for spatial responses over longer timescales. Over the non-exhaustive range of conditions investigated, energetic consequences of spatial disturbance responses for an oystercatcher in the study area are marginal due to low disturbance levels.

1. Spatio-temporal variation in population dynamics of migratory populations is shaped by exposure to different environments during the annual cycle. Hence, exposure to similar environments should translate into synchrony in vital rates. Despite a wide-ranging breeding population, the Baltic/Wadden Sea flyway population of eiders (Somateria m. mollissima) shares wintering grounds in the southern Baltic Sea, inner Danish waters and the Wadden Sea; different colonies within this flyway population are therefore likely to exhibit some degree of synchrony in vital rates. 2. Here we used capture-recapture-recovery data to investigate the impact of hunting, winter climate (the North-Atlantic Oscillation Index), winter temperature, nitrogen runoff, autumn-winter body condition of blue mussels Mytilus spp., natural predation and epidemic disease (avian cholera) on annual survival of adult females in ten study colonies distributed between the Netherlands and Finland. Moreover, we tested how the degree of similarity in spatial winter distributions affected the degree of similarity in annual survival among colonies. 3. None of the covariates universally affected female survival. While the quality of blue mussels in the wintering area explained almost 40 % of the variation in survival of eiders breeding on Christianso in the south-western Baltic Sea, incidence of epidemic disease explained > 60 % in two affected colonies. Furthermore, the spatial winter distribution did not appreciably influence annual survival rates in these ten colonies. 4. The lack of universal effects of spatial winter distribution and winter conditions on survival suggests that local breeding conditions may be more important, and hence prime targets for conservation efforts. Better monitoring of e.g. food quality, predation pressure and epidemic disease at the time of breeding could be the key to better understand the population dynamics in this endangered flyway population.

Este artículo contiene 11 páginas, 2 tablas, 4 figuras., Local biodiversity trends over time are likely to be decoupled from global trends, as local processes may compensate or counteract global change. We analyze 161 long-term biological time series (15–91 years) collected across Europe, using a comprehensive dataset comprising ~6,200 marine, freshwater and terrestrial taxa. We test whether (i) local long-term biodiversity trends are consistent among biogeoregions, realms and taxonomic groups, and (ii) changes in biodiversity correlate with regional climate and local conditions. Our results reveal that local trends of abundance, richness and diversity differ among biogeoregions, realms and taxonomic groups, demonstrating that biodiversity changes at local scale are often complex and cannot be easily generalized. However, we find increases in richness and abundance with increasing temperature and naturalness as well as a clear spatial pattern in changes in community composition (i.e. temporal taxonomic turnover) in most biogeoregions of Northern and Eastern Europe., We are grateful to the ILTER network and the eLTER PLUS project (Grand Agreement No. 871128) for financial support. We acknowledge the E-OBS dataset from the EUFP6 project ENSEMBLES (http://ensembles-eu.metoffice.com) and the data providers in the ECA&D project (http://www.ecad.eu). The evaluation of forest plant diversity was based on data collected by partners of the official UNECE ICP Forests Network (http://icp-forests.net/contributors); part of the data were co-financed by the European Commission, project LIFE 07 ENV/D/000218 “Further Development and Implementation of an EU-level Forest monitoring Systeme (FutMon)”. Data on wintering water birds in Bulgaria were provided by the national Executive Environment Agency with the Ministry of Environment and Waters. Data from the Finnish moth monitoring scheme were supported by the Finnish Ministry of the Environment. Data from the Swedish ICP Integrated Monitoring sites were financed by the Swedish Environmental Protection Agency. Data collection at Esthwaite Water and a subset of UK ECN sites was supported by Natural Environment Research Council award number NE/ R016429/1 as part of the UK-SCaPE programme delivering National Capability. Sponsorship of other UK ECN sites contributing data was provided by Agri-Food and Biosciences Institute, Biotechnology and Biological Sciences Research Council, Department of Environment Food and Rural Affairs, Natural Resources Wales, Defense Science Technology Laboratory, Environment Agency, Forestry Commission, Forest Research, the James Hutton Institute (The Rural & Environment Science & Analytical Services Division of the Scottish Government), Natural England, Rothamsted Research, Scottish Government, Scottish Natural Heritage and the Welsh Government. Data from the Mondego estuary (Portugal) were supported by the Centre for Functional Ecology Strategic Project (UID/BIA/04004/2019) within the PT2020 Partnership Agreement and COMPETE 2020, and by FEDER through the project ReNATURE (Centro 2020, Centro-01-765-0145-FEDER-000007). We would like to thank Limburgse Koepel voor Natuurstudie (LiKoNa) for the data related to the National Park Hoge Kempen (BE). We would like to acknowledge the support for the long-term monitoring program MONEOS in the Scheldt estuary (BE) by ‘De Vlaamse Waterweg’ and ‘Maritieme Toegang’ (Flemish government). We are grateful to the board of the National Park “De Hoge Veluwe” for the permission to conduct our research on their property. We thank Ian J. Winfield and Terje Bongard for contributing data for the sites: Bassenthwaite Lake, Derwent Water (UK) and Atna River (Norway, freshwater invertebrate time series). Open access funding provided by Umeå University.

Assessing the impacts of disturbance over large areas and long time periods is crucial for nature management, but also challenging since impacts depend on both wildlife responses to disturbance and on the spatiotemporal distribution of disturbance sources. Combined tracking of animals and disturbance sources enables quantification of wildlife responses as a function of the distance to a disturbance source. We provide a framework to derive such distance–response curves and combine those with disturbance source presence data to quantify energetic costs of disturbance at a landscape scale. We tracked 90 Eurasian Oystercatchers Haematopus ostralegus and all aircraft in a military training area in the Dutch Wadden Sea. We quantified distance–response curves estimating flight probability and additional displacement for five types of aircraft activities, by comparing bird movement prior to aircraft presence with movement during aircraft presence. We then used the distance–response curves to map mean and variation in additional daily energy expenditure due to cumulative aircraft disturbance across the landscape for a 700‐day period. Flight probability and displacement responses differed strongly among aircraft activities and decreased from transport aeroplanes, through bombing jets, helicopters, jets to small civil aeroplanes. Since the most disturbing aircraft activities were also the rarest ones, mean additional daily energy expenditure did not exceed 0.25%. However, days with substantial (>1%) additional expenditure occurred between 0.1% and 3.7% of all days across high‐tide roosts in the tidal basin. Notably, expenditure particularly spiked on days with transport aeroplane activity (up to 8.5%). Synthesis and applications. We quantified cumulative energetic flight costs due to aircraft disturbance and found that these were low and unlikely to impact survival of oystercatchers in our study area. Our results provide evidence that the legal minimum flight height of 450 m for small civil aeroplanes effectively limits the disturbance of oystercatchers. Mitigation should focus on limiting the number of days when disturbance has a high impact by reducing rare but highly disturbing activities, especially transport aeroplanes. Our approach can be applied to other species and disturbance sources that are automatically tracked, for example boats and walkers, ultimately to quantify the entire anthropogenic disturbance landscape.

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.

Vegetation indices derived from satellite image time series have been extensively used to estimate the timing of phenological events like season onset. Medium spatial resolution (≥250 m) satellite sensors with daily revisit capability are typically employed for this purpose. In recent years, phenology is being retrieved at higher resolution (≤30 m) in response to increasing availability of high-resolution satellite data. To overcome the reduced acquisition frequency of such data, previous attempts involved fusion between high- and medium-resolution data, or combinations of multi-year acquisitions in a single phenological reconstruction. The objectives of this study are to demonstrate that phenological parameters can now be retrieved from single-season high-resolution time series, and to compare these retrievals against those derived from multi-year high-resolution and single-season medium-resolution satellite data. The study focuses on the island of Schiermonnikoog, the Netherlands, which comprises a highly-dynamic saltmarsh, dune vegetation, and agricultural land. Combining NDVI series derived from atmospherically-corrected images from RapidEye (5 m-resolution) and the SPOT5 Take5 experiment (10m-resolution) acquired between March and August 2015, phenological parameters were estimated using a function fitting approach. We then compared results with phenology retrieved from four years of 30 m Landsat 8 OLI data, and single-year 100 m Proba-V and 250 m MODIS temporal composites of the same period. Retrieved phenological parameters from combined RapidEye/SPOT5 displayed spatially consistent results and a large spatial variability, providing complementary information to existing vegetation community maps. Retrievals that combined four years of Landsat observations into a single synthetic year were affected by the inclusion of years with warmer spring temperatures, whereas adjustment of the average phenology to 2015 observations was only feasible for a few pixels due to cloud cover around phenological transition dates. The Proba-V and MODIS phenology retrievals scaled poorly relative to their high-resolution equivalents, indicating that medium-resolution phenology retrievals need to be interpreted with care, particularly in landscapes with fine-scale land cover variability.

Background Ring wear and loss may have important consequences for mark-recapture studies that aim to estimate survival trends. Our study quantifies the rates of wear and loss from a long-running colour-ringing project of the Eurasian Oystercatcher (Haematopus ostralegus) in the Netherlands. Methods Our analysis included 8909 colour-ringed oystercatchers with 118,071 resightings, predominantly ringed and observed by citizen scientists. We quantified how ring wear and loss may vary with ring colour and age, and how this may impact resighting probabilities. We then determined how ring loss may impact survival estimates and resighting probabilities of mark-recapture models by simulating four scenarios of how citizen scientists may resight and report birds with lost colour rings. Results Annual rates of colour ring loss averaged 2.5% compared with 1% for ring wear, but lost rings also had a higher reporting probability (31.2%) than worn rings (10.3%). Lost rings may not directly impact survival estimates since 50% of oystercatchers with lost rings could still be uniquely identified. Ring wear and loss rapidly increased between 10 and 15 years after ringing. Rates of ring loss were comparable amongst ring colours, but the wear rate appeared higher for red and white rings compared to other colours. Rates of ring wear and loss varied significantly amongst different regions in our study, which were characterised as having different habitat features. Differential rates of ring wear may have important implications for studies conducted over large geographical scales or of multiple species. Conclusions Based on simulations of ring loss in our population, estimates of apparent survival may be 0.3–1.2% lower whilst the impact of ring wear was deemed even lower. We developed a table of recommendations containing advice for ring fitting, data collection, study design, and mark-recapture analyses, so that future colour-ringing studies can benefit from our experiences in collecting and analysing data of ring wear and loss through citizen science.

The Sentinel satellite fleet of the Copernicus Programme offers new potential to map and monitor plant traits at fine spatial and temporal resolutions. Among these traits, leaf area index (LAI) is a crucial indicator of vegetation growth and an essential variable in biodiversity studies. Numerous studies have shown that the radiative transfer approach has been a successful method to retrieve LAI from remote-sensing data. However, the suitability and adaptability of this approach largely depend on the type of remote-sensing data, vegetation cover and the ecosystem studied. Saltmarshes are important wetland ecosystems threatened by sea level rise among other human- and animal-induced changes. Therefore, monitoring their vegetation status is crucial for their conservation, yet few LAI assessments exist for these ecosystems. In this study, the retrieval of LAI in a saltmarsh ecosystem is examined using Sentinel-2 and RapidEye data through inversion of the PROSAIL radiative transfer model. Field measurements of LAI and some other plant traits were obtained during two succeeding field campaigns in July 2015 and 2016 on the saltmarsh of Schiermonnikoog, a barrier island of the Netherlands. RapidEye (2015) and Sentinel-2 (2016) data were acquired concurrent to the time of the field campaigns. The broadly employed PROSAIL model was inverted using two look-up tables (LUTs) generated in the spectral band’s settings of the two sensors and in which each contained 500,000 records. Different solutions from the LUTs, as well as, different Sentinel-2 spectral subsets were considered to examine the LAI retrieval. Our results showed that generally the LAI retrieved from Sentinel-2 had higher accuracy compared to RapidEye-retrieved LAI. Utilising the mean of the first 10 best solutions from the LUTs resulted in higher R2 (0.51 and 0.59) and lower normalised root means square error (NRMSE) (0.24 and 0.16) for both RapidEye and Sentinel-2 data respectively. Among different Sentinel-2 spectral subsets, the one comprised of the four near-infrared (NIR) and shortwave infrared (SWIR) spectral bands resulted in higher estimation accuracy (R2 = 0.44, NRMSE = 0.21) in comparison to using other studied spectral subsets. The results demonstrated the feasibility of broadband multispectral sensors, particularly Sentinel-2 for retrieval of LAI in the saltmarsh ecosystem via inversion of PROSAIL. Our results highlight the importance of proper parameterisation of radiative transfer models and capacity of Sentinel-2 spectral range and resolution, with impending high-quality global observation aptitude, for retrieval of plant traits at a global scale.

Migratory connectivity describes linkages between breeding and non-breeding areas. An ongoing challenge is tracking avian species between breeding and non-breeding areas and hence estimating migratory connectivity and seasonal survival. Collaborative color-ringing projects between researchers and citizen scientists provide opportunities for tracking the annual movements of avian species. Our study describes seasonal survival and migratory connectivity using data from more than 4,600 individuals with over 51,000 observations, predominantly collected by citizen scientists. Our study focuses on the Eurasian Oystercatcher (Haematopus ostralegus), a species that has experienced a substantial and ongoing decline in recent decades. Multiple threats have been described, and given that these threats vary in space and time, there is an urgent need to estimate demographic rates at the appropriate spatio-temporal scale. We performed a seasonal multi-state (5 geographical areas within The Netherlands) live- and dead-recoveries analysis under varying model structures to account for biological and data complexity. Coastal breeding populations were largely sedentary, while inland breeding populations were migratory and the direction of migration varied among areas, which has not been described previously. Our results indicated that survival was lower during winter than summer and that survival was lower in inland areas compared with coastal areas. A concerning result was that seasonal survival of individuals over-wintering in the Wadden Sea, an internationally important site for over-wintering shorebirds, appeared to decline during the study period. We discuss the outcomes of our study, and how citizen science was integral for conducting this study. Our findings identify how the demographic rates of the oystercatcher vary in space and time, knowledge that is vital for generating hypotheses and prioritizing future research into the causes of decline.

Recruitment and fate of all 1436 mussel and oyster beds of the Dutch Wadden Sea were studied for the period 1999–2013. Cox’s proportional hazard rate model with covariates such as orbital speed, exposure time and bed size and type showed that large, low-lying beds that experience a low orbital speed live longer. Yet the most striking result was that oyster and mixed beds have a much lower hazard rate than pure mussel beds. Simulation studies, using the observed recruitment series, which was very variable, and the estimated survival curves, showed large variability in total bed area, implying that the present area, though lower than before, does not point to a systematic deviation from the pre-1990 situation, that is, the situation before intensive fisheries on these intertidal beds and the disappearance of them around 1990. Claims that bivalve bed recovery is impossible without restoration efforts are premature and not supported by our analysis. On the contrary, the observed high survival rate of mixed and oyster beds and the expectation that such beds will predominate in the near future can easily result in larger future bed coverage than what has been measured before.

Context: Place-based transdisciplinary research involves multiple academic disciplines and non-academic actors. Long-Term Socio-Ecological Research (LTSER) platform is one concept with ~ 80 initiatives globally. Objectives: As an exercise in learning through evaluation we audited (1) the siting, construction and maintenance of individual LTSER platforms, and (2) them as a distributed infrastructure for place-based transdisciplinary research with focus on the European continent. Methods: First, we defined a normative model for ideal performance at both platform and network levels. Second, four surveys were sent out to the 67 self-reported LTSER platforms officially listed at the end of 2016. Third, with a focus on the network level, we analyzed the spatial distribution of both long-term ecological monitoring sites within LTSER platforms, and LTSER platforms across the European continent. Fourth, narrative biographies of 18 platforms in different stages of development were analyzed. Results: While the siting ofLTSER platforms represented biogeographical regions well, variations in land use history and democratic governance were not well represented. Platform construction was based on 2.1 ecological monitoring sites, with 72% ecosystem and 28% social system research. Maintenance of a platform required three to five staff members, focused mostly on ecosystem research, was based mainly on national funding, and had 1–2 years of future funding secured. Networking with other landscape approach concepts was common. Conclusions: Individually, and as a network, LTSER platforms have good potential for transdisciplinary knowledge production and learning about sustainability challenges. To improve the range of variation of Pan-European social–ecological systems we encourage interfacing with other landscape approach concepts.

Many subtidal predators undertake regular tidal migrations into intertidal areas in order to access abundant prey. One of the most productive habitats in soft bottom intertidal systems is formed by beds of epibenthic bivalves such as blue mussels (Mytilus edulis) and Pacific oysters (Crassostrea gigas). In the Dutch Wadden Sea, these bivalves might face substantial predation pressure by the shore crab (Carcinus maenas), which increased considerably in numbers during the last 20 years. However, the quantification of this species on bivalve beds is challenging, since most methods common for quantifying animal abundance in marine habitats cannot be used. This study investigated the potential of two methods to quantify the abundance of C. maenas on 14 epibenthic bivalve beds across the Dutch Wadden Sea. The use of the number of crabs migrating from subtidal towards intertidal areas as a proxy of abundance on bivalve beds yielded unreliable results. In contrast, crabs caught with traps on the beds were correlated with the abundance assessed on the surrounding bare flats by beam trawl and therefore provided usable results. The estimates, however, were only reliable for crabs exceeding 35 mm in carapace width (CW). The application of these estimates indicated that crab abundances on bivalve beds were influenced by the biogenic structure. Beds dominated by oysters attracted many large crabs (> 50-mm CW), whereas abundances of medium-sized crabs (35–50-mm CW) showed no relationship to the oyster occurrence. The combination of traps and trawls is capable of quantifying crab abundance on bivalve beds, which offers the possibility to study biotic processes such as predator-prey interactions in these complex structures in more detail.

Summary 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.

Aim: Molluscivorous shorebirds supposedly developed their present wintering distribution after the last ice age. Currently, molluscivorous shorebirds are abundant on almost all shores of the world, except for those in the Indo-West Pacific (IWP). Long before shorebirds arrived on the scene, molluscan prey in the IWP evolved strong anti-predation traits in a prolonged evolutionary arms race with durophagous predators including brachyuran crabs. Here, we investigate whether the absence of molluscivorous shorebirds from a site in Oman can be explained by the molluscan community being too well-defended. Location: The intertidal mudflats of Barr Al Hikman, Oman. Methods: Based on samples from 282 locations across the intertidal area the standing stock of the macrozoobenthic community was investigated. By measuring anti-predation traits (burrowing depth, size and strength of armour), the fraction of molluscs available to molluscivorous shorebirds was calculated. Results: Molluscs dominated the macrozoobenthic community at Barr Al Hikman. However, less than 17% of the total molluscan biomass was available to shorebirds. Most molluscs were unavailable either because of their hard-to-crush shells, or because they lived too deeply in the sediment. Repair scars and direct observations confirmed crab predation on molluscs. Although standing stock densities of the Barr Al Hikman molluscs were of the same order of magnitude as at intertidal mudflat areas where molluscivorous shorebirds are abundant, the molluscan biomass available to shorebirds was distinctly lower at Barr Al Hikman. Main conclusions: The established strong molluscan anti-predation traits against crabs precludes molluscan exploitation by shorebirds at Barr Al Hikman. This study exemplifies that dispersal of "novel" predators is hampered in areas where native predators and prey exhibit strongly developed attack and defence mechanisms, and highlights that evolutionary arms races can have consequences for the global distribution of species.

Foraging distributions are thought to be density‐dependent, because animals not only select for a high availability and quality of resources, but also avoid conspecific interference. Since these processes are confounded, their relative importance in shaping foraging distributions remains poorly understood. Here we aimed to rank the contribution of density‐dependent and density‐independent effects on the spatio‐temporal foraging patterns of eurasian oystercatchers. In our intertidal study area, tides caused continuous variation in oystercatcher density, providing an opportunity to disentangle conspecific interference and density‐independent interactions with the food landscape. Spatial distributions were quantified using high‐resolution individual tracking of foraging activity and location. In a model environment that included a realistic reconstruction of both the tides and the benthic food, we tested a family of behaviour‐based optimality models against these tracking data. Density‐independent interactions affected spatial distributions much more strongly than conspecific interference, even in an interference‐prone species like oystercatchers. Spatial distributions were governed by avoidance of bill injury costs, selection for high interference‐free intake rates and a decreasing availability of benthic bivalve prey after their exposure. These density‐independent interactions outweighed interference competition in terms of effect size. We suggest that the bottleneck in our mechanistic understanding of foraging distributions may be primarily the role of density‐independent prey attributes unrelated to intake rates, like damage costs in the case of oystercatchers foraging on perilous prey. At a landscape scale, above the finest inter‐individual distances, effects of conspecific interaction on spatial distributions may have been overemphasised.

The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment1,2,3,4. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions1,5, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators6,7,8,9,10. Individuals can temporally segregate their daily activities (for example, prey avoiding predators, subordinates avoiding dominants) or synchronize their activities (for example, group foraging, communal defence, pairs reproducing or caring for offspring)6,7,8,9,11. The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood5,6,7,9. Here we investigate these rhythms in the context of biparental care, a particularly sensitive phase of social synchronization12 where pair members potentially compromise their individual rhythms. Using data from 729 nests of 91 populations of 32 biparentally incubating shorebird species, where parents synchronize to achieve continuous coverage of developing eggs, we report remarkable within- and between-species diversity in incubation rhythms. Between species, the median length of one parent’s incubation bout varied from 1–19 h, whereas period length—the time in which a parent’s probability to incubate cycles once between its highest and lowest value—varied from 6–43 h. The length of incubation bouts was unrelated to variables reflecting energetic demands, but species relying on crypsis (the ability to avoid detection by other animals) had longer incubation bouts than those that are readily visible or who actively protect their nest against predators. Rhythms entrainable to the 24-h light–dark cycle were less prevalent at high latitudes and absent in 18 species. Our results indicate that even under similar environmental conditions and despite 24-h environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of individuals in captivity5,6,7,9. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms., The study was supported by the Max Planck Society (to B.K.). M.B. is a PhD student in the International Max Planck Research School for Organismal Biology.

The purpose of this study is to experimentally determine the differences between four grazing treatments on the trampling of nests. Additionally, we examine to what extent the trampling probability of nests is higher close to a source of fresh water. We compare the trampling of artificial nests in five different grazing treatments in an experimental design. We use buried clay pigeon targets as artificial mimics of bird nests to obtain reliable estimates of trampling risk and compare these with real nests. We find that horses trample significantly more artificial nests than cattle resulting in lower survival rates of artificial nests under horse grazing than under cattle grazing. For both horses and cattle, we find a clear trend, approaching significance, towards more trampling at higher numbers of livestock. We found that more artificial nests are trampled closer to a freshwater tank. The survival probability of artificial nests in cattle grazed treatments in this study is found to be in the same range as real nests in the study area and very close to the survival probability of Northern Lapwing (Vanellus vanellus) nests under cattle grazing in a different system. We recommend that horses should not be used as grazers for management purposes in areas with high densities of birds' nests in order to minimize the risk of nests being trampled. Additionally, we confirm that the location of freshwater tanks has an important effect on the distribution of livestock and hence on trampling of nests.

In this study we examined the effect of different livestock grazing treatments on breeding bird densities in a salt marsh habitat. To avoid an experiment on the large scale needed to directly measure grazing effects on bird densities, we followed a two-step approach. First, we measured vegetation micro-patterns (mosaic of lower vegetation and taller patches at 4x4 m) around Common Redshank Tringa totanus and Eurasian Oystercatcher Haematopus ostralegus nests and at random sites paired with these nests sites to judge suitability of micro-patterns for nest building. Secondly, we measured micro-patterns at 120 permanent plots in five different experimental grazing treatments to determine how grazing affects micro-patterns. We compared low stocking density of both cattle and of horses, high stocking density of cattle and of horses, and intermittent grazing with a high stocking density of cattle (i.e. yearly intervals of grazing and no grazing). Redshank and Oystercatcher nests occurred in sites with taller vegetation and more pronounced micro-patterns than found at random sites. Paddocks grazed with low densities of livestock or with a high density intermittent grazing treatment had micro-patterns preferred by the birds. We conclude that Redshanks and Oystercatchers may benefit in terms of potential nest sites from grazing at low livestock densities or at intermittent stocking densities through effects of grazing on micro-patterns in the vegetation.

Intertidal mussel beds are important for intertidal ecosystems, because they feature a high taxonomic diversity and abundance of benthic organisms and are important foraging grounds for many avian species. After the introduction of the Pacific oyster (Crassostrea gigas) into the European Wadden Sea, many mussel beds developed into oyster dominated bivalve beds. Despite the fact that oysters have been colonizing many European intertidal areas for about two decades, their impact on the ecosystem is still poorly understood. Here, we investigated the impact of oysters on the condition of mussels and on the spatial distribution of birds on 18 bivalve beds with different grades of oyster occurrence throughout the Dutch Wadden Sea. Moreover, in comparing bird densities on bivalve beds with densities expected on the total intertidal area, we could detect which species exhibit a preference for the structured habitat. Overall, 50 different bird species were observed on the beds, of which about half regularly frequent intertidal flats. Most of these species showed a preference for bivalve beds. The condition of mussels decreased with the oyster dominance, whereas the majority of bird species was not affected by the oyster occurrence. However, three of the four species that were negatively affected depend on intertidal mussels as food source. Even though the Pacific oyster is a nonnative species, attempts to fight it may do more harm to avian biodiversity than good.

Migrating birds are believed to minimize the time spent on migration rather than energy. Birds seem to maximize migration speed in different ways as a noteworthy variation in migration strategies exists. We studied migration strategies of a flight mode and feeding generalist, the Lesser Black-backed Gull Larus fuscus, using GPS-based satellite telemetry. We expected the gulls to achieve very high overall migration speeds by traveling via the shortest direct route, traveling during a large part of the day and night, and making few and short stopovers. Fourteen individuals were tracked between the Dutch breeding colony and the wintering sites in England, southern Europe and northwest Africa. The gulls did not travel via the shortest possible route but made substantial detours by their tendency to follow coasts. Although the gulls traveled during most of the day, and sometimes during the night, they did not achieve long daily distances (177 and 176 km/day in autumn and spring, respectively), which is explained by the gulls stopping frequently on travel days to forage. Furthermore, due to frequent and long migratory stopovers, their overall migration speed was among the lowest recorded for migratory birds (44 and 98 km/day, in autumn and spring, respectively). A possible explanation for the unexpected frequent stopovers and low migration speeds is that gulls do not minimize the duration of migration but rather minimize the costs of migration. Energy rather than time might be important for short-distance migrating birds, resulting in very different migration strategies compared with long-distance migrants.

Summary 1. Climate change encompasses changes in both the means and the extremes of climatic variables, but the population consequences of the latter are intrinsically difficult to study. 2. We investigated whether the frequency, magnitude and timing of rare but catastrophic flooding events have changed over time in Europe’s largest estuary. Subsequently, we quantified how this has affected the flooding risk of six saltmarsh nesting bird species. 3. We show that maximum high tide has increased twice as fast as mean high tide over the past four decades (0AE 8v s. 0AE 4c m year )1 ), resulting in more frequent and more catastrophic flooding of nests, especially around the time when most eggs have just hatched. 4. Using data on species’ nest elevations, on their timing of egg-laying and on the duration that their eggs and chicks are at risk from flooding, we show that flooding risks increased for all six studied species (even after accounting for compensatory land accretion) and this is expected to worsen in the near future if they do not adapt. Moreover, our study provides the first evidence that increasing flooding risks have reduced the reproductive output below stable population levels in at least one species, the Eurasian oystercatcher Haematopus ostralegus. 5. Sensitivity analyses show that currently birds would benefit most from adapting their nest-site selection to higher areas. However, historically the lower marsh has been favoured for its proximity to the feeding grounds and for its low vegetation aiding predator detection. 6. Synthesis and applications. We argue that it is more difficult for birds to infer that habitat quality has decreased from changes in the frequency of rare and unpredictable extreme events than from trends in climatic means. Consequently, at present the lower parts of the saltmarsh may function as an ecological trap. The creation of new (i.e. low) saltmarshes – currently a restoration priority – may thus counteract the goal of increasing the avian biodiversity of an area. Management tools to mitigate the effects of climate change, either by making the higher saltmarsh more attractive (mowing, predator control) or by reducing the flooding risk of the lower marsh (building elevated plots), await to be tested.

Migrants, such as birds or representatives of other taxa, usually make use of several stopover sites to cover the distance between their site of origin and destination. Potentially, multiple routes exist, but often little is known about the causes and consequences of alternative migration routes. Apart from their geographical distribution, the suitability of potential sites might play an important role in the animals' decisions for a particular itinerary. We used an optimal-migration model to test three nonmutually exclusive hypotheses leading to variations in the spring migration routes of a subspecies of Red Knot, Calidris canutus islandica, which migrates from wintering grounds in Western Europe to breeding grounds in Greenland and the Canadian Arctic: the breeding location hypothesis, the energy budget hypothesis, and the predation risk hypothesis. Varying only breeding location, the model predicted that birds breeding in the Canadian Arctic and on West Greenland stop over on Iceland, whereas birds breeding in East and Northeast Greenland migrate via northern Norway, a prediction that is supported by empirical findings. Energy budgets on stopover sites had a strong influence on the choice of route and staging times. Varying foraging-intensity and mass-dependent predation risk prompted the birds to use less risky sites, if possible. The effect of simultaneous changes in the energy budget and predation risk strongly depended on the site where these occurred. Our findings provide potential explanations for the observations that C. canutus islandica uses a diverse array of migration routes. Scrutinizing the three alternative driving forces for the choice of migratory routes awaits further, specific data collection in rapidly developing fields of research (e.g., predation risk assessment, GPS tracking). Generally, the type of modeling presented here may not only highlight alternative explanations, but also direct follow-up empirical research.

Climate change affects both the mean and variability of climatic variables, but their relative impact on the dynamics of populations is still largely unexplored. Based on a long-term study of the demography of a declining Eurasian Oystercatcher (Haematopus ostralegus) population, we quantify the effect of changes in mean and variance of winter temperature on different vital rates across the life cycle. Subsequently, we quantify, using stochastic stage-structured models, how changes in the mean and variance of this environmental variable affect important characteristics of the future population dynamics, such as the time to extinction. Local mean winter temperature is predicted to strongly increase, and we show that this is likely to increase the population's persistence time via its positive effects on adult survival that outweigh the negative effects that higher temperatures have on fecundity. Interannual variation in winter temperature is predicted to decrease, which is also likely to increase persistence time via its positive effects on adult survival that outweigh the negative effects that lower temperature variability has on fecundity. Overall, a 0.1 degrees C change in mean temperature is predicted to alter median time to extinction by 1.5 times as many years as would a 0.1 degrees C change in the standard deviation in temperature, suggesting that the dynamics of oystercatchers are more sensitive to changes in the mean than in the interannual variability of this climatic variable. Moreover, as climate models predict larger changes in the mean than in the standard deviation of local winter temperature, the effects of future climatic variability on this population's time to extinction are expected to be overwhelmed by the effects of changes in climatic means. We discuss the mechanisms by which climatic variability can either increase or decrease population viability and how this might depend both on species' life histories and on the vital rates affected. This study illustrates that, for making reliable inferences about population consequences in species in which life history changes with age or stage, it is crucial to investigate the impact of climate change on vital rates across the entire life cycle. Disturbingly, such data are unavailable for most species of conservation concern.

Fluctuating and disruptive selection are important mechanisms for maintaining intrapopulation trait variation. Nonetheless, few field studies quantify selection pressures over long periods and identify what causes them to fluctuate. Diet specialists in oystercatchers differ in short-term payoffs (intake), but their long-term payoffs are hypothesized to be condition dependent. We test whether phenotypic selection on diet specialization fluctuates between years due to the frequency of specialists, competitor density, prey abundance, and environmental conditions. Short-term payoffs proved to be poor predictors of long-term fitness payoffs of specialization. Sex-differences in diet specialization were maintained by opposing directional fecundity and viability selection between the sexes. Contrasting other studies, selection on individual diet specialization was neither negative frequency- or density-dependent nor dependent on prey abundance. Notwithstanding, viability selection fluctuated strongly (stabilizing disruptive) over the 26-year study period: slightly favoring generalists in most years, but strongly disfavoring generalists in rare harsh winters, suggesting generalists cannot cope with extreme conditions. Although selection fluctuated, mean selection on specialists was weak, which can explain how individual specialization can persist over long periods. Because rare events can dramatically affect long-term selective landscapes, more care should be taken to match the timescale of evolutionary studies to the temporal variability of critical environmental conditions.

Interference competition is often due to kleptoparasitism (food stealing). In which case, the attack distance, the distance over which one animal attacks another in an attempt to steal food, determines to a large extent the competitor density range over which interference significantly affects the intake rate of foraging animals. We develop a simple model of kleptoparasitism containing three parameters: attack distance, the density of foraging animals and a single dimensionless parameter α which summarizes the non-geometrical aspects of the interference process. Dominant and subdominant animals are not considered separately. The model predicts that the average intake rate will decrease exponentially with animal density and that a measure of the strength of interference depends on attack distance squared. The simple model is compared with a much more detailed individual-based foraging model from the literature. Simulated average intake rates are indeed well approximated by an exponential decrease with competitor density. Also the measure of interference behaves in the way expected from the simple model. By explaining the shape of the relationship between intake rate and animal density, the simple model provides insight into the behaviour of the detailed behavioural model. Insight into the role of geometry is important in the interpretation of field results and in the further development of detailed foraging models.

Eurasian Oystercatchers Haematopus ostralegus ostralegus are a classic example of individual feeding specialization. Feeding specialization causes morphological differentiation in Oystercatchers' bill shapes due to varying degrees of abrasion associated with specific handling techniques for each prey species. Consequently, the Oystercatcher's bill shape has been used as a proxy for diet specialization, as it provides a quick and easy way to assess the diet choice of this marine top-predator. However, bill shapes of Oystercatchers are categorized visually in distinct types, while it has been argued that the relevant variation is continuous. Also, it is unclear how comparable the bill-shape classification is among studies and between the sexes and how universal bill shape-diet relationships are. Here we investigate the usefulness of bill-shape types as a proxy for diet choice in Oystercatchers, using four new and two published datasets. We show that quantitative bill-morphometrics provide no evidence that b...

Molecular sexing of birds has been possible for over a decade, but for practical reasons many studies still use biometric data for sex discrimination. In some species, the sexes are easy to distinguish but sexual dimorphism is often more subtle, requiring the use of statistical analyses of biometric measurements to discriminate sexes. These models are usually parameterized and validated using data from a limited number of sites and years. However, the resulting discriminant functions are often applied to other populations and periods. A crucial, but usually untested, assumption is that sexual dimorphism does not vary in time and space. Here we illustrate the consequences of violation of this assumption in Eurasian Oystercatchers Haematopus ostralegus, a species for which most studies have relied on biometric sexing. Using biometric data from captures of known-sex birds, we show that sexual dimorphism varied substantially in time and even reversed in some months and years. Furthermore, some biometric traits used in sexing changed gradually over time, causing a reduction in sexual dimorphism. We show that the consequences of this variation on sex discrimination in Oystercatchers are subtle and easily overlooked, but can result in inaccurate and strongly male- or female-biased sex-ratio estimates. We recommend that biometric sexing should be avoided in Oystercatchers unless specific calibration for each month, year and area is carried out. This recommendation also applies to other species where biometric traits may depend on environmental conditions. We argue that this condition might apply to many bird species and therefore advise caution when interpreting results based on biometric sex discrimination.

Our study investigated how bird species richness and abundance was related to livestock grazing on salt, and brackish marshes, with an emphasis on songbirds, and shorebirds. Survey areas with a high percentage cover of tall vegetation were assumed to have experienced lower livestock grazing intensities than survey areas with a low percentage cover of tall vegetation. This relationship was verified for the tall grass Elytrigia atherica. The species richness, and abundance of birds was related to the percentage cover of tall vegetation on the survey areas. We found that total bird species richness was positively related to the percentage cover of tall vegetation. We also found that all of the investigated species, except Pied Avocet (Recurvirostra avosetta), showed a positive relation to the percentage cover of tall vegetation up to a specific percentage of cover. The abundance of investigated songbird species increased up to an intermediate percentage cover of tall vegetation, and decreased at higher percentage cover of tall vegetation, suggesting that moderate grazing of marshes may maximize the abundance of the investigated songbirds. Abundances of Common Redshank (Tringa totanus) and Eurasian Oystercatcher (Haematopus ostralegus) were positively related to the percentage cover of tall vegetation on salt marshes, but negatively related to the percentage cover of tall vegetation on brackish marshes. With intermediate livestock grazing species number, and abundance of most breeding birds can be maintained in coastal marshes. However, specific goals for management should be set before applying a grazing regime to a marsh.

Intake rate maximization alone is not always sufficient in explaining prey size selection in predators. For example, bivalve-feeding oystercatchers regularly select smaller prey than expected if they aimed to maximize their intake rate. It has been proposed that to these birds large prey are ‘‘risky,’’ in the sense that birds may damage their bills when feeding on large bivalves. Large bivalves yield more energy, but according to this hypothesis this is achieved at the expense of energy yield in the long term when (1) the risk of bill damage increases with prey size and (2) foraging with a damaged bill is less effective. In accordance with this hypothesis, we show that captive oystercatchers feeding on large cockles experienced a high probability of bill tip damage, while bill damage was absent when cockles were small. Moreover, among free-living oystercatchers the prevalence of bill damage was correlated with mean cockle size near the capture site, and the data on captive birds fit in this pattern. Food intake of captive oystercatchers feeding exclusively on cockles was reduced by 23% after bill damage, and free-living birds with damaged bills had 14 g lower mass. Because lower body mass was associated with higher mortality probability, these results indicate long-term costs associated with feeding on large cockles. We conclude that the risk of bill damage can potentially explain why oystercatchers avoid large bivalves and that oystercatchers may maximize long-term intake rate by selecting prey sizes that are ‘‘suboptimal’’ from a short-term rate-maximizing point of view. Key words: prey choice, prey profitability, prey size selection, risky prey. [Behav Ecol]

Wader species wintering on the Banc d'Arguin increased their body mass by about 40% during the 4–6 weeks before their departure in spring. This estimate is based on 1) the empirical fact that most waders which had finished or suspended their body moult into summer plumage were heavy and thus ready to take off, and 2) the assumption that only the heaviest birds in the population left, which allowed the mass of disappearing waders to be estimated from counts and the frequency distribution of body masses in samples of captured birds. The mass gain on the Banc d'Arguin is just over 1% per day, when expressed as a proportion of winter mass. A review of studies on waders preparing for migration shows that 1) the total migratory reserve adds 20–80% to winter mass, 2) the rate of mass gain is 0.1–4% per day and 3. the period of mass increase lasts four weeks on average, but longer if waders prepare for spring migration on the wintering areas. We suggest that all wader species leaving the B anc d'Arguin a...

Animals moving through a complex environment change their movement patterns frequently. Such transitions from one movement behavior to another are a result of multiple factors such as weather, habitat and inter-specific interactions occurring either individually or in tandem. Understanding where and when these changes occur is the first step towards understanding the limiting or key factors acting at different scales. We applied discrete wavelet analysis to find change points in the movement behavior of a Lesser Black-backed gull ( Larus fuscus ). We compared our results with results from the residence method proposed by Barraquand and Benhamou (2008) and the statistical method proposed by Lavielle (1999, 2005). Discrete wavelet analysis allows the identification and localization of change points by decomposing the frequency content of two time series consisting of step length and residence time data, while omitting the problem of redundancy of coefficients consistent with continuous wavelet transformation of movement data. We show that the novel use of discrete wavelet analysis in animal movement studies is widely applicable on different focus variables, allowing changes in the movement behavior and precise change points to be detected.

Human interests often conflict with those of wildlife. In the coastal zone humans often exploit shellfish populations that would otherwise provide food for populations of shorebirds (Charadrii). There has been considerable debate on the consequences of shellfishing for the survival of shorebirds, and conversely the effects of shorebird predation on the shellfish stocks remaining for human exploitation. Until now, it has been difficult to determine the impact of current shellfishery practices on birdsor to investigate how possible alternative policies would affect their survival and numbers. One long-running contentious issue has been how to manage mussel Mytilus edulis and cockle Cerastoderma edule shellfisheries in a way that has least effect on a co-dependent shorebird, the oystercatcher Haematopus ostralegus, which also consumes these shellfish. This study used a behaviour-based model to explore the effects that the present-day management regimes of a mussel (Exe estuary, UK) and a cockle (Burryinlet, UK) fishery have on the survival and numbers of overwintering oystercatchers. It also explored how alternative regimes might affect the birds. The model includes depletion and disturbance as two possibly detrimental effects of shellfishing and some of the longer-term effects on shellfish stocks. Importantly, model birds respond to shellfishing in the same ways as real birds. They increase the time spent feeding at low tide and feed in fields and upshore areas at other times. When shellfishing removes the larger prey, birds eat more smaller prey. The results suggest that, currently, neither shellfishery causes oystercatcher mortality to be higher than it would otherwise be in the absence of shellfishing; at present intensities, shellfishing does not significantly affect the birds. However, they also show that changes in management practices, such as increasing fishing effort, reducing the minimum size of shellfish collected or increasing the daily quota, can greatly affect oystercatcher mortality and population size, and that the detrimental effect of shellfishing can be greatly increased by periods of cold weather or when prey are unusually scarce. By providing quantitative predictions of bird survival and numbers of a range of alternative shellfishery management regimes, the model can guide management policy in these and other estuaries.