Many routes lead to Rome : potential causes for the multi-route migration system of Red Knots, Calidris canutus islandica

Migrants, such as birds or representatives of other taxa, usually make use of several stopover sites to cover the distance between their site of origin and destination. Potentially, multiple routes exist, but often little is known about the causes and consequences of alternative migration routes. Apart from their geographical distribution, the suitability of potential sites might play an important role in the animals’ decisions for a particular itinerary. We used an optimal-migration model to test three nonmutually exclusive hypotheses leading to variations in the spring migration routes of a subspecies of Red Knot, Calidris canutus islandica, which migrates from wintering grounds in Western Europe to breeding grounds in Greenland and the Canadian Arctic: the breeding location hypothesis, the energy budget hypothesis, and the predation risk hypothesis. Varying only breeding location, the model predicted that birds breeding in the Canadian Arctic and on West Greenland stop over on Iceland, whereas birds breeding in East and Northeast Greenland migrate via northern Norway, a prediction that is supported by empirical findings. Energy budgets on stopover sites had a strong influence on the choice of route and staging times. Varying foraging-intensity and mass-dependent predation risk prompted the birds to use less risky sites, if possible. The effect of simultaneous changes in the energy budget and predation risk strongly depended on the site where these occurred. Our findings provide potential explanations for the observations that C. canutus islandica uses a diverse array of migration routes. Scrutinizing the three alternative driving forces for the choice of migratory routes awaits further, specific data collection in rapidly developing fields of research (e.g., predation risk assessment, GPS tracking). Generally, the type of modeling presented here may not only highlight alternative explanations, but also direct follow-up empirical research.

Do Red Knots (Calidris Canutus Islandica) routinely skip Iceland during southward migration?

Subspecies Calidris canutus islandica of the Red Knot breeds on the arctic tundra of northeastern Canada and northern Greenland and winters along the coasts of northwestern Europe. During northward migration, it stops over in either Iceland or northern Norway. It has been assumed that it does the same during southward migration. Using ratios of stable carbon isotopes (δ ¹³ C) in whole blood, blood cells, and plasma, we investigated evidence for a stopover in Iceland en route from the breeding grounds to the Dutch Wadden Sea. With the expected diet (shellfish) and stopover duration at Iceland (12-15 days, maximum 17 days) and the turnover rates of blood cells (15.1 days) and plasma (6.0 days), Red Knots that stopped in Iceland should arrive with a blood (cell) δ ¹³ C midway between a tundra (-24.7[per thousand]) and a marine value (-14.0[per thousand]) and a plasma δ¹³ C approaching the marine value (-15.3[per thousand]). However, many adults arriving at the Wadden Sea had δ¹³ C ratios in blood (cells) and plasma below these levels, and some arrived with clear tundra signals in blood cells, suggesting that they skipped Iceland during southward migration. Surprisingly, available data suggest this also to be true for juveniles during their first southward migration. The δ ¹³ C signature of second-year birds confirmed that they oversummered in the Wadden Sea. Our findings contradict the largely untested idea that juvenile shorebirds make more stopovers than adults as well as the idea that the migration between the Nearctic and Europe is necessarily a two-leg process.

Northward migration of Afro-Siberian Knots Calidris canutus canutus : high variability in Red Knot numbers visiting staging sites on the French Atlantic coast, 1979–2009

The Afro–Siberian Red Knot subspecies, Calidris canutus canutus, winters mainly on Banc d’Arguin, Mauritania, West Africa. An International Wader Study Group project carried out in 1979 suggested that during northward migration Red Knots cover their migration between the wintering grounds and the Siberian breeding grounds in two long-distance non-stop flights, stopping only in the Wadden Sea in Schleswig-Holstein, Germany. Each year Red Knots also visit staging sites along the French Atlantic coast in addition to the German Wadden Sea. Ever since 1979, the French staging sites have been counted on a regular basis and here we present the count data from these 30 years. In some years more than 20% of the population used the French Atlantic coast as a staging site, but numbers are highly variable from one year to the next. We suggest that high numbers in France might occur when birds have to stop short of the Wadden Sea because of head-winds and/or a lack of tail-winds en route from West Africa.

Small home ranges and high site fidelity in red knots (Calidris c. canutus) wintering on the Banc d’Arguin, Mauritania

Using automated and manual radio-telemetry and resightings of individual colour-ringed birds, we assessed the daily use of space of red knots Calidris canutus canutus at a tropical wintering area along the Sahara coast, the Banc d Arguin in Mauritania. Confirming earlier suggestions, we found that birds were very faithful to their roosts and that the daily foraging range was small; in the course of several winter months birds used an area of only 2 16 km2 of intertidal area. We found no differences between their movements in daylight and at night. Additionally, individuals seem to return to exactly the same locations in subsequent winters. This pattern is very different from red knots wintering in the temperate Wadden Sea. Here, they readily change roost sites and easily cover areas of about 800 km2 in the course of weeks but, just as in Mauritania, no differences between day and night are apparent. In northern Patagonia and north-western Australia, red knots have range sizes closer to those on the Banc d Arguin, but here they do show differences in space use between day and night. Ecological explanations for these contrasting patterns require further comparative data based on in-depth studies on the predictability of the food base and the presence of diurnal and nocturnal predators.

Single-point isotope measurements in blood cells and plasma to estimate the time since diet switches

1. Understanding ecological phenomena often requires an accurate assessment of the timing of events. To estimate the time since a diet shift in animals without knowledge on the isotope ratios of either the old or the new diet, isotope ratio measurements in two different tissues (e.g. blood plasma and blood cells) at a single point in time can be used. For this ‘isotopic-clock’ principle, we present here a mathematical model that yields an analytical and easily calculated outcome.

2. Compared with a previously published model, our model assumes the isotopic difference between the old and new diets to be constant if multiple measurements are taken on the same subject at different points in time. Furthermore, to estimate the time since diet switch, no knowledge of the isotopic signature of tissues under the old diet, but only under the new diet is required.

3. The two models are compared using three calibration data sets including a novel one based on a diet shift experiment in a shorebird (red knot Calidris canutus); sensitivity analyses were conducted. The two models behaved differently and each may prove rather unsatisfactory depending on the system under investigation. A single-tissue model, requiring knowledge of both the old and new diets, generally behaved quite reliably.

4. As blood (cells) and plasma are particularly useful tissues for isotopic-clock research, we trawled the literature on turnover rates in whole blood, cells and plasma. Unfortunately, turnover rate predictions using allometric relations are too unreliable to be used directly in isotopic-clock calculations.

5. We advocate that before applying the isotopic-clock methodology, the propagation of error in the ‘time-since-diet-shift’ estimation is carefully assessed for the system under scrutiny using a sensitivity analysis as proposed here.

On 4 June 2008 Siberian Red Knots at Elbe Mouth kissed the canonical evening migration departure rule goodbye

Observations of departing Siberian-breeding Red Knots Calidris canutus canutus from their central staging site during northward migration, the Schleswig-Holstein Wadden Sea, Germany, in early June 2008, challenge the established notion that departing long-distance migrating waders only leave around sunset. During four days we scanned several thousand Red Knots for colour-ringed individuals and found a total of 20 different individuals that were previously ringed at either their main wintering site, the Banc d'Arguin in Mauritania, or at stopover sites on the Atlantic coast of France. Body masses of captured Red Knots in Schleswig-Holstein were higher than 200 g and hematocrite values showed an average of 58%, clearly indicating that they were ready for take-off. On all except one evening, we noted impressive departure movements during the incoming tide. On that exceptional evening a cold front thunderstorm passed over the area. Late the next morning, thousands of Red Knots departed during the incoming tide. We assume that the birds avoided taking off in adverse weather conditions and elaborate why Red Knots presumably traded off advantages from departing during twilight. We suggest that during spring migration, schedules are so tight that further delays decrease fitness, either because it would cause another full day of exposure to high predation risk by falcons, or because of conditions upon arrival on the tundra.

Stochastic atmospheric assistance and the use of emergency staging sites by migrants

Numerous animals move vast distances through media with stochastic dynamic properties. Avian migrants must cope with variable wind speeds and directions en route, which potentially jeopardize fine-tuned migration routes and itineraries. We show how unpredictable winds affect flight times and the use of an intermediate staging site by red knots (Calidris canutus canutus) migrating from west Africa to the central north Siberian breeding areas via the German Wadden Sea. A dynamic migration model incorporating wind conditions during flight shows that flight durations between Mauritania and the Wadden Sea vary between 2 and 8 days. The number of birds counted at the only known intermediate staging site on the French Atlantic coast was strongly positively correlated with simulated flight times. In addition, particularly light-weight birds occurred at this location. These independent results support the idea that stochastic wind conditions are the main driver of the use of this intermediate stopover site as an emergency staging area. Because of the ubiquity of stochastically varying media, we expect such emergency habitats to exist in many other migratory systems, both airborne and oceanic. Our model provides a tool to quantify the effect of winds and currents en route.

Densities of individually marked migrants away from the marking site to estimate population sizes : a test with three wader populations

Capsule Population estimates based on the mark–resighting method can be a useful alternative to population-wide counts.

Aims To investigate whether the mark–resighting method can be used as an alternative to counts to estimate the size of wader populations.

Methods Individual colour-marking and subsequent resightings allowed accurate estimates of annual survival for three populations of waders, on which basis we could estimate the actual number of marked birds alive. Densities of marked birds were determined on sites away (2000–4300 km) from the ringing locations expecting marked birds to be randomly distributed among non-marked conspecifics. Population sizes are estimated by combining these densities with the number of marked birds alive.

Results We found indications that the distribution of marked birds was indeed random in the locations away from the site of marking. The estimated population size of Red Knot Calidris canutus canutus was in accordance with the most recent estimates based on counts. Our estimate of the Calidris c. islandica population was somewhat lower, and that of the Bar-tailed Godwit Limosa lapponica taymyrensis population was considerably lower than the latest estimates based on counts.

Conclusion Population estimates based on the mark–resighting method can be a useful alternative for, or addition to, population-wide counts, as long as the assumption of random distribution of marked birds at the reading sites is taken into account. We conclude that the Afro-Siberian Bar-tailed Godwit population has recently decreased in size or has been substantially overestimated during the counts.

Basal metabolic rate, food intake, and body mass in cold- and warm-acclimated Garden Warblers

We address the question of whether physiological flexibility in relation to climate is a general feature of the metabolic properties of birds. We tested this hypothesis in hand-raised Garden Warblers (Sylvia borin), long-distance migrants, which normally do not experience great temperature differences between summer and winter. We maintained two groups of birds under cold and warm conditions for 5 months, during which their body mass and food intake were monitored. When relatedness (siblings vs. non-siblings) of the experimental birds was taken into account, body mass in cold-acclimated birds was higher than in warm-acclimated birds. BMR, measured at the end of the 5-month temperature treatment, was also higher in the cold- than the warm-acclimated group. Migrant birds thus seem to be capable of the same metabolic cold-acclimation response as has been reported in resident birds. The data support the hypothesis that physiological flexibility is a basic trait of the metabolic properties of birds.

Variation in energy intake and basal metabolic rate of a bird migrating in a wind tunnel

1. We studied the changes in body mass, metabolizable energy intake rate (ME) and basal metabolic rate (BMR) of a Thrush Nightingale, Luscinia luscinia, following repeated 12-h migratory flights in a wind tunnel. In total the bird flew for 176 h corresponding to 6300 km. This is the first study where the fuelling phase has been investigated in a bird migrating in captivity.

2. ME was very high, supporting earlier findings that migrating birds have among the highest intake rates known among homeotherms. ME was significantly higher the second day of fuelling, indicating a build-up of the capacity of the digestive tract during the first day of fuelling.

3. Further indications of an increase in size or activity level of metabolically active structures during fuelling come from the short-term variation in BMR, which increased over the 2-day fuelling period with more than 20%, and in almost direct proportion to body mass. However, mass-specific BMR decreased over the season.

4. The patterns of mass change, ME and BMR of our focal bird following two occasions of 12-h fasts were the same as after flights, indicating that fast and flight may involve similar physiological processes.

5. The relatively low ME the first day following a flight may be a contributing factor to the well-known pattern that migrating birds during stopover normally lose mass the first day of fuelling.

Is salt stress a problem for waders wintering on the Banc d'Arguin, Mauritania?

Knots Calidris canutus and sanderlings C. alba were used in cage experiments in which water and food consumption were measured under different salt regimes. Food consumption decreased and water consumption increased after changing the water provided from fresh to salt. Knots have the capability of adapting to salt water. Swallowing of adherent water with the prey, as well as evaporative cooling in heat stressed birds, might increase salt stress. -from Authors

High basal metabolic rates of shorebirds while in the Arctic: a circumpolar view

The basal metabolic rate (BMR) of Old World long-distance-migrant shorebirds has been found to vary along their migration route. On average, BMR is highest in the Arctic at the start of fall migration, intermediate at temperate latitudes, and lowest on the tropical wintering grounds. As a test of the generality of this pattern, we measured the BMR of one adult and 44 juvenile shorebirds of 10 species (1-18 individuals of each species, body-mass range 19-94 g) during the first part of their southward migration in the Canadian Arctic (68-76°N). The interspecific relationship between BMR and body mass was almost identical to that found for juvenile shorebirds in the Eurasian Arctic (5 species), although only one species appeared in both data sets. We conclude that high BMR of shorebirds in the Arctic is a circumpolar phenomenon. The most likely explanation is that the high BMR reflects physiological adaptations to low ambient temperatures. Whether the BMR of New World shorebirds drops during southward migration remains to be investigated.

O uso de praias arenosas com diferentes concentrações humanas por espécies de aves limícolas (Charadriidae e Scolopacidae) neárticas no sudeste do Brasil

As aves migratórias podem reconhecer humanos e animais domésticos como possíveis predadores, alterando seus padrões de abundância e comportamento de formação de bandos. O objetivo do presente estudo foi comparar a abundância de aves migratórias neárticas, a freqüência de bandos e o número médio de aves por bando em áreas com alta e baixa concentração humana em uma região costeira de praia arenosa no sudeste do Brasil. As aves, pessoas e cães foram contados mensalmente entre novembro de 2006 a abril de 2007. Foram registradas seis espécies de aves (Arenaria interpres, Calidris alba, Calidris canutus, Calidris pusilla, Charadrius semipalmatus, Pluvialis dominica) nas duas áreas, no entanto somente C. canutus foi registrado exclusivamente na área com baixa concentração humana. Houve diferença significativa no número médio de pessoas e cães entre as áreas, mas não no número médio de aves. Não houve correlação entre o número de humanos e aves, e entre cães e aves. Adicionalmente, não houve diferença significativa na freqüência de bandos e número de indivíduos por bando entre as áreas. Os resultados deste estudo destacaram a sensibilidade de C. canutus na área com alta concentração humana e a necessidade de futuras investigações que determinem os limites máximos de concentração de pessoas e cães domésticos que as aves migratórias neárticas podem tolerar para a tomada de ações de proteção em áreas costeiras com ocupação humana.