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The ecological reasons for variation in avian migration, with some populations migrating across thousands of kilometres between breeding and non-breeding areas with one or few refuelling stops, in contrast to others that stop more often, remain to be pinned down. Red Knots Calidris canutus are a textbook example of a shorebird species that makes long migrations with only a few stops. Recognizing that such behaviours are not necessarily species-specific but determined by ecological context, we here provide a description of the migrations of a relatively recently described subspecies (piersmai). Based on data from tagging of Red Knots on the terminal non-breeding grounds in northwest Australia with 4.5- and 2.5-g solar-powered Platform Terminal Transmitters (PTTs) and 1.0-g geolocators, we obtained information on 19 route-records of 17 individuals, resulting in seven complete return migrations. We confirm published evidence that Red Knots of the piersmai subspecies migrate from NW Australia and breed on the New Siberian Islands in the Russian Arctic and that they stage along the coasts of southeastern Asia, especially in the northern Yellow Sea in China. Red Knots arrived on the tundra breeding grounds from 8 June onwards. Southward departures mainly occurred in the last week of July and the first week of August. We documented six non-stop flights of over c. 5000 km (with a maximum of 6500 km, lasting 6.6 days). Nevertheless, rather than staging at a single location for multiple weeks halfway during migration, piersmai-knots made several stops of up to a week. This was especially evident during northward migration, when birds often stopped along the way in southeast Asia and 'hugged' the coast of China, thus flying an additional 1000-1500 km compared with the shortest possible (great circle route) flights between NW Australia and the Yellow Sea. The birds staged longest in areas in northern China, along the shores of Bohai Bay and upper Liaodong Bay, where the bivalve Potamocorbula laevis, known as a particularly suitable food for Red Knots, was present. The use of multiple food-rich stopping sites during northward migration by piersmai is atypical among subspecies of Red Knots. Although piersmai apparently has the benefit of multiple suitable stopping areas along the flyway, it is a subspecies in decline and their mortality away from the NW Australian non-breeding grounds has been elevated.

Migratory shorebirds show highly organized seasonal cycles in physiological and morphological traits (body mass and composition, plumage, hormone levels, etc.), which in captivity is accompanied by restless behaviour at times when free‐living birds would start migration. We introduce the idea that seasonally changing preference for habitat could motivate migrants to embark on migration and that this cognitive process could also guide them to seasonally appropriate places. We explored this by testing whether red knots (Calidris canutus), which also in captivity maintain marked circannual phenotypic rhythms, show evidence of seasonal change in preference for pictures of seasonally appropriate habitats. We first developed a method to verify whether red knots are able to memorize and discriminate contrasting pictures projected by LCD projectors. This was followed by two different experiments in which we tested for a seasonally changing preference for breeding or non‐breeding habitat. When carried out during the pre‐breeding season, the red knots are expected to prefer pictures of mudflats, their non‐breeding habitat. At the start of the breeding season, they should prefer pictures of the tundra breeding habitat. We established that knots are able to distinguish and memorize projected images. We failed to demonstrate the predicted change in vision‐based habitat preference, but for reasons of test design we do not interpret this as a strong rejection of the hypothesis. Instead, we suggest that experiments with greater numbers of individuals tested once, perhaps in combination with the provision of additional cues such as smells and sounds, will help the development of these ideas further.

Mass regulation is 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, or BMR), 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/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 on 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.

Despite the wealth of studies on seasonal movements of birds between southern nonbreeding locations and High Arctic breeding locations, the key mechanisms of navigation during these migrations remain elusive. A flight along the shortest possible route between pairs of points on a sphere (‘orthodrome’) requires a bird to be able to assess its current location in relation to its migration goal and to make continuous adjustment of heading to reach that goal. Alternatively, birds may navigate along a vector with a fixed orientation (‘loxodrome’) based on magnetic and/or celestial compass mechanisms. Compass navigation is considered especially challenging for summer migrations in Polar regions, as continuous daylight and complexity in the geomagnetic field may complicate the use of both celestial and magnetic compasses here. We examine the possible use of orientation mechanisms during migratory flights across the Greenland Icecap. Using a novel 2 g solar‐powered satellite transmitter, we documented the flight paths travelled by a female red knot Calidris canutus islandica during two northward and two southward migrations. The geometry of the paths suggests that red knots can migrate across the Greenland Icecap along the shortest‐, orthodrome‐like, path instead of the previously suggested loxodrome path. This particular bird's ability to return to locations visited in a previous year, together with its sudden course changes (which would be appropriate responses to ambient wind fields), suggest a map sense that enables red knots to determine location, so that they can tailor their route depending on local conditions.

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 (

Red knots (Calidris canutus) are shorebirds that spent most of the year at intertidal areas such as the Wadden Sea. However, once a year to and from their northern breeding grounds. Red knots yearly undergo a remodeling of their body to be physiologically prepared for the long flight and following reproductive season. Previous work has shown that many experienced migrants are very consistent in their migratory routines.The aim of this thesis is to increase understanding of the development of individual migratory routines. In the first part of this thesis, I first explore individual variation in migratory behaviour present in wild knots using a novel solar-powered satellite transmitter. In the second part of this thesis I investigate how differences in experience effect the development of physiological and behavioural traits in birds temporarily held in captivity. By combining these results I show that environmental conditions play a key role in shaping individual migratory routines. Expending on the role of experience in shaping individual differences I present a testing protocol aimed at testing cognitive aspects of migration, a fairly under represented topic in studies on migration. For future studies I suggest to combine tracking of individual birds with standardized assays to stimulate research into the cognitive aspects of migratory behaviour. Because I believe that the closer we are to understanding ‘what it is like to be a bird (or any other animal) the more we are able, and willing, to appreciate and protect the animals and the habitats that are around us.

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.

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.