Survival costs of reproduction are independent of energy costs in a seabird, the pelagic cormorant.

Life-history theory predicts that investment in reproduction should decrease survival (the 'cost of reproduction'). It is often assumed that energy allocation drives such trade-offs, with limited energy available for both reproduction and survival. However, the underlying mechanisms remain poorly understood, maybe because survival costs of reproduction are only apparent when resources are limited. Here, we took advantage of a natural experiment created by fluctuating environmental conditions to compare energy expenditure of a seabird, the pelagic cormorant (Phalacrocorax pelagicus), between contrasting population-scale scenarios of survival costs of reproduction. We used multi-state capture-recapture modelling across 16 years to identify which breeding seasons induced high survival costs (survival ratebreeders < survival ratenon/failed breeders) and we concomitantly estimated energy expenditure of chick-rearing males using time-energy budget models across 4 years. Daily energy expenditure (DEE) of chick-rearing pelagic cormorants varied significantly among years. However, survival costs of reproduction were observed in only 1 year, and contrary to our expectations, variation in DEE was not associated with population-level survival costs. Similarly, at the individual level, DEE in 1 year did not predict the probability of being observed again at the colony in following years (apparent survival). Finally, DEE was independent of brood size and brood age, but older individuals tended to expend less energy than younger ones. Given the lack of an apparent energetic 'cost of reproduction', lower DEE in older birds could be due to improved efficiency rather than avoidance of costs in old birds. Although future studies should account for potential sex-specific energetic constraints by including data on female energy expenditure, we conclude that a direct link between the rate of energy expenditure during breeding and subsequent survival is unlikely in this system.

Red knots in Europe: a dead end host species or a new niche for highly pathogenic avian influenza?

The 2020/2021 epidemic in Europe of highly pathogenic avian influenza virus (HPAIV) of subtype H5 surpassed all previously recorded European outbreaks in size, genotype constellations and reassortment frequency and continued into 2022 and 2023. The causative 2.3.4.4b viral lineage proved to be highly proficient with respect to reassortment with cocirculating low pathogenic avian influenza viruses and seems to establish an endemic status in northern Europe. A specific HPAIV reassortant of the subtype H5N3 was detected almost exclusively in red knots (Calidris canutus islandica) in December 2020. It caused systemic and rapidly fatal disease leading to a singular and self-limiting mass mortality affecting about 3500 birds in the German Wadden Sea, roughly 1 % of the entire flyway population of islandica red knots. Phylogenetic analyses revealed that the H5N3 reassortant very likely had formed in red knots and remained confined to this species. While mechanisms of virus circulation in potential reservoir species, dynamics of spill-over and reassortment events and the roles of environmental virus sources remain to be identified, the year-round infection pressure poses severe threats to endangered avian species and prompts adaptation of habitat and species conservation practices.

Variations in inter-specific and sex-related niche partitioning in pelagic boobies during their annual cycle.

Animals that co-occur in a region (sympatry) may share the same environment (syntopy), and niche differentiation is expected among closely related species competing for resources. The masked booby (Sula dactylatra) and smaller congeneric red-footed booby (Sula sula) share breeding grounds. In addition to the inter-specific size difference, females of both species are also larger than the respective males (reversed sexual size dimorphism). Although both boobies consume similar prey, sometimes in mixed-species flocks, each species and sex may specialize in terms of their diet or foraging habitats. We examined inter- and intra-specific differences in isotopic values (δ13C and δ15N) in these pelagically feeding booby species during the incubation period at Clarion Island, Mexico, to quantify the degrees of inter- and intra-specific niche partitioning throughout the annual cycle. During incubation, both species preyed mainly on flyingfish and squid, but masked boobies had heavier food loads than red-footed boobies. There was no overlap in isotopic niches between masked and red-footed boobies during breeding (determined from whole blood), but there was slight overlap during the non-breeding period (determined from body feathers). Female masked boobies had a higher trophic position than conspecific males during breeding; however, no such pattern was detected in red-footed boobies. These results provide evidence of inter- and intra-specific niche partitioning in these tropical seabird species, particularly during the breeding period and in the more-dimorphic species. Our results suggest that these closely related species use different strategies to cope with the same tropical marine environment.

Behavioral responses to offshore windfarms during migration of a declining shorebird species revealed by GPS-telemetry.

EU member countries and the UK are currently installing numerous offshore windfarms (OWFs) in the Baltic and North Seas to achieve decarbonization of their energy systems. OWFs may have adverse effects on birds; however, estimates of collision risks and barrier effects for migratory species are notably lacking, but are essential to inform marine spatial planning. We therefore compiled an international dataset consisting of 259 migration tracks for 143 Global Positioning System-tagged Eurasian curlews (Numenius arquata arquata) from seven European countries recorded over 6 years, to assess individual response behaviors when approaching OWFs in the North and Baltic Seas at two different spatial scales (i.e. up to 3.5 km and up to 30 km distance). Generalized additive mixed models revealed a significant small-scale increase in flight altitudes, which was strongest at 0-500 m from the OWF and which was more pronounced during autumn than during spring, due to higher proportions of time spent migrating at rotor level. Furthermore, four different small-scale integrated step selection models consistently detected horizontal avoidance responses in about 70% of approaching curlews, which was strongest at approximately 450 m from the OWFs. No distinct, large-scale avoidance effects were observed on the horizontal plane, although they could possibly have been confounded by changes in flight altitudes close to land. Overall, 28.8% of the flight tracks crossed OWFs at least once during migration. Flight altitudes within the OWFs overlapped with the rotor level to a high degree in autumn (50%) but to a significantly lesser extent in spring (18.5%). Approximately 15.8% and 5.8% of the entire curlew population were estimated to be at increased risk during autumn and spring migration, respectively. Our data clearly show strong small-scale avoidance responses, which are likely to reduce collision risk, but simultaneously highlight the substantial barrier effect of OWFs for migrating species. Although alterations in flight paths of curlews due to OWFs seem to be moderate with respect to the overall migration route, there is an urgent need to quantify the respective energetic costs, given the massive ongoing construction of OWFs in both sea areas.

Large-scale effects of offshore wind farms on seabirds of high conservation concern.

The North Sea is a key area worldwide for the installation of offshore wind farms (OWFs). We analysed data from multiple sources to quantify the effects of OWFs on seabirds from the family Gaviidae (loons) in the German North Sea. The distribution and abundance of loons changed substantially from the period before to the period after OWF construction. Densities of loons were significantly reduced at distances of up to 9-12 km from the OWF footprints. Abundance declined by 94% within the OWF + 1 km zone and by 52% within the OWF + 10 km zone. The observed redistribution was a large-scale effect, with birds aggregating within the study area at large distances from the OWFs. Although renewable energies will be needed to provide a large share of our energy demands in the future, it is necessary to minimize the costs in terms of less-adaptable species, to avoid amplifying the biodiversity crisis.

Active breeding seabirds prospect alternative breeding colonies.

Compared to other animal movements, prospecting by adult individuals for a future breeding site is commonly overlooked. Prospecting influences the decision of where to breed and has consequences on fitness and lifetime reproductive success. By analysing movements of 31 satellite- and GPS-tracked gull and tern populations belonging to 14 species in Europe and North America, we examined the occurrence and factors explaining prospecting by actively breeding birds. Prospecting in active breeders occurred in 85.7% of studied species, across 61.3% of sampled populations. Prospecting was more common in populations with frequent inter-annual changes of breeding sites and among females. These results contradict theoretical models which predict that prospecting is expected to evolve in relatively predictable and stable environments. More long-term tracking studies are needed to identify factors affecting patterns of prospecting in different environments and understand the consequences of prospecting on fitness at the individual and population level.

Effects of agricultural practices on foraging habitats of a seabird species in the Baltic Sea.

Omnivorous and opportunistic species may be good indicators of food availability. Gulls often use human-impacted landscapes and may respond to changes by altering their feeding ecology. We investigated the foraging behavior of individual common gulls (Larus canus), focusing on their distribution during foraging and their selected habitat types. We tracked adult common gulls using GPS telemetry at their largest breeding colony in the southwestern Baltic Sea, Germany. Foraging habitats were analyzed from tracking data for three breeding seasons 2016, 2017, and 2019 and were compared with potentially available foraging habitats. Most breeding birds flew toward terrestrial areas. Feeding sites were located on average 11.7-14.3 km from the colony (range 0.9-36.5 km). Corn and sugar beet fields were used significantly and extensively compared with their availability in 2016 and 2017, while wheat, rape, and barley fields were used significantly less. Data from 2019 suggested seasonal shifts in habitat use. Birds spent between 30 and 1300 min per week at their preferred feeding sites, with significant differences between the major habitats selected. We found a stable, clear, multiyear pattern in common gull foraging behavior in relation to agricultural practices. Fields with little or no crop cover and thus access to the soil were preferred over fields with high crop cover. These results suggest that local food availability may be limiting further population increases in this species.

Timing of spring departure of long distance migrants correlates with previous year's conditions at their breeding site.

Precise timing of migration is crucial for animals targeting seasonal resources at locations encountered across their annual cycle. Upon departure, long-distance migrants need to anticipate unknown environmental conditions at their arrival site, and they do so with their internal annual clock. Here, we tested the hypothesis that long-distance migrants synchronize their circannual clock according to the phenology of their environment during the breeding season and therefore adjust their spring departure date according to the conditions encountered at their breeding site the year before. To this end, we used tracking data of Eurasian curlews from different locations and combined movement data with satellite-extracted green-up dates at their breeding site. The spring departure date was better explained by green-up date of the previous year, while arrival date at the breeding site was better explained by latitude and longitude of the breeding site, suggesting that other factors impacted migration timing en route. On a broader temporal scale, our results suggest that long-distance migrants may be able to adjust their migration timing to advancing spring dates in the context of climate change.

Risks to different populations and age classes of gannets from impacts of offshore wind farms in the southern North Sea.

The southern North Sea holds the world's highest concentration of offshore wind farms (OWFs). Northern gannets (Morus bassanus), a species considered at high risk from OWF impacts, show a strong seasonal peak there in November, but it is unclear which populations and age classes are most at risk of collision with wind turbines. We tagged adult and juvenile gannets at the world's largest colony (Bass Rock) and reviewed two sources of survey data for different age classes to study their movements through southern North Sea waters. Tracked birds showed peak numbers in the southern North Sea in mid-October, with much smaller numbers there during November. Adults were distributed throughout the area, including waters close to OWFs, whereas juveniles were confined to the coast. Survey data indicated high proportions of immature gannets in southern North Sea waters, suggesting higher collision risk than for adults. Gannets present in November may be predominantly from colonies further north than Bass Rock.

Analysis of local habitat selection and large-scale attraction/avoidance based on animal tracking data: is there a single best method?

BACKGROUND: New wildlife telemetry and tracking technologies have become available in the last decade, leading to a large increase in the volume and resolution of animal tracking data. These technical developments have been accompanied by various statistical tools aimed at analysing the data obtained by these methods. METHODS: We used simulated habitat and tracking data to compare some of the different statistical methods frequently used to infer local resource selection and large-scale attraction/avoidance from tracking data. Notably, we compared spatial logistic regression models (SLRMs), spatio-temporal point process models (ST-PPMs), step selection models (SSMs), and integrated step selection models (iSSMs) and their interplay with habitat and animal movement properties in terms of statistical hypothesis testing. RESULTS: We demonstrated that only iSSMs and ST-PPMs showed nominal type I error rates in all studied cases, whereas SSMs may slightly and SLRMs may frequently and strongly exceed these levels. iSSMs appeared to have on average a more robust and higher statistical power than ST-PPMs. CONCLUSIONS: Based on our results, we recommend the use of iSSMs to infer habitat selection or large-scale attraction/avoidance from animal tracking data. Further advantages over other approaches include short computation times, predictive capacity, and the possibility of deriving mechanistic movement models.

Migrating curlews on schedule: departure and arrival patterns of a long-distance migrant depend on time and breeding location rather than on wind conditions.

BACKGROUND: Departure decisions in long-distance migratory bird species may depend on favourable weather conditions and beneficial resources at the destination location, overarched by genetic triggers. However, few studies have tried to validate the significance of these three concepts simultaneously, and long-term, high-resolution tagging datasets recording individual movements across consecutive years are scarce. We used such a dataset to explore intraspecific and intra-individual variabilities in departure and arrival decisions from/to wintering grounds in relation to these three different concepts in bird migration. METHODS: We equipped 23 curlews (Numenius arquata) wintering in the Wadden Sea with Global Positioning System data loggers to record their spatio-temporal patterns of departure from and arrival at their wintering site, and the first part of their spring migration. We obtained data for 42 migrations over 6 years, with 12 individuals performing repeat migrations in consecutive years. Day of year of departure and arrival was related to 38 meteorological and bird-related predictors using the least absolute shrinkage and selection operator (LASSO) to identify drivers of departure and arrival decisions. RESULTS: Curlews migrated almost exclusively to Arctic and sub-Arctic Russia for breeding. They left their wintering site mainly during the evening from mid- to late April and returned between the end of June and mid-July. There was no difference in departure times between the sexes. Weather parameters did not impact departure decisions; if departure days coincided with headwind conditions, the birds accounted for this by flying at higher altitudes of up to several kilometres. Curlews breeding further away in areas with late snowmelt departed later. Departures dates varied by only < 4 days in individual curlews tagged over consecutive years. CONCLUSIONS: These results suggest that the trigger for migration in this long-distance migrant is largely independent of weather conditions but is subject to resource availability in breeding areas. The high intra-individual repeatability of departure days among subsequent years and the lack of relationship to weather parameters suggest the importance of genetic triggers in prompting the start of migration. Further insights into the timing of migration in immatures and closely related birds might help to further unravel the genetic mechanisms triggering migration patterns.

Northern gannets (Morus bassanus) are strongly affected by operating offshore wind farms during the breeding season.

Northern gannets (Morus bassanus) have been ranked as one of the most vulnerable species in terms of collision with offshore wind farm (OWF) turbines, and strong avoidance of OWFs has been documented for this species. Gannets increasingly encounter OWFs within the ranges of their largest breeding colonies along the European coasts. However, information on their actual reactions to OWFs during the breeding season is lacking. We investigated the possible effects of OWFs located 23-35 km north of the colony on Helgoland in the southern North Sea on breeding gannets. GPS tags were applied to 28 adult gannets breeding on Helgoland for several weeks over 2 years. Most gannets (89%) predominantly avoided the OWFs in both years, but 11% frequently entered them when foraging or commuting between the colony and foraging areas. Flight heights inside the OWFs were close to the rotor-blade zone, especially for individuals predominantly avoiding the OWFs. Gannets preferred distances of 250-450 m to the turbines when being inside the OWF. A point process modelling approach revealed that the gannets resource selection of the OWF area compared with the surroundings (outside OWF = up to 15 km from the OWF border) was reduced by 21% in 2015 and 37% in 2016. This study provides the first detailed characterisation of individual reactions of gannets to OWFs during the breeding season and one of the first comprehensive studies of OWF effects on this species based on telemetry data. The documented effects need to be considered during the planning processes for future OWFs, especially those located close to large seabird breeding colonies.

Effects of offshore windfarms on seabird abundance: Strong effects in spring and in the breeding season.

The increasing development of offshore wind farms (OWFs) worldwide leads to possible conflicts with the ecological requirements of top predators that largely depend on offshore areas. Seabird species exhibit different behavioural reactions to OWFs, ranging from avoidance resulting in habitat loss, to attraction causing an increased risk of colliding with the turbines. We investigated how OWFs affected the densities and distributions of guillemots and kittiwakes breeding in the southern North Sea and if the effects varied among seasons using a 'before-after control impact' (BACI) analysis approach based on a large-scale and long-term dataset covering 14 years before and 3 years after the construction of OWFs. Guillemot relative density in the OWF decreased by 63% in spring, and by 44% in the breeding season. Kittiwake relative density in the OWF decreased by 45% in the breeding season, and not significantly by 10% in spring. We furthermore estimated the response radii to the OWF for both species and seasons, finding that guillemots showed a response radius of ~9 km in spring and kittiwakes a radius of ~20 km in the breeding season. The results underline the value of large-scale and long-term assessments considering seasonal variation throughout the yearly cycle. The here provided information on the seasonally different reactions of seabirds to OWFs adds substantially to our current knowledge and provides the necessary basis for reliable estimations of OWF effects on guillemots and kittiwakes. Such evaluations are urgently needed for future planning and management recommendations to decision-makers.

So far, so good Similar fitness consequences and overall energetic costs for short and long-distance migrants in a seabird.

Although there is a consensus about the evolutionary drivers of animal migration, considerable work is necessary to identify the mechanisms that underlie the great variety of strategies observed in nature. The study of differential migration offers unique opportunities to identify such mechanisms and allows comparisons of the costs and benefits of migration. The purpose of this study was to compare the characteristics of short and long-distance migrations, and fitness consequences, in a long-lived seabird species. We combined demographic monitoring (survival, phenology, hatching success) of 58 Northern Gannets (Morus bassanus) breeding on Bonaventure Island (Canada) and biologging technology (Global Location Sensor or GLS loggers) to estimate activity and energy budgets during the non-breeding period for three different migration strategies: to the Gulf of Mexico (GM), southeast (SE) or northeast (NE) Atlantic coast of the U.S. Survival, timing of arrival at the colony and hatching success are similar for short (NE, SE) and long-distance (GM) migrants. Despite similar fitness consequences, we found, as expected, that the overall energetic cost of migration is higher for long-distance migrants, although the daily cost during migration was similar between strategies. In contrast, daily maintenance and thermoregulation costs were lower for GM migrants in winter, where sea-surface temperature of the GM is 4-7o C warmer than SE and NE. In addition, GM migrants tend to fly 30 min less per day in their wintering area than other migrants. Considering lower foraging effort and lower thermoregulation costs during winter for long-distance migrants, this suggests that the energetic benefits during the winter of foraging in the GM outweigh any negative consequences of the longer-distance migration. These results support the notion that the costs and benefits of short and long-distance migration is broadly equal on an annual basis, i.e. there are no apparent carry-over effects in this long-lived bird species, probably because of the favourable conditions in the furthest wintering area.

Operational offshore wind farms and associated ship traffic cause profound changes in distribution patterns of Loons (Gavia spp.).

Seabirds select suitable habitats at sea, but these habitats may be strongly impacted by marine spatial planning, including the construction of offshore wind farms (OWFs) and the associated ship traffic. Loons (Gavia spp.) are particularly vulnerable to anthropogenic activities and are also of high conservation status, making them particularly relevant to marine planning processes. We investigated the effects of OWF construction and ship traffic on Loon distributions in the German North Sea on a large spatial scale, using a 'before-after' control impact analysis approach and a long-term data set. Many OWFs were built in or close to core areas of Loon distributions. Loons showed significant shifts in their distribution in the 'after' period and subsequently aggregated between two OWF clusters, indicating the remaining suitable habitat. The decrease in Loon abundance became significant as far as about 16 km from the closest OWF. Ship traffic also had a significant negative impact on Loons, indicating that OWFs deterred Loons through the combined effect of ship traffic and the wind turbines themselves. This study provides the first analysis of the extensive effects of OWFs and ships on Loons on a large spatial scale. The results provide an essential baseline for future marine spatial planning processes in the German North Sea and elsewhere.

Intercolony variations in movement patterns and foraging behaviors among herring gulls (Larus argentatus) breeding in the eastern Wadden Sea.

Herring gulls (Larus argentatus) are opportunistic predators that prefer to forage in the intertidal zone, but an increasing degree of terrestrial foraging has recently been observed. We therefore aimed to analyze the factors influencing foraging behavior and diet composition in the German Wadden Sea. Gulls from three breeding colonies on islands at different distances from the mainland were equipped with GPS data loggers during the incubation seasons in 2012-2015. Logger data were analyzed for 37 individuals, including 1,115 foraging trips. Herring gulls breeding on the island furthest from the mainland had shorter trips (mean total distance = 12.3 km; mean maximum distance = 4.2 km) and preferred to feed on the tidal flats close to the colony, mainly feeding on common cockles (Cerastoderma edule) and shore crabs (Carcinus maenas). In contrast, herring gulls breeding close to the mainland carried out trips with a mean total distance of 26.7 km (mean maximum distance = 9.2 km). These gulls fed on the neobiotic razor clams (Ensis leei) in the intertidal zone, and a larger proportion of time was spent in distant terrestrial habitats on the mainland, feeding on earthworms. δ13C and δ15N values were higher at the colony furthest from the mainland and confirmed a geographical gradient in foraging strategy. Analyses of logger data, pellets, and stable isotopes revealed that herring gulls preferred to forage in intertidal habitats close to the breeding colony, but shifted to terrestrial habitats on the mainland as the tide rose and during the daytime. Reduced prey availability in the vicinity of the breeding colony might force herring gulls to switch to feed on razor clams in the intertidal zone or to use distant terrestrial habitats. Herring gulls may thus act as an indicator for the state of the intertidal system close to their breeding colony.

Decreasing δ13C and δ15N values in four coastal species at different trophic levels indicate a fundamental food-web shift in the southern North and Baltic Seas between 1988 and 2016.

Marine ecosystems are exposed to increasing human pressures and climatic change worldwide. It has therefore become essential to describe ecosystem statuses with respect to multinational protection schemes, often necessitating long-term monitoring programmes. Changes in the food-web structure, which can be monitored via stable isotope measurements, represent an important descriptor of the status of marine ecosystems. We investigated long-term changes (29 years) in isotopic values (δ13C and δ15N) in four indicative organisms at different trophic levels in the southern North and Baltic Seas: bladderwrack (Fucus vesiculosus), blue mussel (Mytilus ssp.), eelpout (Zoarces viviparus), and herring gull (Larus argentatus). Time series analyses using generalised additive models revealed largely consistent declines in δ13C and δ15N throughout all trophic levels of the coastal food web at all study sites, indicating a clear change in these coastal regions from 1988 to 2016. There were no clear long-term patterns in egg biometrics for herring gulls, except for a consistent increase in eggshell thickness. The declines in stable isotope values were in line with the results of previous long-term studies of single higher-trophic-level species, which suggested that the noted changes were mainly caused by altered foraging patterns of the studied species. The current results demonstrate that declines in δ13C and δ15N have occurred throughout the whole food web, not just in particular species. We discuss the possible reasons for the decrease in stable isotope values, including decreasing eutrophication and an increase in terrestrial carbon sources.

Interaction between birds and macrofauna within food webs of six intertidal habitats of the Wadden Sea.

The determination of food web structures using Ecological Network Analysis (ENA) is a helpful tool to get insight into complex ecosystem processes. The intertidal area of the Wadden Sea is structured into diverse habitat types which differ in their ecological functioning. In the present study, six different intertidal habitats (i.e. cockle field, razor clam field, mud flat, mussel bank, sand flat and seagrass meadow) were analyzed using ENA to determine similarities and characteristic differences in the food web structure of the systems. All six systems were well balanced between their degree of organization and their robustness. However, they differed in their detailed features. The cockle field and the mussel bank exhibited a strong dependency on external imports. The razor clam field appeared to be a rather small system with low energy transfer. In the mud flat microphytobenthos was used as a main food source and the system appeared to be sensitive to perturbations. Bird predation was the most pronounced in the sand flat and the seagrass meadow and led to an increase in energy transfer and parallel trophic cycles in these habitats. Habitat diversity appears to be an important trait for the Wadden Sea as each subsystem seems to have a specific role in the overall functioning of the entire ecosystem.

A fundamental study revisited: Quantitative evidence for territory quality in oystercatchers (Haematopus ostralegus) using GPS data loggers.

A fundamental study by Ens et al. (1992, Journal of Animal Ecology, 61, 703) developed the concept of two different nest-territory qualities in Eurasian oystercatchers (Haematopus ostralegus, L.), resulting in different reproductive successes. "Resident" oystercatchers use breeding territories close to the high-tide line and occupy adjacent foraging territories on mudflats. "Leapfrog" oystercatchers breed further away from their foraging territories. In accordance with this concept, we hypothesized that both foraging trip duration and trip distance from the high-tide line to the foraging territory would be linearly related to distance between the nest site and the high tide line. We also expected tidal stage and time of day to affect this relationship. The former study used visual observations of marked oystercatchers, which could not be permanently tracked. This concept model can now be tested using miniaturized GPS devices able to record data at high temporal and spatial resolutions. Twenty-nine oystercatchers from two study sites were equipped with GPS devices during the incubation periods (however, not during chick rearing) over 3 years, providing data for 548 foraging trips. Trip distances from the high-tide line were related to distance between the nest and high-tide line. Tidal stage and time of day were included in a mixing model. Foraging trip distance, but not duration (which was likely more impacted by intake rate), increased with increasing distance between the nest and high-tide line. There was a site-specific effect of tidal stage on both trip parameters. Foraging trip duration, but not distance, was significantly longer during the hours of darkness. Our findings support and additionally quantify the previously developed concept. Furthermore, rather than separating breeding territory quality into two discrete classes, this classification should be extended by the linear relationship between nest-site and foraging location. Finally, oystercatcher's foraging territories overlapped strongly in areas of high food abundance.