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Tracking animal movement is important for understanding how animals interact with their (changing) environment, and crucial for predicting and explaining how animals are affected by anthropogenic activities. The Wadden Sea is a UNESCO World Heritage Site and a region of global importance for millions of shorebirds. Due to climate change and anthropogenic activity, understanding and predicting movement and space-use in areas like the Wadden Sea is increasingly important. Monitoring and predicting animal movement, however, requires high-resolution tracking of many individuals. While high-resolution tracking has been made possible through GPS, trade-offs between tag weight and battery life limit its use to larger species. Here, we introduce WATLAS (the Wadden Sea deployment of the ATLAS tracking system) capable of monitoring the movements of hundreds of (small) birds simultaneously in the Dutch Wadden Sea. WATLAS employs an array of receiver stations that can detect and localize small, low-cost tags at fine spatial (metres) and temporal resolution (seconds). From 2017 to 2021, we tracked red knots, sanderlings, bar-tailed godwits, and common terns. We use parts of these data to give four use-cases revealing its performance and demonstrating how WATLAS can be used to study numerous aspects of animal behaviour, such as, space-use (both intra- and inter-specific), among-individual variation, and social networks across levels of organization: from individuals, to species, to populations, and even communities. After describing the WATLAS system, we first illustrate space-use of red knots across the study area and how the tidal environment affects their movement. Secondly, we show large among-individual differences in distances travelled per day, and thirdly illustrate how high-throughput WATLAS data allows calculating a proximity-based social network. Finally, we demonstrate that using WATLAS to monitor multiple species can reveal differential space use. For example, despite sanderlings and red knots roosting together, they foraged in different areas of the mudflats. The high-resolution tracking data collected by WATLAS offers many possibilities for research into the drivers of bird movement in the Wadden Sea. WATLAS could provide a tool for impact assessment, and thus aid nature conservation and management of the globally important Wadden Sea ecosystem. [ABSTRACT FROM AUTHOR]

1. Fine-scale tracking of animal movement is important to understand the proximate mechanisms of animal behaviour. The reverse-GPS system--ATLAS--uses inexpensive (~€25), lightweight (<1 g) and low-power (~0.4 mJ/transmission) tags. Six systems are now operational worldwide and have successfully tracked over 50 species in various landscape types. The growing use of ATLAS to track animal movement motivates further refinement of best-practice application and an assessment of its accuracy. 2. Here, we test the accuracy and precision of the largest ATLAS system, located in the Dutch Wadden Sea, using concurrent GPS measurements as a reference. This large-scale ATLAS system consists of 26 receivers and covers 1,326 km2 of intertidal region, with almost no physical obstacles for radio signals, providing a useful baseline for other systems. We compared ATLAS and GPS location estimates for a route (mobile test) and 16 fixed locations (stationary test) on the Griend mudflat. Precision was estimated using standard deviation during the stationary tests. We also give examples of tracked red knots Calidris canutus islandica to illustrate the use of the system in tracking small shorebirds (~120 g). 3. ATLAS-derived location estimates differed from GPS by a median of 4.2 m (stationary test) and 5.7 m (mobile test). Signals that were collected by more receiver stations were more accurate, although even three-receiver localisations were comparable with GPS localisations (~10 m difference). Receivers that detected 90% of the 1 Hz transmissions from our test tag were within 5 km of their furthest detection but height of both receiver and tag seemed to influence detection distance. The test tag (1 Hz) had a fix rate of >90% at 15 of 16 stationary sites. Tags on birds (1/6 Hz) on the Griend mudflat had a mean fix rate of 51%, yielding an average sampling rate of 0.085 Hz. Fix rates were higher in more central parts of the receiver array. 4. ATLAS provides accurate, regional-scale tracking with which hundreds of relatively small-bodied species can be tracked simultaneously for long periods of time. Future ATLAS users should consider the height of receivers, their spatial arrangement, density and the movement modes of their study species (e.g. ground-dwelling or flying). [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]

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]

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]

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]

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. [ABSTRACT FROM AUTHOR]

The article focuses on a study which described the use of the behavioural anti-predation trait of the marine bivalve Macoma balthica as an indicator of population size change. The study utilized the burrowing depth character of Baltic tellin to forecast population change. It analyzed three components of population change, namely the decline in the risk of predation at a given depth, increased in the marginal value of energy, and the decline in Macoma balthica's expected future fitness. Study authors concluded that behavioural ecology can be used to predict the perception of animals about their environment.

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]