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Deciding when to depart on long-distance, sometimes global, movements can be especially important for flying species. Adverse weather conditions can affect energetic flight costs and navigational ability. While departure timings and conditions have been well-studied for migratory flights to and from the breeding range, few studies have focussed on flights within the non-breeding season. Yet in some cases, overwintering ranges can be large enough that ecological barriers, and a lack of resting sites en route, may resist movement, especially in unfavorable environmental conditions. Understanding the conditions that will enable or prohibit flights within an overwintering range is particularly relevant in light of climate change, whereby increases in extreme weather events may reduce the connectivity of sites. We tracked 495 ( n  = 251 in 2019; n  = 244 in 2020) overwintering red knots ( Calidris canutus islandica ) in the Dutch Wadden Sea and investigated how many departed towards the UK (on westward relocation flights), which requires flying over the North Sea. For those that departed, we used a resource selection model to determine the effect of environmental conditions on the timing of relocation flights. Specifically, we investigated the effects of wind, rain, atmospheric pressure, cloud cover, and migratory timing relative to sunset and tidal cycle, which have all been shown to be crucial to migratory departure conditions. Approximately 37% (2019) and 36% (2020) of tagged red knots departed on westward relocation flights, indicating differences between individuals' space use within the overwintering range. Red knots selected for departures between 1 and 2.5 h after sunset, approximately 4 h before high tide, with tailwinds and little cloud cover. However, rainfall and changes in atmospheric pressure appear unimportant. Our study reveals environmental conditions that are important for relocation flights across an ecological barrier, indicating potential consequences of climate change on connectivity., Competing Interests: The authors declare no conflicts of interest., (© 2024 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Variation in foraging tactics and diet is usually attributed to differences in morphology, experience and prey availability. Recently, consistent individual differences in behaviour (personality) have been shown to be associated with foraging strategies. Bolder or more exploratory individuals are predicted to have a faster pace-of-life and offset the costs of moving more or in risky areas, with higher energetic gains by encountering profitable foraging opportunities and prey. However, the relationship between personality, foraging and diet is poorly understood. We investigated how exploratory behaviour in red knots Calidris canutus is associated with foraging tactics and diet by combining laboratory experiments, field observations and stable isotope analysis. First, we developed a mobile experimental arena to measure exploration speed in controlled settings. We validated the method by repeated testing of individuals over time and contexts. This setup allowed us to measure exploratory personality at the field site, eliminating the need to bring birds into captivity for long periods of time. After releasing birds within days of their capture, we asked whether exploration speed was associated with differences in foraging tactics and diet in the wild. We found that tactile foraging red knots mainly caught hard-shelled prey that are buried in the sediment, whereas visual foraging knots only captured soft preys located close to or on the surface. We also found that faster explorers showed a higher percentage of visual foraging than slower explorers. By contrast, morphology (bill length and gizzard size) had no significant effect on foraging tactics. Diet analysis based on δ 15 N and δ 13 C stable isotope values of plasma and red blood cells confirmed our field observations with slower explorers mainly consumed hard-shelled prey while faster explorers consumed more soft than hard-shelled prey. Our results show that foraging tactics and diet are associated with a personality trait, independent of morphological differences. We discuss how consistent behaviour might develop early in life through positive feedbacks between foraging tactics, prey type and foraging efficiency., (© 2021 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Mass regulation in birds is well documented. For example, birds can increase body mass in response to lower availability and/or predictability of food and decrease body mass in response to increased predation danger. Birds also demonstrate an ability to maintain body mass across a range of food qualities. Although the adaptive significance of mass regulation has received a great deal of theoretical and empirical attention, the mechanisms by which birds achieve this have not. Several non-exclusive mechanisms could facilitate mass regulation in birds. Birds could regulate body mass by adjusting food intake (dieting), activity, baseline energetic requirements (basal metabolic rate), mitochondrial efficiency or assimilation efficiency. Here, we present the results of two experiments in captive red knots ( Calidris canutus islandica ) that assess three of these proposed mechanisms: dieting, activity and up- and down-regulation of metabolic rate. In the first experiment, knots were exposed to cues of predation risk that led them to exhibit presumably adaptive mass loss. In the second experiment, knots maintained constant body mass despite being fed alternating high- and low-quality diets. In both experiments, regulation of body mass was achieved through a combination of changes in food intake and activity. Both experiments also provide some evidence for a role of metabolic adjustments. Taken together, these two experiments demonstrate that fine-scale management of body mass in knots is achieved through multiple mechanisms acting simultaneously., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Age-related increases in the repeatable expression of labile phenotypic traits are often assumed to arise from an increase in among-individual variance due to differences in developmental plasticity or by means of state-behavior feedbacks. However, age-related increases in repeatability could also arise from a decrease in within-individual variance as a result of stabilizing trait expression, that is, canalization. Here we describe age-related changes in within-individual and among-individual variance components in two correlated traits-gizzard mass and exploration behavior-in a medium-sized shorebird, the red knot ( Calidris canutus ). Increased repeatability of gizzard mass came about due to an increase in among-individual variance, unrelated to differences in developmental plasticity, together with decreases in within-individual variance consistent with canalization. We also found canalization of exploration but no age-related increase in overall repeatability, which suggests that showing predictable expression of exploration behavior may be advantageous from a very young age onward. Contrasts between juveniles and adults in the first year after their capture provide support for the idea that environmental conditions play a key role in generating among-individual variation in both gizzard mass and exploration behavior. Our study shows that stabilization of traits occurs under constant conditions: with increased exposure to predictable cues, individuals may become more certain in their assessment of the environment allowing traits to become canalized.

Migratory birds undergo impressive body remodelling over the course of an annual cycle. Prior to long-distance flights, red knots ( Calidris canutus islandica) reduce gizzard mass while increasing body mass and pectoral muscle mass. Although body mass and pectoral muscle mass are functionally linked via their joint effects on flight performance, gizzard and pectoral muscle mass are thought to be independently regulated. Current hypotheses for observed negative within-individual covariation between gizzard and pectoral muscle mass in free-living knots are based on a common factor (e.g. migration) simultaneously affecting both traits, and/or protein limitation forcing allocation decisions. We used diet manipulations to generate within-individual variation in gizzard mass and test for independence between gizzard and pectoral muscle mass within individuals outside the period of migration and under conditions of high protein availability. Contrary to our prediction, we observed a negative within-individual covariation between gizzard and pectoral muscle mass. We discuss this result as a potential outcome of an evolved mechanism underlying body remodelling associated with migration. Although our proposed mechanism requires empirical testing, this study echoes earlier calls for greater integration of studies of function and mechanism, and in particular, the need for more explicit consideration of the evolution of mechanisms underlying phenotypic design.

[This corrects the article DOI: 10.1186/s40462-018-0142-4.].

Background: Space use strategies by foraging animals are often considered to be species-specific. However, similarity between conspecific strategies may also result from similar resource environments. Here, we revisit classic predictions of the relationships between the resource distribution and foragers' space use by tracking free-living foragers of a single species in two contrasting resource landscapes. At two main non-breeding areas along the East-Atlantic flyway (Wadden Sea, The Netherlands and Banc d'Arguin, Mauritania), we mapped prey distributions and derived resource landscapes in terms of the predicted intake rate of red knots ( Calidris canutus ), migratory molluscivore shorebirds. We tracked the foraging paths of 13 and 38 individual red knots at intervals of 1 s over two and five weeks in the Wadden Sea and at Banc d'Arguin, respectively. Mediated by competition for resources, we expected aggregation to be strong and site fidelity weak in an environment with large resource patches. The opposite was expected for small resource patches, but only if local resource abundances were high., Results: Compared with Banc d'Arguin , resource patches in the Wadden Sea were larger and the maximum local resource abundance was higher. However, because of constraints set by digestive capacity, the average potential intake rates by red knots were similar at the two study sites. Space-use patterns differed as predicted from these differences in resource landscapes. Whereas foraging red knots in the Wadden Sea roamed the mudflats in high aggregation without site fidelity (i.e. grouping nomads ), at Banc d'Arguin they showed less aggregation but were strongly site-faithful (i.e. solitary residents) ., Conclusion: The space use pattern of red knots in the two study areas showed diametrically opposite patterns. These differences could be explained from the distribution of resources in the two areas. Our findings imply that intraspecific similarities in space use patterns represent responses to similar resource environments rather than species-specificity. To predict how environmental change affects space use, we need to understand the degree to which space-use strategies result from developmental plasticity and behavioural flexibility. This requires not only tracking foragers throughout their development, but also tracking their environment in sufficient spatial and temporal detail., Competing Interests: All research, including animal experiments, was carried out according to Dutch law (DEC license NIOZ 10.04).Not applicable.The authors declare that they have no competing interests.Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

In the original HTML version of this Article, the order of authors within the author list was incorrect. The consortium VRS Castricum was incorrectly listed after Theunis Piersma and should have been listed after Cornelis J. Camphuysen. This error has been corrected in the HTML version of the Article; the PDF version was correct at the time of publication.

Under climate warming, migratory birds should align reproduction dates with advancing plant and arthropod phenology. To arrive on the breeding grounds earlier, migrants may speed up spring migration by curtailing the time spent en route, possibly at the cost of decreased survival rates. Based on a decades-long series of observations along an entire flyway, we show that when refuelling time is limited, variation in food abundance in the spring staging area affects fitness. Bar-tailed godwits migrating from West Africa to the Siberian Arctic reduce refuelling time at their European staging site and thus maintain a close match between breeding and tundra phenology. Annual survival probability decreases with shorter refuelling times, but correlates positively with refuelling rate, which in turn is correlated with food abundance in the staging area. This chain of effects implies that conditions in the temperate zone determine the ability of godwits to cope with climate-related changes in the Arctic.

Many monitoring programmes of species abundance and biomass increasingly face financial pressures. Occupancy is often easier and cheaper to measure than abundance or biomass. We, therefore, explored whether measuring occupancy is a viable alternative to measuring abundance and biomass. Abundance- or biomass-occupancy relationships were studied for sixteen macrozoobenthos species collected across the entire Dutch Wadden Sea in eight consecutive summers. Because the form and strength of these relationships are scale-dependent, the analysis was completed at different spatiotemporal scales. Large differences in intercept and slope of abundance- or biomass-occupancy relationships were found. Abundance, not biomass, was generally positively correlated with occupancy. Only at the largest scale, seven species showed reasonably strong abundance-occupancy relationships with large coefficients of determination and small differences in observed and predicted values (RMSE). Otherwise, and at all the other scales, intraspecific abundance and biomass relationships were poor. Our results showed that there is no generic relationship between a species' abundance or biomass and its occupancy. We discuss how ecological differences between species could cause such large variation in these relationships. Future technologies might allow estimating a species' abundance or biomass directly from eDNA sampling data, but for now, we need to rely on traditional sampling technology.

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., (© 2016 The Author(s).)

Behavioural variation within a species is usually explained as the consequence of individual variation in physiology. However, new evidence suggests that the arrow of causality may well be in the reverse direction: behaviours such as diet preferences cause the differences in physiological and morphological traits. Recently, diet preferences were proposed to underlie consistent differences in digestive organ mass and movement patterns (patch residence times) in red knots (Calidris canutus islandica). Red knots are molluscivorous and migrant shorebirds for which the size of the muscular stomach (gizzard) is critical for the food processing rate. In this study, red knots (C. c. canutus, n = 46) were caught at Banc d'Arguin, an intertidal flat ecosystem in Mauritania, and released with radio-tags after the measurement of gizzard mass. Using a novel tracking system (time-of-arrival), patch residence times were measured over a period of three weeks. Whether or not gizzard mass determined patch residence times was tested experimentally by offering 12 of the 46 tagged red knots soft diets prior to release; this reduced an individual's gizzard mass by 20-60%. To validate whether the observed range of patch residence times would be expected from individual diet preferences, we simulated patch residence times as a function of diet preferences via a simple departure rule. Consistent with previous empirical studies, patch residence times in the field were positively correlated with gizzard mass. The slope of this correlation, as well as the observed range of patch residence times, was in accordance with the simulated values. The 12 birds with reduced gizzard masses did not decrease patch residence times in response to the reduction in gizzard mass. These findings suggest that diet preferences can indeed cause the observed among-individual variation in gizzard mass and patch residence times. We discuss how early diet experiences can have cascading effects on the individual expression of both behavioural and physiomorphic traits. This emphasizes that to understand the ecological consequences of individual differences, knowledge of the environment during development is required., (© 2016 The Authors. Journal of Animal Ecology © 2016 British Ecological Society.)

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., (© 2016 The Author(s).)

Digestive capacity often limits food intake rate in animals. Many species can flexibly adjust digestive organ mass, enabling them to increase intake rate in times of increased energy requirement and/or scarcity of high-quality prey. However, some prey species are defended by secondary compounds, thereby forcing a toxin limitation on the forager's intake rate, a constraint that potentially cannot be alleviated by enlarging digestive capacity. Hence, physiological flexibility may have a differential effect on intake of different prey types, and consequently on dietary preferences. We tested this effect in red knots (Calidris canutus canutus), medium-sized migratory shorebirds that feed on hard-shelled, usually mollusc, prey. Because they ingest their prey whole and crush the shell in their gizzard, the intake rate of red knots is generally constrained by digestive capacity. However, one of their main prey, the bivalve Loripes lucinalis, imposes a toxin constraint due to its symbiosis with sulphide-oxidizing bacteria. We manipulated gizzard sizes of red knots through prolonged exposure to hard-shelled or soft foods. We then measured maximum intake rates of toxic Loripes versus a non-toxic bivalve, Dosinia isocardia. We found that intake of Dosinia exponentially increased with gizzard mass, confirming earlier results with non-toxic prey, whereas intake of Loripes was independent of gizzard mass. Using linear programming, we show that this leads to markedly different expected diet preferences in red knots that try to maximize energy intake rate with a small versus a large gizzard. Intra- and inter-individual variation in digestive capacity is found in many animal species. Hence, the here proposed functional link with individual differences in foraging decisions may be general. We emphasize the potential relevance of individual variation in physiology when studying trophic interactions.

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.

At what phenotypic level do closely related subspecies that live in different environments differ with respect to food detection, ingestion and processing? This question motivated an experimental study on rock sandpipers (Calidris ptilocnemis). The species' nonbreeding range spans 20 deg of latitude, the extremes of which are inhabited by two subspecies: C. p. ptilocnemis that winters primarily in upper Cook Inlet, Alaska (61°N) and C. p. tschuktschorum that overlaps slightly with C. p. ptilocnemis but whose range extends much farther south (∼40°N). In view of the strongly contrasting energetic demands of their distinct nonbreeding distributions, we conducted experiments to assess the behavioral, physiological and sensory aspects of foraging and we used the bivalve Macoma balthica for all trials. C. p. ptilocnemis consumed a wider range of prey sizes, had higher maximum rates of energy intake, processed shell waste at higher maximum rates and handled prey more quickly. Notably, however, the two subspecies did not differ in their abilities to find buried prey. The subspecies were similar in size and had equally sized gizzards, but the more northern ptilocnemis individuals were 10-14% heavier than their same-sex tschuktschorum counterparts. The higher body mass in ptilocnemis probably resulted from hypertrophy of digestive organs (e.g. intestine, liver) related to digestion and nutrient assimilation. Given the previously established equality of the metabolic capacities of the two subspecies, we propose that the high-latitude nonbreeding range of ptilocnemis rock sandpipers is primarily facilitated by digestive (i.e. physiological) aspects of their foraging ecology rather than behavioral or sensory aspects., (© 2015. Published by The Company of Biologists Ltd.)

The evolutionary function and maintenance of variation in animal personality is still under debate. Variation in the size of metabolic organs has recently been suggested to cause and maintain variation in personality. Here, we examine two main underlying notions: (i) that organ sizes vary consistently between individuals and cause consistent behavioural patterns, and (ii) that a more exploratory personality is associated with reduced survival. Exploratory behaviour of captive red knots (Calidris canutus, a migrant shorebird) was negatively rather than positively correlated with digestive organ (gizzard) mass, as well as with body mass. In an experiment, we reciprocally reduced and increased individual gizzard masses and found that exploration scores were unaffected. Whether or not these birds were resighted locally over the 19 months after release was negatively correlated with their exploration scores. Moreover, a long-term mark-recapture effort on free-living red knots with known gizzard masses at capture confirmed that local resighting probability (an inverse measure of exploratory behaviour) was correlated with gizzard mass without detrimental effects on survival. We conclude that personality drives physiological adjustments, rather than the other way around, and suggest that physiological adjustments mitigate the survival costs of exploratory behaviour. Our results show that we need to reconsider hypotheses explaining personality variation based on organ sizes and differential survival.

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.

When a diet switch results in a change in dietary isotopic values, isotope ratios of the consumer's tissues will change until a new equilibrium is reached. This change is generally best described by an exponential decay curve. Indeed, after a diet switch in captive red knot shorebirds (Calidris canutus islandica), the depletion of (13)C in both blood cells and plasma followed an exponential decay curve. Surprisingly, the diet switch with a dietary (15)N/(14)N ratio (δ(15)N) change from 11.4 to 8.8 ‰ had little effect on δ(15)N in the same tissues. The diet-plasma and diet-cellular discrimination factors of (15)N with the initial diet were very low (0.5 and 0.2 ‰, respectively). δ(15)N in blood cells and plasma decreased linearly with increasing body mass, explaining about 40 % of the variation in δ(15)N. δ(15)N in plasma also decreased with increasing body-mass change (r (2)=.07). This suggests that the unusual variation in δ(15)N with time after the diet switch was due to interferences with simultaneous changes in body-protein turnover.

Timing "in the real world" must cope with the temporal complexity of natural environments. Extreme examples for the resultant "multitasking" are migratory birds, which precisely time movements to remote areas. New field technologies highlight temporal accuracy, while captivity studies emphasize underlying programs and plasticity of schedules. After reviewing these findings, we focus on waders, which undertake spectacular long-distance migrations, have robust circannual clocks, and cope with diel, tidal, and polar environments. To explore features that may facilitate such multitasking, we speculated that melatonin amplitudes are low and damped during seasons when entrainment to subtle Zeitgebers occurs. We measured melatonin profiles under European daylength in two species with different ecologies and found low-amplitude melatonin cycles that changed over the year. Annual patterns neither fully supported our hypothesis, nor simply reflected daylight availability. While migratory birds are inspiring models for chronobiology, mechanistic understanding of their multitasking is still poor., (Copyright © 2012 Elsevier B.V. All rights reserved.)

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., (This journal is © 2011 The Royal Society)

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 (M(sum)), 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, M(sum) 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.

1. Studies of diet choice usually assume maximization of energy intake. The well-known 'contingency model' (CM) additionally assumes that foraging animals only spend time searching or handling prey. Despite considerable empirical support, there are many foraging contexts in which the CM fails, but such cases were considered exceptions rather than the rule. 2. For animals constrained by the rate at which food is digested, CM does not necessarily lead to maximal energy intake rates because the time for digestion is not part of the selection criteria. In the main model developed to explain diet choice under a digestive constraint, the 'digestive rate model' (DRM), time lost to digestive breaks is minimized so that energy intake over total time (searching, handling, digestive breaks) is maximized. 3. It is increasingly acknowledged that most animals may face digestive constraints as prey capture rates vary over time and as it would be a waste to carry around heavy digestive machinery that can rapidly process food under all circumstances: this is only needed in times of high demand, provided that enough food can be found. 4. In molluscivore shorebirds ingesting hard-shelled prey such as red knots (Calidris canutus), the predictions of DRM were held up so far, whereas those of CM were rejected. However, most tests were carried out under controlled experimental conditions. Red knots overwinter in coastal areas over much of Western Europe and we capitalized on this variation by comparing, during a single winter, observed diet composition with predictions of DRM, CM and a null model assuming no prey selection ('no-selection model', NSM). 5. The observed diets were best predicted by DRM followed by CM. NSM poorly predicted observed diet choice. Under the present conditions, diet choice based on DRM would on average have yielded an energy intake rate twice as large as one based on CM. By adjusting the size of their gizzard (held constant in the present simulations), red knots could have lifted their energy intake rate further. We suggest that application of the DRM can help many diet studies forward, especially those previously seen as exceptions to the classical CM-based rule.

1. Whether intertidal areas are used to capacity by shorebirds can best be answered by large-scale manipulation of foraging areas. The recent overexploitation of benthic resources in the western Dutch Wadden Sea offers such an 'experimental' setting. 2. We review the effects of declining food abundances on red knot Calidris canutus islandica numbers, based on a yearly large-scale benthic mapping effort, long-term colour-ringing and regular bird-counts from 1996 to 2005. We focus on the three-way relationships between suitable foraging area, the spatial predictability of food and red knot survival. 3. For each benthic sampling position, red knot intake rate (mg AFDM s(-1)) was predicted by a multiple prey species functional response model, based on digestive rate maximization (this model explained diet and intake rate in earlier studies on red knots). This enabled us to derive the spatial distribution of the suitable foraging area, which in each of the 10 years was analysed with a measure of autocorrelation, i.e. Moran's I. 4. Over the 10 years, when accounting for a threshold value to meet energetic demands, red knots lost 55% of their suitable foraging area. This ran parallel to a decrease in red knot numbers by 42%. Although there was also a decrease in patchiness (i.e. less information about the location of the suitable feeding sites), this did not yet lead to additional loss of birds. 5. To cope with these landscape-scale declines in food stocks, an increase in the capacity for instantaneous food processing would be required. Although we show that red knots indeed enlarged their muscular gizzards, the increase in gizzard size was not enough to compensate for the decreased feeding area. 6. Survival of islandica knots in the western Dutch Wadden Sea, based on colour-ring resightings, declined from 89% in the first half of our study period to 82% in the second half of our study period and could account for almost half of the decline in red knot numbers; the rest must have moved elsewhere in winter. 7. Densities of red knots per unit suitable foraging area remained constant at 10 knots ha(-1) between 1996 and 2005, which suggests that red knots have been using the Dutch Wadden Sea to full capacity.

Previous experiments showed reduction of basal metabolic rate (BMR) in birds facing energetic challenges. We alternately exposed two groups of red knots (Calidris canutus) to either 6 h or 22 h of food availability for periods of 22 d. Six h of access to food led to a 6%-10% loss of body mass over the first 8 d, with nearly all of the birds' daily energy expenditures supported by body nutrient stores during the first 2 d. Birds responded by increasing feeding behavior and food intake, but the response was slow. There were no gains of mass before day 15, which suggests a digestive bottleneck and a period of physiological adjustment. Food-restricted birds exhibited decreases in pectoral-muscle thickness and BMR in association with a loss of body mass. Although a decrease in BMR saves energy, savings represented only 2%-7% of the daily energy spent in excess of that acquired during the deficit period. Red knots did not downregulate mass-independent BMR. On the bases of recent independent findings and the pattern of mass gain observed when food access was switched from 6 h to 22 h, we suggest that these birds routinely maintain nutrient stores as a buffer against periods of energy shortages, thereby precluding the need for downregulation of mass-independent BMR.

The winter immunoenhancement hypothesis associates long nights and increased exposure to melatonin with enhanced immune function in winter when resource availability is low and the chances of becoming ill are high. Thus, increased exposure to melatonin in the winter could be adaptive for species facing difficult winter conditions. This idea has found some support in studies of resident mammals. In birds, the link between day length and melatonin over the annual cycle is weaker, and contributions of melatonin to seasonal timing are unclear. Furthermore, many species, especially migrants, do not experience the most difficult conditions of their annual cycle in winter. In this study, we tested whether the winter immunoenhancement hypothesis holds in an avian species, the red knot Calidris canutus. We found that melatonin duration and amplitude varied significantly over the annual cycle with the highest values occurring in winter. However, peaks did not correspond to the winter solstice or with annual variation in immune function. Our findings do not support the winter immunoenhancement hypothesis in knots and question whether the idea that immune function should be bolstered in winter can be generalized to systems where winter is not the most difficult time of the year.

Optimality reasoning from behavioural ecology can be used as a tool to infer how animals perceive their environment. Using optimality principles in a 'reversed manner' may enable ecologists to predict changes in population size before such changes actually happen. Here we show that a behavioural anti-predation trait (burrowing depth) of the marine bivalve Macoma balthica can be used as an indicator of the change in population size over the year to come. The per capita population growth rate between years t and t+1 correlated strongly with the proportion of individuals living in the dangerous top 4 cm layer of the sediment in year t: the more individuals in the top layer, the steeper the population decline. This is consistent with the prediction based on optimal foraging theory that animals with poor prospects should accept greater risks of predation. This study is among the first to document fitness forecasting in animals.

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 degrees C, 5 degrees 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 (M(sum))] 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 M(sum). 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.

There is a widespread concern about the direct and indirect effects of industrial fisheries; this concern is particularly pertinent for so-called "marine protected areas" (MPAs), which should be safeguarded by national and international law. The intertidal flats of the Dutch Wadden Sea are a State Nature Monument and are protected under the Ramsar convention and the European Union's Habitat and Birds Directives. Until 2004, the Dutch government granted permission for ~75% of the intertidal flats to be exploited by mechanical dredgers for edible cockles (Cerastoderma edule). Here we show that dredged areas belonged to the limited area of intertidal flats that were of sufficient quality for red knots (Calidris canutus islandica), a long-distance migrant molluscivore specialist, to feed. Dredging led to relatively lower settlement rates of cockles and also reduced their quality (ratio of flesh to shell). From 1998 to 2002, red knots increased gizzard mass to compensate for a gradual loss in shellfish quality, but this compensation was not sufficient and led to decreases in local survival. Therefore, the gradual destruction of the necessary intertidal resources explains both the loss of red knots from the Dutch Wadden Sea and the decline of the European wintering population. This study shows that MPAs that do not provide adequate protection from fishing may fail in their conservation objectives., Competing Interests: Competing interests. The authors have declared that no competing interests exist.

Seasonal acclimatization and experimental acclimation to cold in birds typically results from increased shivering endurance and elevated thermogenic capacity leading to improved resistance to cold. A wide array of physiological adjustments, ranging from biochemical transformations to organ mass variations, are involved in this process. Several studies have shown that improved cold endurance is accompanied by increases in summit metabolic rate (M(sum)), a measure of maximal heat production and an indicator of the level of sustainable thermogenic capacity. However, improved endurance to cold can also be achieved without significant changes in M(sum). The same is true for basal metabolic rate (BMR), which is known to increase in association with cold acclimatization or acclimation in some species but not in others. We investigated cold acclimation in a migrant shorebird known for extreme physiological flexibility, the red knot (Calidris canutus, the northerly wintering subspecies islandica). We measured BMR and M(sum) over two months in birds caught in the wild and transferred to experimentally controlled conditions representative of aspects of their seasonal thermal environment (two groups at constant 25 degrees C, one group at constant 4 degrees C and two groups experiencing variable outdoor temperatures). Birds maintained in both cold and variable ambient temperatures showed a 14-15% higher body mass, 33-45% higher food intake, and 26% and 13% elevations in BMR and M(sum), respectively, compared with birds kept at thermoneutrality. These results, together with data on alimentary tract size and pectoral muscle thickness measured by ultrasonography, suggest that red knots acclimate to cold primarily through modulation of (lean) body mass components. Heavier individuals have larger muscles, which allow higher maximal heat production and better thermal compensation. Cold acclimation effects on BMR are most probably due to changes in the size of visceral organs, although not the alimentary tract in this specific case. The liver, known for its thermogenic capacity, is a probable candidate. Overall, our results indicate that relatively small changes in body mass and muscle size allow enough reserve capacity in terms of heat production to cope with typical wintering ambient temperature variations as measured on the red knot's wintering grounds.

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.

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 2H, (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 2H and (18)O only) and a stable isotope ratio infrared laser spectrometry (2H, (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.

Aiming to interpret functionally the large variation in gizzard masses of red knots Calidris canutus, we experimentally studied how the digestive processing rate is influenced by the size of the gizzard. During their non-breeding season, red knots feed on hard-shelled molluscs, which they ingest whole and crush in their gizzard. In three experiments with captive birds we tested predictions of the hypothesis that gizzard size, via the rate of shell crushing and processing, constrains intake rate in red knots (against the alternative idea that external handling times constrain intake rate). Gizzard size within individual birds was manipulated by varying the hardness of the diet on offer, and was confirmed by ultrasonography. The results upheld the "shell-crushing hypothesis" and rejected the "handling time hypothesis". Intake rates on with-shell prey increased with gizzard size, and decreased with shell mass per prey. Intake rates on soft (without shell) prey were higher than on with-shell prey and were unaffected by gizzard size. Offering prey that were heavily shelled relative to their flesh mass led to energy intake rates that were marginally sufficient to balance the daily energy budget within the time that is naturally available in a tidal system. We predicted the optimal gizzard sizes that are required to either (1) balance energy income with energy expenditure, or (2) to maximise net daily energy intake. The gizzard mass of free-living red knots in the Wadden Sea is such that it maximises daily net energy intake in spring when fuelling for migration, while it balances energy budget throughout the remainder of the year.

Although the energy costs of foraging and food processing in vertebrates may be considerable, they have rarely been quantified separately. Here we present estimates for both cost factors based on a series of trials with a shorebird, the red knot Calidris canutus, fed natural and artificial prey types under naturalistic but fully controlled indoor aviary conditions. During eight 1-day trials we successfully manipulated the extent to which the five red knots were (1) actively probing and walking (i.e. foraging) and (2) actually ingesting prey (i.e. processing food) that was (3) either hard-shelled or not (i.e. crushing). Energy expenditures, estimated by the doubly labelled water (DLW) method, calibrated for use in this particular condition, varied between 1.5 and 4 W. A hierarchical analysis of variance indicated that the crushing of hard-shelled prey entailed no extra cost. We arrived at the following breakdown of cost components under the thermoneutral conditions of the experiment: a cost of active rest/maintenance of 1.665 W, an additional cost of foraging of 0.602 W and an additional digestive processing cost of 1.082 W. These cost levels are all well within the range of expectation and are consistent with the results of a separate outdoor aviary experiment in which the thermostatic costs needed separate estimation. On the basis of the cost and performance functions of gizzards of different mass, it was shown that under the conditions of this experiment the red knots expended the bare minimum for a balanced budget, maintaining the smallest possible gizzard. Under field conditions a larger gizzard would be required.