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Animals frequently exhibit consistent among-individual differences in behavioural and physiological traits that are inherently flexible. Why should individuals differ consistently in their expression of labile traits? Recently, positive feedbacks between state and behaviour have been proposed as a possible explanation for the maintenance of consistent among-individual differences in both state and behaviour. If state affects behaviour, and behaviour reciprocally affects state, then differences in either state or behaviour that arise among individuals even by chance could be maintained over extended periods of time., We tested for positive feedbacks experimentally using wild-caught red knots ( Calidris canutus islandica). In the wild, knots exhibit consistent among-individual differences in digestive physiology (the mass of the muscular part of the stomach, the gizzard) and foraging behaviour (diet), two inherently labile traits., Experimentally manipulated diet quality had a large effect on gizzard mass. Experimentally manipulated gizzard mass reciprocally influenced total food eaten during ad libitum trials., The effect of gizzard mass on diet choice, though in the predicted direction, was not statistically significant. Individuals exhibited consistent differences in foraging behaviour of unknown origin independent of current gizzard mass, as well as large residual unexplained variance in foraging behaviour. These two sources of variation in foraging behaviour overruled the gizzard mass-dependent foraging behaviour and hence eroded the treatment-related differences in gizzard mass., We conclude that positive feedbacks between diet choice and gizzard mass play at best a limited role in maintaining among-individual variation in gizzard mass in red knots. Furthermore, we suggest that many models of state-behaviour feedbacks likely overestimate their potential importance in maintaining long-term among-individual variation in labile traits because most models of state-behaviour feedbacks fail to account for the effects of additional factors that may act to disrupt the feedback loops., The among-individual differences in diet choice observed during solitary foraging trials eroded the consistent among-individual differences in gizzard mass observed following periods of staple diet treatments in which knots foraged in social groups. Social foraging interactions may play an important role determining the expression of foraging behaviours such as intake rate that in turn influence gizzard mass. Further studies are needed to experimentally test the role of social interactions as a mechanism generating consistent among-individual differences in foraging behaviours and gizzard mass., A is available for this article. [ABSTRACT FROM AUTHOR]

Because of the complications in achieving the necessary long-term observations and experiments, the nature and adaptive value of seasonal time-keeping mechanisms in long-lived organisms remain understudied. Here we present the results of a 20-year-long study of the repeated seasonal changes in body mass, plumage state, and primary molt of 45 captive red knots Calidris canutus islandica, a High Arctic breeding shorebird that spends the nonbreeding season in temperate coastal areas. Birds kept outdoors and experiencing the natural photoperiod of the nonbreeding area maintained sequences of life-cycle stages, roughly following the timing in nature. For 6 to 8 years, 14 of these birds were exposed to unvarying ambient temperature (12 °C) and photoperiodic conditions (12:12 LD). Under these conditions, for at least 5 years they expressed free-running circannual cycles of body mass, plumage state, and wing molt. The circannual cycles of the free-running traits gradually became longer than 12 months, but at different rates. The prebreeding events (onset and offset of prealternate molt and the onset of spring body mass increase) occurred at the same time of the year as in the wild population for 1 or several cycles. As a result, after 4 years in 12:12 LD, the circannual cycles of prealternate plumage state had drifted less than the cycles of prebasic plumage state and wing molt. As the onset of body mass gain drifted less than the offset, the period of high body mass became longer under unvarying conditions. We see these differences between the prebreeding and postbreeding life-cycle stages as evidence for adaptive seasonal time keeping in red knots: the life-cycle stages linked to the initiation of reproduction rely mostly on endogenous oscillators, whereas the later stages rather respond to environmental conditions. Postbreeding stages are also prone to carryover effects from the earlier stages. [ABSTRACT FROM AUTHOR]

Negative density-dependence is generally studied within a single trophic level, thereby neglecting its effect on higher trophic levels. The 'functional response' couples a predator's intake rate to prey density. Most widespread is a type II functional response, where intake rate increases asymptotically with prey density; this predicts the highest predator densities at the highest prey densities. In one of the most stringent tests of this generality to date, we measured density and quality of bivalve prey (edible cockles Cerastoderma edule) across 50 km² of mudflat, and simultaneously, with a novel time-of-arrival methodology, tracked their avian predators (red knots Calidris canutus). Because of negative density-dependence in the individual quality of cockles, the predicted energy intake rates of red knots declined at high prey densities (a type IV, rather than a type II functional response). Resource-selection modelling revealed that red knots indeed selected areas of intermediate cockle densities where energy intake rates were maximized given their phenotype-specific digestive constraints (as indicated by gizzard mass). Because negative density-dependence is common, we question the current consensus and suggest that predators commonly maximize their energy intake rates at intermediate prey densities. Prey density alone may thus poorly predict intake rates, carrying capacity and spatial distributions of predators. [ABSTRACT FROM AUTHOR]

Migratory shorebirds inhabit environments that may yield contrasting salinity-temperature regimes-with widely varying osmoregulatory demands, even within a given species-and the question is: by which physiological means and at which organisational level do they show adjustments with respect to these demands? Red knots Calidris canutus winter in coastal areas over a range of latitudes. The nominal subspecies winters in salty areas in the tropics, whereas the subspecies Calidris canutus islandica winters in north-temperate regions of comparatively lower salinities and temperatures. In this study, both subspecies of red knot were acclimated to different salinity (28/40 ‰)-temperature (5/35 °C) combinations for 2-week periods. We then measured food/salt intakes, basal metabolic rate (BMR), body mass and temperature, fat and salt gland scores, gizzard mass, heat-shock proteins, heterophils/lymphocytes (H/L) ratio and plasma Na to assess the responses of each taxon to osmoregulatory challenges. High salinity (HS)-warm-acclimated birds reduced food/salt intake, BMR, body mass, fat score and gizzard mass, showing that salt/heat loads constrained energy acquisition rates. Higher salt gland scores in saltier treatments indicated that its size was adjusted to higher osmoregulatory demands. Elevated plasma Na and H/L ratio in high-salinity-warm-acclimated birds indicated that salt/heat loads might have a direct effect on the water-salt balance and stress responses of red knots. Subspecies had little or no effect on most measured parameters, suggesting that most adjustments reflect phenotypic flexibility rather than subspecific adaptations. Our results demonstrate how salinity and temperature affect various phenotypic traits in a migrant shorebird, highlighting the importance of considering these factors jointly when evaluating the environmental tolerances of air-breathing marine taxa. [ABSTRACT FROM AUTHOR]

Under the ideal-free distribution, omniscient individuals with similar habitat requirements that are free to move should be distributed such that no individual can improve fitness by changing sites; deviations would indicate trade-offs and constraints on ranging behavior. We studied site occupancy and annual survival in red knots Calidris c. canutus at their main wintering area Banc d’Arguin, Mauritania. We collected mark-resighting data at 2 high-tide roosts (A and B) that were only 3 km apart and within sight. Birds were faithful to their roosts and foraged in nearby intertidal areas, with no overlap between birds from A and B. Shellfish-rich seagrass beds were of greater abundance for birds roosting at A than at B. During 8 winters, we found different sex ratios (48% and 58% males at A and B, respectively) and different proportion of juveniles (22% and 45%) at the 2 roosts. Adult annual survival was higher at A (0.83±0.01 standard error [SE]) than at B (0.81±0.03). Though rare, between winter season movements were 3 times more frequent from B to A than vice versa, indicating that knots can assess the differences in site quality: birds behaved as if they were “ideal”. As larger females and older birds occurred more at A, differences in competitive ability might maintain the site occupancy pattern. As females return from the high Arctic breeding grounds first, and adults return before juveniles, priority of occupancy may also play a role. Such an advantage of arriving earlier would represent a seasonal carryover effect. [ABSTRACT FROM PUBLISHER]

Summary 1. Birds of marine environments have specialized glands to excrete salt, the saltglands. Located on the skull between the eyes, the size of these organs is expected to reflect their demand, which will vary with water turnover rates as a function of environmental (heat load, salinity of prey and drinking water) and organismal (energy demand, physiological state) factors. On the basis of inter- and intraspecific comparisons of saltgland mass ( msg) in 29 species of shorebird (suborder Charadrii) from saline, fresh and mixed water habitats, we assessed the relative roles of organism and environment in determining measured msg species. 2. The allometric exponent, scaling dry msg to shorebird total body mass ( mb), was significantly higher for coastal marine species (0·88, N = 19) than for nonmarine species (0·43, N = 14). Within the marine species, those ingesting bivalves intact had significantly higher msg than species eating soft-bodied invertebrates, indicating that seawater contained within the shells added to the salt load. 3. In red knots ( Calidris canutus), dry msg varied with monthly averaged ambient temperature in a U-shaped way, with the lowest mass at 12·5 °C. This probably reflects increased energy demand for thermoregulation at low temperatures and elevated respiratory water loss at high temperatures. In fuelling bar-tailed godwits ( Limosa lapponica), dry msg was positively correlated with intestine mass, an indicator of relative food intake rates. These findings suggest once more that saltgland masses vary within species (and presumably individuals) in relation to salt load, that is a function of energy turnover (thermoregulation and fuelling) and evaporative water needs. 4. Our results support the notion that msg is strongly influenced by habitat salinity, and also by factors influencing salt load and demand for osmotically free water including ambient temperature, prey type and energy intake rates. Saltglands are evidently highly flexible organs. The small size of saltglands when demands are low suggests that any time costs of adjustment are lower than the costs of maintaining a larger size in this small but essential piece of metabolic machinery. [ABSTRACT FROM AUTHOR]

Local studies have shown that the distribution of red knots Calidris canutus across intertidal mudflats is consistent with the predictions of an ideal distribution, but not a free distribution. Here, we scale up the study of feeding distributions to their entire wintering area in western Europe. Densities of red knots were compared among seven wintering sites in The Netherlands, UK and France, where the available mollusc food stocks were also measured and from where diets were known. We tested between three different distribution models that respectively assumed (i) a uniform distribution of red knots over all areas, (ii) a uniform distribution across all suitable habitat (based on threshold densities of harvestable mollusc prey), and (iii) an ideal and free distribution (IFD) across all suitable habitats. Red knots were not homogeneously distributed across the different European wintering areas, also not when considering suitable habitats only. Their distribution was best explained by the IFD model, suggesting that the birds are exposed to interference and have good knowledge about their resource landscape at the spatial scale of NW Europe, and that the costs of movement between estuaries, at least when averaged over a whole winter, are negligible. [ABSTRACT FROM AUTHOR]

Phenotypic flexibility in shorebirds has been studied mainly in the context of adjustments to migration and to quality of food; little is known on how birds adjust their phenotype to harsh winter conditions. We showed earlier that red knot (Calidris canutus islandica) can acclimate to cold by elevating body mass. This goes together with larger pectoral muscles, i.e., greater shivering machinery, and thus, better thermogenic capacity. Here, we present results of a yearlong experiment with indoor captive knots to determine whether this strategy is part of their natural seasonal phenotypic cycle. We maintained birds under three thermal regimes: constant cold (5°C), constant thermoneutrality (25°C) and natural seasonal variation between these extremes (9–22°C). Each month we measured variables related to the birds’ endurance to cold and physiological maintenance [body mass, thickness of pectoral muscles, summit metabolic rate (Msum), food intake, gizzard size, basal metabolic rate (BMR)]. Birds from all treatments expressed synchronized and comparable variation in body mass in spite of thermal treatments, with a 17–18% increase between the warmest and coldest months of the year; which appeared regulated by an endogenous driver. In addition, birds living in the cold exhibited a 10% higher average body mass than did those maintained at thermoneutrality. Thickness of the pectoral muscle tracked changes in body mass in all treatments and likely contributed to greater capacity for shivering in heavier birds. Consequently, Msum was 13% higher in cold-acclimated birds compared to those experiencing no thermoregulation costs. However, our data also suggest that part of maximal heat production comes from nonshivering processes. Birds facing cold conditions ate up to 25% more food than did birds under thermoneutral conditions, yet did not develop larger gizzards. Seasonal variation in BMR followed changes in body mass, probably reflecting changes in mass of metabolically active tissues. Just as cold-exposed birds, red knots in the variable treatment increased body mass in winter, thereby improving cold endurance. During summer, however, they maintained a lower body mass and thermogenic capacity compared to cold-exposed birds, similar to individuals kept at thermoneutrality. We conclude that red knots acclimate to seasonal variations in ambient temperature by modulating body mass, combining a preprogrammed increase in mass during winter with a capacity for fine-tuning body mass and thermogenic capacity to temperature variations. [ABSTRACT FROM AUTHOR]

The article presents a study which examines the factors that prompted Red knots birds to stop over in Iceland and northern Norway during its migration. It notes the use of the species' stable carbon isotypes in whole blood, blood cells and plasma ratios to generate evidences for its stopover activities. The result indicates that the levels of the bird's blood cells and plasma played a significant role which prompted it to stop over in the area.

Phenotypic flexibility is a phenomenon where physiological functions in animals are reversibly adjusted in response to ecological constraints. Research usually focuses on effects of single constraints, but under natural conditions animals face a multitude of restrictions acting simultaneously, and potentially generating conflicting demands on the phenotype. We investigated the conflicting demands of low temperatures and a low quality diet on the phenotype of a shorebird, the red knot Calidris canutus. We tested the effects of switching diet from a high quality trout food to low quality hard-shelled bivalves in captive birds acclimated to temperatures reflecting natural winter conditions. Feeding on bivalves generated a digestive constraint forcing the birds to increase the height and width of their gizzard by 66% and 71%, respectively, over 30 days. The change in gizzard size was associated with an initial 15% loss of body mass and a reduction in size of the pectoral muscles by 11%. Because pectoral muscle size determines summit metabolic rate (Msum, an indicator of cold endurance), measured Msum declined by 9%. Therefore, although the birds were acclimated to cold, gizzard growth led to a loss of cold endurance. We propose that cold-acclimated knots facing a digestive constraint made a phenotypic compromise by giving-up cold hardiness for digestive capacity. Field studies suggest that phenotypic compromises occur in free-living red knots as well and help improve survival. [ABSTRACT FROM AUTHOR]

1. Millions of shorebirds migrate each year through a small number of highly productive staging areas where they often conflict with fisheries interests. Delaware Bay, USA, is a major shorebird stopover site where, in spring, many thousands of shorebirds undergo rapid mass gain by feeding on the eggs of commercially harvested horseshoe crabs Limulus polyphemus. 2. Environmental factors may cause deviations from the best migration schedule. We used within-year mass gain data from red knot Calidris canutus caught in Delaware Bay between 1998 and 2005 to determine the degree of flexibility individuals have to vary migration speed. 3. Mass gain by birds below 133 g was shown to comprise 15·3% fat (39 kJ g−1), the remainder being lean mass (6 kJ g−1). Above this critical level, fat comprised 83·9% of mass deposition. The rates of energy deposition (kJ d−1) were therefore fundamentally different between the two states but were among the highest ever recorded among vertebrates (5–7 × basic metabolic rate). 4. A total of 36–62% of the variation in observed rates of energy deposition between 1998 and 2002 was explained by a year factor, date and mass at initial capture and interaction terms, such that light-weight birds at the end of May had rates of mass gain or energy deposition two to three times higher than birds of similar mass in mid-May, indicating that birds were attempting to achieve a certain mass by a certain date. In 2003 and 2005, this relationship broke down as a result of lower densities of eggs. 5. Synthesis and application. The maintenance of high densities of crab eggs required for high rates of mass gain in red knot requires severe cuts in, or the complete cessation of, the crab harvest, reduced human and raptor-related disturbance as well as management of beaches to provide sufficient crab-spawning habitat. These findings are widely applicable to other systems where harvesting activities come into conflict with migrating animals and show that certain sections of the population, in this case the long-distance migrants from South America, will be impacted more than short-distance migrants whose physiology may give them access to alternative food resources. [ABSTRACT FROM AUTHOR]

In the calidrine sandpiper red knot (Calidris canutus), the weeks preceding takeoff for long-distance migration are characterized by a rapid increase in body mass, largely made up of fat but also including a significant proportion of lean tissue. Before takeoff, the pectoral muscles are known to hypertrophy in preparation for endurance flight without any specific training. Because birds facing cold environments counterbalance heat loss through shivering thermogenesis, and since pectoral muscles represent a large proportion of avian body mass, we asked the question whether muscle hypertrophy in preparation for long-distance endurance flight would induce improvements in thermogenic capacity. We acclimated red knots to different controlled thermal environments: 26°C, 5°C, and variable conditions tracking outdoor temperatures. We then studied within-individual variations in body mass, pectoral muscle size (measured by ultrasound), and metabolic parameters [basal metabolic rate (BMR) and summit metabolic rate (Msum)] throughout a 3-mo period enclosing the migratory gain and loss of mass. The gain in body mass during the fattening period was associated with increases in pectoral muscle thickness and thermogenic capacity independent of thermal acclimation. Regardless of their thermal treatment, birds showing the largest increases in body mass also exhibited the largest increases in Msum. We conclude that migratory fattening is accompanied by thermoregulatory side effects. The gain of body mass and muscle hypertrophy improve thermogenic capacity independent of thermal acclimation in this species. Whether this represents an ecological advantage depends on the ambient temperature at the time of fattening. [ABSTRACT FROM AUTHOR]

Besides the ‘normal’ challenge of obtaining adequate intake rates in a patchy and dangerous world, shorebirds foraging in intertidal habitats face additional environmental hurdles. The tide forces them to commute between a roosting site and feeding grounds, twice a day. Moreover, because intertidal food patches are not all available at the same time, shorebirds should follow itineraries along the best patches available at a given time. Finally, shorebirds need additional energy stores in order to survive unpredictable periods of bad weather, during which food patches are covered by extreme tides. In order to model such tide- specific decisions, we applied stochastic dynamic programming in a spatially explicit context. Two assumptions were varied, leading to four models. First, birds had either perfect (ideal) or no (non-ideal) information about the intake rate at each site. Second, traveling between sites was either for free or incurred time and energy costs (non-free). Predictions were generated for three aspects of foraging: area use, foraging routines, and energy stores. In general, non-ideal foragers should feed most intensely and should maintain low energy stores. If traveling for such birds is free, they should feed at a random site; otherwise, they should feed close to their roost. Ideal foragers should concentrate their feeding around low tide (especially when free) and should maintain larger energy stores (especially when non-free). If traveling for such birds is free, they should feed at the site offering the highest intake rate; otherwise, they should trade off travel costs and intake rate. Models were parameterized for Red Knots (Calidris canutus) living in the Dutch Wadden Sea in late summer, an area for which detailed, spatially explicit data on prey densities and tidal heights are available. Observations of radio-marked knots (area use) and unmarked knots (foraging routines, energy stores) showed the closest match with the ideal/non-free model. We conclude that knots make state-dependent decisions by trading off starvation against foraging-associated risks, including predation. Presumably, knots share public information about resource quality that enables them to behave in a more or less ideal manner. We suggest that our modeling approach may be applicable in other systems where resources fluctuate in space and time. [ABSTRACT FROM AUTHOR]

We have used the isotope dilution technique to study changes in the body composition of a migratory shorebird species (Red Knot, Calidris canutus ) through an assessment of the amount of body water in it. Birds were quantitatively injected with a dose of water with elevated concentrations of 2 H, 17 O, and 18 O. Thereafter, blood samples were taken and distilled. The resulting water samples were analysed using an isotope ratio mass spectrometry (for 2 H and 18 O only) and a stable isotope ratio infrared laser spectrometry ( 2 H, 17 O, and 18 O) to yield estimates of the amount of body water in the birds, which in turn could be correlated to the amount of body fat. Here, we validate laser spectrometry against mass spectrometry and show that all three isotopes may be used for body water determinations. This opens the way to the extension of the doubly labelled water method, used for the determination of energy expenditure, to a triply labelled water method, incorporating an evaporative water loss correction on a subject-by-subject basis or, alternatively, the reduction of the analytical errors by statistically combining the 17 O and 18 O measurements. [ABSTRACT FROM AUTHOR]

1. When prey occur at high densities, energy assimilation rates are generally constrained by rates of digestion rather than by rates of collection (i.e. search and handle). As predators usually select patches containing high prey densities, rates of digestion will play an important role in the foraging ecology of a species.2. The red knotCalidris canutusshows tremendous inter- and intra-individual variation in maximum rates of digestion due to variation in the size of the processing machinery (gizzard and intestine), which makes it a suitable species to study the effects of digestive processing rate on foraging decisions.3. Here we report on patch use, prey choice, and daily foraging times as a function of gizzard size in free-ranging, radio-marked, red knots. As knots crush their bulky bivalve prey in their gizzard, the size of this organ, which we measured using ultrasonography, determines digestive processing rate.4. Using the digestive rate model, we a priori modelled patch use, prey choice, and daily foraging times as a function of gizzard mass. Focusing on two contrasting patches, birds with small gizzards were expected to feed on high-quality (soft-bodied) prey found in low densities in the one patch, while birds with large gizzards were expected to feed on low-quality (hard-shelled) prey found in high densities in the other patch. Assuming that red knots aim to balance their energy budget on a daily basis, we expected daily foraging time to decline with gizzard mass.5. Observed patch and prey choices were in quantitative agreement with these theoretical predictions. Observed daily foraging times were only in qualitative agreement: they declined with gizzard mass but less steeply than predicted.6. We discuss that red knots might be aiming for a slightly positive energy budget in order to (i) refuel their stores depleted during migration, and (ii) to insure against unpredictability in supply and demand during winter. Red knots arriving from their breeding grounds with small gizzards are only able to realize this aim when densities of soft-bodied prey are high, which is the case in late July and early August. Rapidly declining soft-bodied prey densities throughout late summer pose a major penalty for individuals arriving late at their wintering grounds.7. The long daily foraging periods required by knots with small gizzards are only feasible through‘tide-extension’. In our study area, birds can and do raise the daily low tide period from 12 h to almost 17 h by moving along with the tide in an easterly direction, subsequently flying back to their starting point at the high tide roost.Journal of Animal Ecology(2005) doi: 10.1111/j.1365-2656.2004.00904.x [ABSTRACT FROM AUTHOR]

We measured basal metabolic rate (BMR), body mass, lean mass, and gizzard mass of captive red knots Calidris canutus islandica maintained on a trout chow diet (soft-texture, low ash and water content) for several years and then shifted to small mussels Mytilus edulis (hard-texture, high ash and water content). During a 3-week period of feeding on mussels, body mass, lean mass, and gizzard mass increased 7.3 g (+7%), 10.5 g (+12%), and 4.9 g (+213%), respectively, yet BMR decreased from 0.96 to 0.89 W (−8%). Under the new mussel regime, red knots must have reduced the metabolic intensity of some of the tissues. This suggests that the experimental red knots experienced the transition to a mussel diet as stressful and energy limiting, resulting in an energy-saving strategy by reducing BMR in spite of hypertrophy of the gizzard and other organs. [ABSTRACT FROM AUTHOR]

Studies reversibility and time course of changes in the gizzard size of red knots, Calidris canutus. Effect of hard diet of shellfish on gizzard size; Effect of diet of soft foods on gizzard size; Number of days for gizzard size and mass to change from hard to soft food versus soft food to hard food.

Focuses on a study which evaluated the application of deuterium as a tracer to measure water turnover rates in the Red Knot shorebird. Methods; Changes in body mass and total body water estimates; Behavioral observations.

Recent insights suggest that predators should include (mildly) toxic prey when non-toxic food is scarce. However, the assumption that toxic prey is energetically as profitable as non-toxic prey misses the possibility that non-toxic prey have other ways to avoid being eaten, such as the formation of an indigestible armature. In that case, predators face a trade-off between avoiding toxins and minimizing indigestible ballast intake. Here, we report on the trophic interactions between a shorebird (red knot, Calidris canutus canutus) and its two main bivalve prey, one being mildly toxic but easily digestible, and the other being non-toxic but harder to digest. A novel toxin-based optimal diet model is developed and tested against an existing one that ignores toxin constraints on the basis of data on prey abundance, diet choice, local survival and numbers of red knots at Banc d'Arguin (Mauritania) over 8 years. Observed diet and annual survival rates closely fit the predictions of the toxin-based model, with survival and population size being highest in years when the non-toxic prey is abundant. In the 6 of 8 years when the non-toxic prey is not abundant enough to satisfy the energy requirements, red knots must rely on the toxic alternative. [ABSTRACT FROM AUTHOR]

In birds and mammals, juvenile and adult foragers are often found apart from each other. In this study, we found this is also true for red knots, Calidris canutus canutus, wintering on the intertidal flats of Banc d'Arguin, Mauritania. Not only did juveniles feed separately from adults, they also fed at places where they were more vulnerable to predation by large falcons. That the dangerous areas used by juveniles were no better feeding areas led us to reject the food–safety trade-off that explained age-related distribution differences in many earlier studies. Instead, juveniles were displaced by adults in dyadic interactions which suggests that they suffered from interference from adults. Juveniles retreated to feeding areas that were more dangerous and yielded lower intake rates, and coped by extending foraging time by using higher, nearshore intertidal areas that were exposed for longer. When disturbed by predators in these nearshore areas, juveniles continued feeding whereas adults left. Thus, rather than compensating for increased predation danger by higher intake rates, on the Banc d'Arguin red knot juveniles foraged for longer. [ABSTRACT FROM AUTHOR]