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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. [ABSTRACT FROM AUTHOR]

Rock Sandpipers Calidris ptilocnemis have the most northerly non‐breeding distribution of any shorebird in the Pacific Basin (upper Cook Inlet, Alaska; 61°N, 151°W). In terms of freezing temperatures, persistent winds and pervasive ice, this site is the harshest used by shorebirds during winter. We integrated physiological, metabolic, behavioural and environmental aspects of the non‐breeding ecology of Rock Sandpipers at the northern extent of their range to determine the relative importance of these factors in facilitating their unique non‐breeding ecology. Not surprisingly, estimated daily energetic demands were greatest during January, the coldest period of winter. These estimates were greatest for foraging birds, and exceeded basal metabolic rates by a factor of 6.5, a scope of increase that approaches the maximum sustained rate of energetic output by shorebirds during periods of migration, but far exceeds these periods in duration. We assessed the quality of their primary prey, the bivalve Macoma balthica, to determine the daily foraging duration required by Rock Sandpipers to satisfy such energetic demands. Based on size‐specific estimates of M. balthica quality, Rock Sandpipers require over 13 h/day of foraging time in upper Cook Inlet in January, even when feeding on the highest quality prey. This range approaches the average daily duration of mudflat availability in this region (c. 18 h), a maximum value that annually decreases due to the accumulation of shore‐fast ice. Rock Sandpipers are likely to maximize access to foraging sites by following the exposure of ice‐free mudflats across the upper Cook Inlet region and by selecting smaller, higher quality M. balthica to minimize foraging times. Ultimately, this unusual non‐breeding ecology relies on the high quality of their prey resources. Compared with other sites across their range, M. balthica from upper Cook Inlet have relatively light shells, potentially the result of the region's depauperate invertebrate predator community. Given the delicate balance between environmental and prey conditions that currently make Cook Inlet a viable wintering area for Rock Sandpipers, small variations in these variables may affect the suitability of the site in the future. [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]

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. [ABSTRACT FROM AUTHOR]

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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]

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 baithica 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. [ABSTRACT FROM AUTHOR]

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 of13C in both blood cells and plasma followed an exponential decay curve. Surprisingly, the diet switch with a dietary15N/14N ratio (δ15N) change from 11.4 to 8.8 ‰ had little effect on δ15N in the same tissues. The diet-plasma and diet-cellular discrimination factors of15N with the initial diet were very low (0.5 and 0.2 ‰, respectively). δ15N in blood cells and plasma decreased linearly with increasing body mass, explaining about 40 % of the variation in δ15N. δ15N in plasma also decreased with increasing body-mass change (r2=.07). This suggests that the unusual variation in δ15N with time after the diet switch was due to interferences with simultaneous changes in body-protein turnover. [ABSTRACT FROM PUBLISHER]

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]

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]

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. [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]

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 (Msum), 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 Msum. 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 Msum 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°C, one group at constant 4°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 Msum, 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. [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]

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 1day 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. [ABSTRACT FROM AUTHOR]

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 nonbreeding 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 dally net energy intake in spring when fuelling for migration, while it balances energy budget throughout the remainder of the year. [ABSTRACT FROM AUTHOR]

Tests the hypothesis that ingested water equilibrates isotopically with the body pool of a shorebird with unrivaled water fluxes. Changes in body mass; Relationship between average body mass and average amount of body water; Percentage of time the birds spent foraging, resting, and flying.

Organs, even of fully grown adult birds, mammals, and reptiles, may show substantial size changes in relation to specific performances. These changes are difficult to study, because measurements usually can only be obtained following the death of the animal. We explored the use of ultrasonographic imaging, a relatively simple noninvasive technique, to measure size of pectoral muscles and stomach in two small shorebird species (red knots Calidris canutus and golden plovers Pluvialis apricaria). Accuracy of ultrasound measurements in estimating organ mass in red knots was reasonably high. Depending on the equipment used, the error of individual measurements was 20%-25% for the pectoral muscles and 26%-44% for the stomach. In plovers the technique was less accurate, probably because of the low variability of the organs involved. Ultrasound scanning is particularly suited to measure rapidly changing organ sizes over short time intervals. We demonstrate this with an example in which changes in individuals in size of pectoral muscle and stomach were monitored in captive red knots following a change in diet. Ultrasound measures will enable studies on the links between body composition and future behavior and physiology. [ABSTRACT FROM AUTHOR]

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]