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Kolguev Island (69˚05′ N 49˚15′ E) is located in the Pechora Sea, the southeastern part of the Barents Sea. The island’s ecosystem is unusual due to the total absence of rodents and specialized predators such as weasels, while non-specialized predators such as Arctic (Vulpes lagopus) and red (V. vulpes) foxes and Rough-legged Hawk (Buteo lagopus) are common. Currently, 111 bird species have been registered here, of which 58 are nesting. The absence of rodents and the relatively stable predation pressure have resulted in the high abundance of many bird species: Willow Ptarmigan (Lagopus lagopus), several goose species, some waders, and passerines. Over the 125-year history of ornithological studies on Kolguev, the island avifauna has changed significantly. The trend of an increase in the proportion of widespread and Siberian species together with a decrease in the proportion of Arctic species was observed. Since 2006, a thorough monitoring of Kolguev avifauna has been carried out, during which the dynamics of the bird population densities have been traced. The abundance of Black-bellied Plover (Pluvialis squatarola) and Dunlin (Calidris alpina) decreased, while the numbers of Barnacle Goose (Branta leucopsis) have increased sharply since the 1980s. The breeding density of Rough-legged Hawk has also increased in recent years. The long-term monitoring of Kolguev ecosystems has indicated the high international conservation value of the island due to the high breeding density of many bird species. Our study, covering more than a century of avifaunal studies with almost annual monitoring over the past three decades, provides an unusually long and detailed time-series for an Arctic island.

Salt marshes are vulnerable to climate change-associated sea-level rise and storm-induced surges. Their degradation will likely affect shorebirds relying on this ecosystem. Least Sandpiper (Calidris minutilla) and Pectoral Sandpiper (Calidris melanotos) migrating along coastline habitats typically use salt marshes to rest and replenish their body reserves. Our objective was to test if width of the different vegetation zones within salt marshes affects the occurrence of Least and Pectoral Sandpipers stopping along the St. Lawrence River Estuary, Quebec, Canada, during fall migration. We established 26 survey sites, each 600 m in length, along the shoreline. Shorebird surveys were conducted in 2011 and 2012. We characterized salt marshes by measuring the width of each vegetation zone (lower marsh and upper marsh). We analyzed shorebird presence/not detected data with generalized estimating equations to test the predictions that occurrence of Least Sandpipers and Pectoral Sandpipers increases with width of both the lower and upper marsh. Upper marsh width was positively associated with probability of occurrence in each species. Our results highlight the importance of protecting the integrity of salt marshes for these two species. In the St. Lawrence River Estuary, where landward migration of salt marshes is no longer possible (coastal squeeze), effective management of shorelines is much needed. Otherwise, salt marshes and these two species could be locally jeopardized.

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.

Many migratory shorebird species using the East Asian–Australasian Flyway are declining rapidly. While the loss of staging habitats in East Asia is considered the primary cause, stressors to fitness often occur throughout the geographic range of declining species, and threats in the non-breeding grounds have been comparatively poorly studied. Three species of migratory shorebird, Curlew Sandpiper (Calidris ferruginea), Sharp-tailed Sandpiper (C. acuminata) and Red-necked Stint (C. ruficollis), use Australia’s dynamic temporary wetland systems opportunistically, yet these large wetland systems have become increasingly degraded, with reduced frequency and extent of flooding. Here, we test whether variables related to wetland availability in Australia’s interior can explain annual variation in apparent survival, abundance or immature to adult ratios at three well-monitored coastal shorebird areas in southern Australia (total area > 1315 km2). We show that coastal annual bird abundance and ratios of immatures at the coast were higher when inland Australia was relatively hot and dry. Also, a small but significant amount of variation in annual apparent survival can be explained by annual variation in inland conditions, with higher survival rates in years when inland conditions were relatively wet and cool. For the endangered Curlew Sandpiper, the impacts of Australian environmental conditions may be exacerbating the impacts of conditions experienced in other parts of its range on fitness and survival. While the effects we document here are relatively weak, they do suggest that management of inland wetlands for these shorebirds may positively affect survival rates of these sharply declining species.

Dietary specialization is common in animals and has important implications for individual fitness, inter‐ and intraspecific competition, and the adaptive potential of a species. Diet composition can be influenced by age‐ and sex‐related factors including an individual's morphology, social status, and acquired skills; however, specialization may only be necessary when competition is intensified by high population densities or increased energetic demands.To better understand the role of age‐ and sex‐related dietary specialization in facilitating seasonal resource partitioning, we inferred the contribution of biofilm, microphytobenthos, and benthic invertebrates to the diets of western sandpipers (Calidris mauri) from different demographic groups during mid‐winter (January/February) and at the onset of the breeding migration (April) using stable isotope mixing models. Western sandpipers are sexually dimorphic with females having significantly greater body mass and bill length than males.Diet composition differed between seasons and among demographic groups. In winter, prey consumption was similar among demographic groups, but, in spring, diet composition differed with bill length and body mass explaining 31% of the total variation in diet composition. Epifaunal invertebrates made up a greater proportion of the diet in males which had lesser mass and shorter bills than females. Consumption of Polychaeta increased with increasing bill length and was greatest in adult females. In contrast, consumption of microphytobenthos, thought to be an important food source for migrating sandpipers, increased with decreasing bill length and was greatest in juvenile males.Our results provide the first evidence that age‐ and sex‐related dietary specialization in western sandpipers facilitate seasonal resource partitioning that could reduce competition during spring at the onset of the breeding migration.Our study underscores the importance of examining resource partitioning throughout the annual cycle to inform fitness and demographic models and facilitate conservation efforts., Our results provide the first evidence of seasonal differences in diet composition among demographic groups of western sandpipers, supporting the conclusion that age‐ and sex‐related dietary specialization facilitate seasonal resource partitioning in this species. Differences in diet composition among demographic groups were more pronounced at the onset of migration in spring compared to mid‐winter following the expectation that the consumption of biofilm and microphytobenthos provides energy for sandpipers during their breeding migration and reduces competition when high densities of birds occupy foraging sites. In spring, juveniles, particularly juvenile males, consumed more biofilm and microphytobenthos than adults supporting our hypothesis that sandpipers with shorter bills, lesser mass, and less prey handling experience would consume the greatest amount of biofilm and microphytobenthos. Photo credit: William Chan.

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.

The gut microbiota of birds is known to be characterized for different species, although it may change with feeding items. In this study, we compared the gut microbiota of birds with different feeding behaviors in the same habitat. We collected fecal samples from three Arctic species, snow buntings Plectrophenax nivalis, sanderlings Calidris alba, and pink‐footed geese Anser brachyrhynchus that are phylogenetically quite distant in different families to evaluate effects of diet on gut microbiota. Also, we characterized the prevalence of fecal bacteria using the Illumina MiSeq platform to sequence bacterial 16S rRNA genes. Our NMDS results showed that fecal bacteria of snow buntings and sanderlings were significantly distant from those of pink‐footed geese. Although all three birds were occupied by three bacterial phyla, Proteobacteria, Firmicutes, and Bacteroidetes, dominant taxa still varied among the species. Our bacterial sequences showed that snow buntings and sanderlings were dominated by Firmicutes and Bacteroidetes, while pink‐footed geese were dominated by Proteobacteria. In addition, the bacterial diversity in snow buntings and sanderlings was significantly higher than that in pink‐footed geese. Our results suggest that insectivorous feeding diet of snow buntings and sanderlings could be responsible for the similar bacterial communities between the two species despite the distant phylogenetic relationship. The distinctive bacterial community in pink‐footed geese was discussed to be related with their herbivorous diet., We showed that fecal bacteria of snow buntings and sanderlings were distant from those of pink‐footed geese. In addition, the bacterial diversity in snow buntings and sanderlings was significantly higher than that in geese. Our results suggest that diet as well as host phylogeny may affect the gut microbiota of Arctic birds.

Antipredator vigilance may be affected by feeding techniques that obstruct animal’s vision, such as underwater foraging. I examined the effect of underwater foraging on head-up position in spring staging dunlins (Calidris alpina), predicting that this type of foraging would positively affect the frequency of head-up vigilance in this species. The results of analysis showed that significant predictors of the occurrence of head-up positions were underwater foraging and head-up position that occurred in the imminent past. Results of this study imply that dunlins are less likely to raise their heads above the shoulder line if they have not obstructed their vision by underwater foraging in the previous moment and that birds are able to acquire visual information from their surroundings not only when their heads are in the upright position but also during head-down foraging and/or during horizontal head position.

n Focus: Reneerkens, J., Versluijs, T. S. L., Piersma, T., Alves, J. A., Boorman, M., Corse, C., ... Lok, T. (2020). Low fitness at low latitudes: wintering in the tropics in-creases migratory delays and mortality rates in an Arctic breeding shorebird. Journal of Animal Ecology, 89, 691–703. A central question in migratory ecology has been to understand the fitness consequences of individual variation in migration distance among different species and populations. Reneerkens et al. (2020) investigated the demographic consequences of long-distance migration for Sanderlings Calidris alba, an Arctic-breeding species of sandpiper. Their study population has a remark-able geographic distribution with a breeding range that is concentrated in north-east Greenland and Ellesmere Island, Canada but a nonbreeding range that extends across 85° of latitude from Scotland to Namibia. The authors report on unexpected patterns of latitudinal variation in three demographic parameters: timing of passage on northward migration, probability of juvenile migration and apparent survival of adults. Sanderlings travelling 1,800–2,800 km to settle at north temperate sites dur-ing the nonbreeding season had earlier passage dates, and also higher probabilities of migration and apparent survival. In contrast, birds travelling 6,000–7,800 km to equato-rial sites experienced later passage dates, delayed maturity and lower apparent survival. However, if Sanderlings migrated even farther and flew over 11,000 km to nonbreed-ing sites in Namibia, then their performance was restored to early passage dates and higher survival. Movement tracks from birds tagged with geolocators showed that birds wintering in Namibia make nonstop flights of 7,500 km that bypass West Africa during northward migration. Thus, all lines of evidence suggest that Sanderlings face adversity when spending the nonbreeding season at equatorial latitudes. Moreover, the central finding that components of fitness can have nonlinear relationships with migration dis-tance is a novel discovery that leads to many additional questions. The new findings have broader implications for theoretical models of migration, and for understanding how different patterns of movements may arise or be maintained in migratory species

Present-day ecology and population structure are the legacies of past climate and habitat perturbations, and this is particularly true for species that are widely distributed at high latitudes. The red knot, Calidris canutus, is an arctic-breeding, long-distance migratory shorebird with six recognized subspecies defined by differences in morphology, migration behavior, and annual-cycle phenology, in a global distribution thought to have arisen just since the Last Glacial Maximum (LGM). We used nextRAD sequencing of 10,881 single-nucleotide polymorphisms (SNPs) to assess the neutral genetic structure and phylogeographic history of 172 red knots representing all known global breeding populations. Using population genetics approaches, including model-based scenario-testing in an approximate Bayesian computation (ABC) framework, we infer that red knots derive from two main lineages that diverged ca. 34,000 years ago, and thus persisted at the LGM in both Palearctic and Nearctic refugia, followed by at least two instances of secondary contact and admixture. In two flyways, we detected clear genetic structure between population pairs with similar migrations and substantial geographic overlap in the non-breeding season. Conversely, other populations were only weakly differentiated despite clearly divergent migratory phenotypes and little or no apparent contact throughout the annual cycle. In general, the magnitude of genetic differentiation did not match that of phenotypic differences among populations, suggesting that flyway-specific phenotypes developed quite rapidly and do not necessarily impose barriers to gene flow. Our results suggest that population structure and migratory phenotypes in red knots arose from a complex interplay among phylogeography, plasticity, and selective processes.

Estuaries have historically been subject to considerable habitat loss, and continue to be subjected to such in areas where the natural landward migration of intertidal habitats is constrained by hard coastal defences. Thus, in estuaries where direct (e.g., port development) or indirect (e.g., sea level rise) processes are predicted to threaten intertidal habitats and associated waterbird species, there is a regulatory requirement to produce compensatory intertidal habitats. Managed realignment (MR) is a shoreline management practise that is undertaken to build sustainable coastal defences and create intertidal habitats in estuaries. This nature-based solution brings multiple benefits in the form of carbon storage, increased resilience to flooding, and, potentially, the formation of new habitats, which is the topic of this study. A 75-ha site at the Paull Holme Strays (Humber Estuary, United Kingdom) was monitored over a 10-year period following MR to examine the change in the abundance of waterbirds in the chosen site in response to the physical processes occurring there. Using digital terrain models (DTMs) collected via light detection and ranging (LiDAR), we examined how four compensatory target species responded to changes in elevation after the creation of the site. It was shown that the very rapid accretion of estuarine sediment occurred in the first decade of the new re-created intertidal, which, over time, led to changes in the numbers of benthic foraging birds supported. Furthermore, elevation change was also driven by this sediment accretion, the rate of which depended on the initial bed elevation of the sectors within the site. Ten years after the recreation of the habitat, the spatial heterogeneity in the bed elevation remained high; however, the sectors with the lowest elevations accreted the most over the 10-year period. The foraging number of the four waterbird species that colonised the MR site significantly declined above a certain elevation, with this effect being most pronounced for the Eurasian curlew (Numenius arquata). The number of common shelducks (Tadorna tadorna), dunlins (Calidris alpina), and common redshanks (Tringa totanus) declined significantly after initial peaks 5–7 years after the creation of the site, reflecting the ongoing elevation changes. Thus, this study highlighted the need for long-term studies to understand how species respond to large-scale habitat construction. It can also aid in predicting the suitability of an MR site for waterbirds in the medium and long term.

Previous studies have highlighted the importance of intertidal habitats on the coast of Jiangsu Province, China for the Critically Endangered Spoon-billed Sandpiper Calidris pygmaea during the post-breeding moult. We report Lincoln-Petersen closed-population estimates of numbers of Spoon-billed Sandpipers Calidris pygmaea at the three principal sites in the region for the species: Tiaozini, Yangkou and Dongling. Our estimates were based upon resightings and scan surveys of individually-marked birds. Surveys were conducted in September–October when nearly all of the birds present were adults (≥1 yr old). We estimated the number present at Tiaozini in each of the years 2017–2020, during which numbers declined from 225 to 151, mostly between 2018 and 2019. To identify parts of the intertidal area used by Spoon-billed Sandpipers, we mapped locations of satellite-tagged birds in 2017 and 2019. Data from seven tagged birds indicated that they moved to intertidal flats up to 7 km seaward of the nearest point of the seawall at low tide. We then analysed satellite imagery for 2017–2020 for intertidal areas up to this distance from the coast and found evidence for changes in topography of the intertidal mudflats and positions of major channels. The mean extent of exposed mudflats per tidal cycle appeared to have declined by about 20% between 2017 and 2020. Surveys at two other sites in 2020 indicated that there were 26 birds at Yangkou and 63 birds at Dongling, with a total of 240 birds at all three sites combined in 2020. This is approximately half of the estimated world population of adults. The coast of Jiangsu Province is therefore of global importance for the Spoon-billed Sandpiper. Although birds no longer at Tiaozini may have been displaced to other sites, the observed population decline, which co-incided with changes in intertidal topography, raises concerns about the future conservation status of the species. Uncertainty about future changes to the topography and extent of intertidal mudflats makes the co-ordinated protection of all sites in Jiangsu Province known to hold Spoon-billed Sandpipers during the post-breeding moult period a conservation priority. This applies particularly to Tiaozini, Yangkou and Dongling, but also to other sites at which future surveys may reveal to hold concentrations of the species.

High‐quality staging sites are critical for long‐distance migratory shorebirds to rest and refuel but are under threat from human development, including expansion of wind energy projects. However, predicting migration timing and movements in relation to weather conditions at staging sites can increase our understanding and mitigate effects of wind turbine collisions. Here we assessed northward migration timing and orientation in relation to environmental conditions at an inland staging area in Saskatchewan, Canada, with active and proposed wind energy developments. The area is known to host ~25% of North America's Sanderling Calidris alba population and 16 other Arctic‐breeding migrant shorebird species. We quantified arrival and departure time of day in relation to weather using data from 140 of 237 Sanderlings radiotagged locally and at a southern staging site in the Gulf of Mexico with the Motus Wildlife Tracking System (April–June, 2015–2017). Although Sanderling arrival times were not related to time of day or weather, departures were more likely at sunset in winds blowing towards the northwest at intermediate speeds (

Protected areas (PAs) are a cornerstone of modern conservation. For PAs that are established to conserve mobile species, it is important to cover all the key areas regularly used by these species. However, zonation and boundaries of PAs have often been established with limited knowledge of animal movements, leaving the effectiveness of some PAs doubtful. We used radio tracking data to evaluate the extent to which two coastal PAs in mainland China encompassed the full range of habitats used by migratory shorebirds during non-breedingseasons. The core zone (highest restriction on human activities) of the Yalu Jiang Estuary National Nature Reserve (Liaoning) incorporated only 22 ± 6% (n=34) of the diurnal home range (95% kernel density) of the endangered great knots Calidris tenuirostris. In contrast, the core zone of Chongming Dongtan (Shanghai) incorporated 73 ± 24% (n=25) of the home range of dunlins Calidris alpine. During high tide, great knots in Yalu Jiang mostly occurred in the experimental zone (least restriction on human activities) or sometimes outside the PA boundary altogether, where the birds could face substantial threats. By investigating satellite tracking records, consulting published literature, interviewing local experts and mapping habitat composition in different coastal PAs in China, we found that wet artificial supratidal habitats were frequently used by migratory shorebirds but the coverage of these habitats in coastal PAs was low. These PA boundaries and/or zonations should be revised to conserve mobile species more effectively. With the increasing number of tracking studies, analysing the spatial relationships between PAs and the movement ranges of mobile species can increasingly inform the development of a representative, comprehensive PA network.

Dunlin Calidris alpina has been recorded as breeding in China based on a pullus collected on 7 May 1927 at Tsingtao, Shandong province; a remarkably early date for a nearly fledged young. The record appears highly unlikely to be an example of deliberate fraud, and there is no obvious evidence of mislabelling. We have been unable to determine the validity of the claimed breeding record, however, based on a review of available information, it appears certain that the species does not currently breed in China.

Estuaries of major rivers provide important stopover habitat for migratory birds throughout the world. These estuaries experience large amounts of freshwater inputs from spring runoff. Understanding how freshwater inputs affect food supply for migrating birds, and how birds respond to these changes will be essential for effective conservation of critical estuarine habitats. We estimated trends over time in counts of Western Sandpiper (Calidris mauri) and Pacific Dunlin (Calidris alpina pacifica) during northward migration on the Fraser River estuary, British Columbia, Canada, where shorebirds feed extensively on intertidal biofilm and invertebrates. We also examined whether counts were correlated with a suite of environmental variables related to local conditions (precipitation, temperature, wind speed and direction, solar radiation, tidal amplitude, and discharge rates from the Fraser River) during a total of 540 surveys from 1991 to 2019. Counts of Western Sandpiper declined ~54% (−2.0% per annum) over the entire study period, and 23% from 2009 to 2019 (−0.9% per annum). Counts of Pacific Dunlin did not show a statistically significant change over the study period. Counts of shorebirds were lower when discharge from the Fraser River was high, which we propose results from a complex interaction between the abrupt changes in salinity and the estuarine food web related to the quantity or quality of intertidal biofilm. Counts were also higher when tidal amplitude was lower (neap tides), potentially related to longer exposure times of the mudflats than during spring tides. Effects of wind are likely related to birds delaying departure from the stopover site during unfavorable wind conditions. The negative trend in migrating Western Sandpipers is consistent with declines in nonbreeding areas as observed in Christmas Bird Counts. Understanding causes of population change in migratory shorebirds highlights the need for research on mechanistic pathways in which freshwater inputs affect food resources at estuarine stopovers., Shorebird counts at a critical coastal stopover site have declined over a 29 year period (1991–2019). Counts were correlated with tidal amplitude, wind, and river discharge. We hypothesize a complex interaction between the abrupt changes in salinity from freshwater inputs, a decrease in quality and quantity of intertidal biofilm, and shorebird counts.

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.

The degree to which individuals migrate among particular breeding, migration, and wintering sites can have important implications for prioritizing conservation efforts. Four subspecies of Dunlin (Calidris alpina) migrate along the East Asian-Australasian Flyway. Each subspecies has a distinct and well-defined breeding range, but their migration and winter ranges are poorly defined or unknown. We assessed the migratory connectivity of 3 of these subspecies by evaluating a dataset that encompasses 57 yr (1960-2017), and comprises more than 28,000 Dunlin banding records and 818 observations (71 recaptures and 747 band resightings). We present some of the first evidence that subspecific segregation likely occurs, with arcticola Dunlin wintering in areas of Japan, and other arcticola, actites, and sakhalina Dunlin wintering in areas of the Yellow and China seas. Observations indicate that whether an arcticola Dunlin winters in Japan or the Yellow and China seas is independent of their breeding location, sex, or age. Furthermore, observations indicate that ≥83% of arcticola Dunlin exhibit interannual site fidelity to specific wintering sites. This suggests that the degradation of specific wetland areas may negatively affect particular individuals of a particular subspecies (or combination of subspecies), and, if widespread, could result in population declines. Given the possible biases inherent in analyzing band recovery data, we recommend additional flyway-wide collaboration and the use of lightweight tracking devices and morphological and genetic assignment techniques to better quantify subspecies' migratory movements and nonbreeding distributions. This information, when combined, will enable effective conservation efforts for this species across the East Asian-Australasian Flyway. LAY SUMMARY The East Asian-Australasian Flyway has more threatened and near-Threatened migratory waterbird species than any other flyway in the world; however, developing flyway conservation plans has been challenging, in part due to limited information regarding population-specific migration patterns. Four subspecies of Dunlin migrate and winter along the East Asian-Australasian Flyway. Each has a well-defined breeding range; their migration and winter ranges are poorly defined or unknown. We assessed the migratory connectivity of 3 subspecies using data from 57 yr (1960-2017), which comprises more than 28,000 Dunlin banding records, 71 recaptures, and 747 band resightings. Subspecific segregation likely occurs, with arcticola Dunlin wintering in Japan, and other arcticola, actites, and sakhalina Dunlin wintering in the Yellow and China seas. It is likely that ≥83% of arcticola Dunlin exhibit interannual site fidelity to specific wintering sites. Our findings suggest that degradation of specific wetland areas may negatively affect particular individuals of a particular subspecies (or combination of subspecies) and could result in population declines.

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.

Background Age at maturity and the timing of first breeding are important life history traits. Most small shorebird species mature and breed as ‘yearlings’, but have lower reproductive success than adults. In some species, yearlings may defer northward migration and remain in non-breeding regions (‘oversummering’) until they reach 2 years of age. Some adults also oversummer. Oversummering would be favoured by natural selection if survival were as a result raised sufficiently to compensate for the missed breeding opportunity. Several thousand Semipalmated Sandpipers (Calidris pusilla) spend the non-breeding period at Paracas, Perú, including individuals with long bills (likely from eastern Arctic breeding populations ~ 8000 km distant) and short bills (likely from western Arctic breeding populations, up to 11,000 km distant), with short-billed birds more likely to oversummer. We tested the prediction that oversummering birds have higher survival than migrants, and that the magnitude of this higher survival for oversummering birds is enough to compensate for their lost breeding season. Methods We used a Multi-State Mark-Recapture model based on 5 years of encounter data (n = 1963 marked birds, and 3229 resightings) obtained year-round at Paracas, Perú, to estimate seasonal (i.e. breeding and non-breeding) survivorship for migrant and oversummering birds. We calculated the magnitude of the oversummering survival advantage required to compensate, for both yearlings and adults, based on published measures of annual survival and reproductive success. Using bill length as a proxy for migration distance, we investigated whether migratory survival is distance-dependent. Results We estimate that 28% of yearlings and 19% of adults oversummer. Survival is higher for oversummering birds than for migrants, and the oversummering survival advantage is greater for adults (0.215) than for yearlings (0.140). The theoretical thresholds predicted by the size of the missed reproductive opportunity are 0.240 for adults and 0.134 for yearlings. Migratory survival decreases and the oversummering rate increases with migration distance, as assessed by culmen length. Conclusions Our results support the life history hypothesis that oversummering raises survival enough to compensate for the loss of a breeding opportunity. Greater migration distance lowers survival and increases the probability of oversummering.

Many migratory birds are declining worldwide. In line with the general causes for the global biodiversity crisis, habitat loss, pollution, hunting, over-exploitation and climate change are thought to be at the basis of these population declines. Long-distance migrants seem especially vulnerable to rapid anthropogenic change, yet, the rate of decline across populations and species varies greatly within flyways. We hypothesize that differences in migration strategy, and notably stopover-site use, may be at the basis of these variations in resilience to global change. By identifying and comparing the migration strategies of two very closely related shorebird species, the Curlew sandpiper Calidris ferruginea and the Red-necked stint Calidris ruficollis, migrating from the same non-breeding site in Australia to similar breeding sites in the high Russian Arctic, we aimed to explain why these two species express differential resilience to rapid changes within their flyway resulting in different population trajectories in recent times. Based on 13 Curlew sandpiper and 16 Red-necked stint tracks from light-level geolocator tags, we found that individual Curlew sandpipers make use of fewer stopover areas along the flyway compared to Red-necked stints. Furthermore, and notably during northward migration, Curlew sandpipers have a higher dependency on fewer sites, both in terms of the percentage of individuals visiting key stopover sites and the relative time spent at those sites. While Curlew sandpipers rely mainly on the Yellow Sea region, which has recently experienced a sharp decline in suitable habitat, Red-necked stints make use of additional sites and spread their relative time en-route across sites more evenly. Our results indicate that differential migration strategies may explain why Curlew sandpipers within the East Asian-Australasian Flyway are declining rapidly (9.5%-5.5% per year) while Red-necked stints remain relatively stable (-3.1%-0%). We consider that more generally, the number of sites per individual and among a population, the spatial distribution across the flyway, as well as the relationship between the time spent over sites may prove to be key variables explaining populations and species' differential resilience to environmental change.

Waders normally fly with their legs stretched out under the tail, with their feet pointing backwards. Whereas there are a few published records of waders flying with their legs folded forward (like a passerine), with legs and feet tucked in under the belly feathers, the causes and consequences of waders flying with folded legs are poorly understood. Through searching the literature, asking ornithologists in person and through public calls, and from searching photos on the internet, we retrieved 90 records of 17 different species where one or more individuals flew with folded legs. It was most common in Spotted Redshank Tringa erythropus (in up to 10% of all birds) and Common Redshank T. totanus, but was also observed in short-legged species such as Dunlin Calidris alpina and Common Ringed Plover Charadrius hiaticula. The behaviour was recorded in most months of the year, in different age classes, in single birds as well as in flocks, at different ambient temperatures, and in both migratory flights and local move-ments. Whereas reducing heat loss at low temperatures is a likely, but not omnipo-tent, explanation for waders flying with folded legs, aerodynamics and leg muscle fatigue may also affect how waders keep their legs in flight. (Less)

The Bay of Fundy, Canada is a critical migratory stopover for Semipalmated Sandpipers (Calidris pusilla) during fall migration. Recent ecological changes combined with range-wide population declines indicate that a re-evaluation of habitat use and regional fidelity is required. The Bay of Fundy is divided into three regions, each of which contains multiple foraging and roost sites. In this study, we examined local movement and regional fidelity within and between years using radio-telemetry and field-readable markers. We deployed 194 radio-transmitters and 2295 field-readable flags on Semipalmated Sandpipers during migration in 2013-2014, and resighted these birds, along with additional birds flagged in 2012, between 2013 and 2016. Radio-tracked birds used on average 1.3 to 4.1 foraging and roost sites per day, with most making multiple daily movements, but only 2% made permanent moves between geographically distant regions within the bay during stopover. Additionally, 97% of birds returned to their original region in subsequent years. The use of multiple sites suggests Semipalmated Sandpipers are capable of adapting to dynamic conditions in the Bay of Fundy on a local scale, but the high regional fidelity suggests that their ability to adapt to changes occurring at a regional scale may be limited. Uncertainty about the capability of Semipalmated Sandpipers to flexibly move among different regions of the Bay of Fundy supports the need for conservation at both the site and regional level.

The spoon-billed sandpiper Calidris pygmaea, a migratory Arctic-breeding shorebird, is one of the rarest birds and its population has declined since the 1970s. We surveyed its most important known wintering area in the Upper Gulf of Mottama in Myanmar to estimate recent (2009–2016) changes in its numbers there. The total number of small shorebirds present in the Upper Gulf was counted and the proportion of them that were spoon-billed sandpipers was estimated from sample scans. These two quantities were multiplied together to give the estimated number of spoon-billed sandpipers in each of 4 years. Total numbers of combined small shorebird species tripled from 21,000 to 63,000 between 2009 and 2016, coincident with efforts to reduce hunting pressure on waterbirds. However, the proportion of small shorebirds that were spoon-billed sandpipers declined and their estimated absolute numbers fell by about half, from 244 to 112 individuals. It is probable that loss of intertidal habitat and shorebird hunting elsewhere on the migration route of the spoon-billed sandpipers wintering at Mottama is causing a continued decline, although this is occurring at a less rapid rate than that recorded from Arctic Russia before 2010. The number of spoon-billed sandpipers wintering on the Upper Gulf of Mottama remains the highest single-site total for this species from any known wintering site. Preventing resurgence of illegal shorebird hunting and ensuring long-term protection of the intertidal feeding habitats and roost sites in the Gulf are high priorities if extinction of this species is to be averted.

We examined the influence of the density of foragers on feeding rates of Semipalmated Sandpipers (Calidris pusilla (Linnaeus, 1766)) while using different foraging modes at a spring stopover site in Delaware Bay, USA. Using dynamic estimates of interindividual distances obtained at short intervals of time, we explored how forager density affected feeding rates when Semipalmated Sandpipers used visual pecking or tactile probing. Pecking rate significantly increased with interindividual distances, whereas probe rate was not affected by density. Our study also showed that in fast-moving foragers, such as Semipalmated Sandpipers, in which the number of nearby foragers and distance to the nearest neighbour continuously change throughout the foraging bout, pecking rates are more affected by nearest neighbour distance than by the number of foragers in their immediate vicinity. In addition, our study implies that foragers using different foraging modes might be differently affected by nearby competitors perhaps in response to prey disturbance by neighbours.

Understanding spatial connectivity of long-distance migrants is important for effective management and conservation of both game and nongame species. Hunting of Nearctic-breeding shorebirds occurs in the Caribbean and northern South America; however, the origins of harvested individuals are generally unknown. We used stable hydrogen isotopes (δ2H) in feathers of juvenile shorebirds to infer the origins of birds harvested at 2 sites in Barbados using probabilistic assignments based on a terrestrial–freshwater δ2H isoscape. We used tissue δ13C and δ15N values to filter individuals that had derived nutrients from marine sources. Natal origins of juvenile American Golden-Plover (Pluvialis dominica), Stilt Sandpiper (Calidris himantopus), Short-billed Dowitcher (Limnodromus griseus), and Lesser Yellowlegs (Tringa flavipes) were predicted to be mainly from the eastern parts of their breeding ranges in eastern Canada, with American Golden-Plover, Stilt Sandpiper, and Short-billed Dowitcher also having h...

Changes in demographic rates underpin changes in population size, and understanding demographic rates can greatly aid the design and development of strategies to maintain populations in the face of environmental changes. However, acquiring estimates of demographic parameters at relevant spatial scales is difficult. Measures of annual survival rates can be particularly challenging to obtain because large-scale, long-term tracking of individuals is difficult and the resulting data contain many inherent biases. In recent years, advances in both tracking and analytical techniques have meant that, for some taxonomic groups, sufficient numbers of survival estimates are available to allow variation within and among species to be explored. Here we review published estimates of annual adult survival rates in shorebird species across the globe, and construct models to explore the phylogenetic, geographical, seasonal and sex-based variation in survival rates. Models of 295 survival estimates from 56 species show that survival rates calculated from recoveries of dead individuals or from return rates of marked individuals are significantly lower than estimates from mark–recapture models. Survival rates also vary across flyways, largely as a consequence of differences in the genera that have been studied and the analytical methods used, with published estimates from the Americas and from smaller shorebirds (Actitis, Calidris and Charadrius spp.) tending to be underestimated. By incorporating the analytical method used to generate each estimate within a mixed model framework, we provide method-corrected species-specific and genus-specific adult annual survival estimates for 52 species of 15 genera.

Many north American shorebird populations are declining. it is therefore urgent to identify major sites used during their annual cycle to achieve effective conservation measures. our objective was to expand some aspects of the knowledge base needed to assess the ecological value of the St. Lawrence River Estuary for shorebird conservation. Here, we present the results of the most intensive shorebird survey ever conducted in the St. Lawrence River Estuary during fall migration. Surveys were conducted between St-Jean-Port-Joli and St-Simon-sur-Mer, Quebec, Canada, in 2011 and 2012, from late June/early July through late november, corresponding to the migration period of all species potentially present in the study area. The Semipalmated Sandpiper (Calidris pusilla) was one of the two most abundant species during both years of our study (most abundant species, followed by Dunlin [Calidris alpina] and Black-bellied Plover [Pluvialis squatarola] in 2011; second to Blackbellied Plover in 2012). Considering the entire shorebird community, abundance of individuals peaked in early September. Peak abundance occurred earlier for adults than for juveniles. For most species, juveniles largely outnumbered adults. Juveniles were relatively less abundant in 2012 than in 2011. This reflected a general trend observed in northeastern north America between those years, suggesting a lower breeding success in 2012. Given its importance as a staging site for juvenile birds (study area used annually by up to a few hundred thousand shorebirds) and therein, its conservation value, we recommend that the St. Lawrence River Estuary should be included within the Western Hemisphere Shorebird Reserve network.

Gut microbes are increasingly recognised for their role in regulating an animal's metabolism and immunity. However, identifying repeatable associations between host physiological processes and their gut microbiota has proved challenging, in part because microbial communities often respond stochastically to host physiological stress (e.g. fasting, forced exercise or infection). Migratory birds provide a valuable system in which to test host-microbe interactions under physiological extremes because these hosts are adapted to predictable metabolic and immunological challenges as they undergo seasonal migrations, including temporary gut atrophy during long-distance flights. These physiological challenges may either temporarily disrupt gut microbial ecosystems, or, alternatively, promote predictable host-microbe associations during migration. To determine the relationship between migration and gut microbiota, we compared gut microbiota composition between migrating and non-migrating ("resident") conspecific shorebirds sharing a flock. We performed this across two sandpiper species, Calidris ferruginea and Calidris ruficollis, in north-western Australia, and an additional C. ruficollis population 3,000 km away in southern Australia. We found that migrants consistently had higher abundances of the bacterial genus Corynebacterium (average 28% abundance) compared to conspecific residents (average

We recorded 2 new bird species in Entre Ríos province, Argentina: Calidris canutus (Linnaeus, 1758) and Trogonsurrucura Vieillot, 1817. In addition, we present 25 noteworthy records for bird species in El Palmar National Park,Pre Delta National Park, and other parts of the province.

Regular counts of migrating animals at stopover sites have been used as a measure of site importance at the global scale as well as for monitoring long-term population changes. However, migratory passage can last for several weeks and the turnover rate of individuals is often high, preventing the use of peak counts to estimate the total number of migrants. This estimate can be achieved, however, by combining count data with information on stopover length. Here, we developed a new method to quantify the total number of migrant birds using stopover areas hosting overlapping populations of local (breeding or wintering) and passage individuals of a given species. We illustrate the application of this method by estimating the number of spring migrant dunlins Calidris alpina stopping over at the Tagus estuary (Portugal). We used carbon and nitrogen isotopic signatures in toenails to identify migrants and to estimate their proportion in total counts. We then employed isotopic clock models to determine time since arrival (TSA) of individuals from signatures in red blood cells and plasma and, using a simulation approach, we derived the relationship between TSA and stopover length. Finally, by dividing the number of migrants.day (obtained from counts) by the stopover length, we assessed the size of the migrant population. We demonstrated that stopover length can be directly predicted from mean TSA values of birds sampled during migration, regardless of variations in (1) migratory rate, that is, the phenology of arrivals throughout the migratory period and (2) the number of birds involved in the migratory event. Migrant dunlins stayed on average 7.5 days at the Tagus estuary during spring migration, which combined with count data resulted in an estimate of c. 30,000 passage dunlins, using the study site. Our novel approach is not strict in its assumptions, and therefore can be customised to the specificities of different taxa and study areas. Estimates of total number of migrants along migratory flyways are critical to identify priority sites for conservation action. In fact, single stopover sites may host a very significant proportion of a global population albeit for a short period.

10 pages; International audience; Birds exhibit a wide diversity of breeding strategies. During incubation or chick-rearing, parental care can be either uniparental, by either the male or the female, or biparental. Understanding the selective pressures that drive these different strategies represents an exciting challenge for ecologists. In this context, assigning the type of parental care at the nest (e.g. biparental or uniparental incubation strategy) is often a prerequisite to answering questions in evolutionary ecology. The aim of this study was to produce a standardized method unequivocally to assign an incubation strategy to any Sanderling Calidris alba nest found in the field by monitoring nest temperature profiles. Using drops of >3 °C in nest temperature (recorded with thermistors) to distinguish incubation and recess periods, we showed that the number of recesses and the total duration of these recesses from 09:00 to 17:00 h UTC allowed us reliably (99.1% after 24 h and 100% when monitoring the nest for at least 4 days) to assign the incubation strategy at the nest for 21 breeding adults (14 nests). Monitoring nest temperature for at least 24 h is an effective method to assign an incubation strategy without having to re-visit nests, thereby saving time in the field and minimizing both disturbance and related increase in predation risk of clutches. Given the advantages of our method, we suggest that it should be used more widely in studies that aim to document incubation strategies and patterns in regions where ambient temperatures are at least 3 °C below the median nest temperature.

Migratory birds encounter a broad range of pathogens during their journeys, making them ideal models for studying immune gene evolution. Despite the potential value of these species to immunoecology and disease epidemiology, previous studies have typically focused on their adaptive immune gene repertoires. In this study, we examined the evolution of innate immune genes in three long-distance migratory waders (order Charadriiformes). We analysed two parts of the extracellular domains of two Toll-like receptors (TLR3 and TLR7) involved in virus recognition in the Sanderling (Calidris alba), Red-necked Stint (Calidris ruficollis), and Ruddy Turnstone (Arenaria interpres). Our analysis was extended to 50 avian species for which whole-genome sequences were available, including two additional waders. We found that the inferred relationships among avian TLR3 and TLR7 do not match the whole-genome phylogeny of birds. Further analyses showed that although both loci are predominantly under purifying selection, the evolution of the extracellular domain of avian TLR3 has also been driven by episodic diversifying selection. TLR7 was found to be duplicated in all five wader species and in two other orders of birds, Cuculiformes and Passeriformes. The duplication is likely to have occurred in the ancestor of each order, and the duplicated copies appear to be undergoing concerted evolution. The phylogenetic relationships of wader TLR7 matched those of the five wader species, but that of TLR3 did not. Instead, the tree inferred from TLR3 showed potential associations with the species' ecology, including migratory behaviour and exposure to pathogens. Our study demonstrates the importance of combining immunological and ecological knowledge to understand the impact of immune gene polymorphism on the evolutionary ecology of infectious diseases.

Determining the sex of individuals through molecular methods is an increasinglyimportant research tool, particularly for studies of sexually monomorphic species.However, currently available primer sets to molecularly sex birds are known tocause difficulties in sexing Red Knots Calidris canutus and several other waders,due to amplification failure of, and polymorphisms on, the CHD-Z gene. Tocounter these problems, we developed a new primer set, 2602F/2669R, andshow that these primers can be used to reliably sex Red Knots and eight otherwader species in four genera. We expect these primers to be useful in a widerange of wader and possibly non-wader species.

Semipalmated Sandpipers (Calidris pusilla (L., 1766)) that migrate through the upper Bay of Fundy, Canada, depend on a rich food supply to fuel their continued migration. Although past studies have reported a diet dominated by the amphipod Corophium volutator (Pallas, 1766), an animal rich in n–3 polyunsaturated fatty acids (PUFAs), recent evidence suggests that sandpiper diets are broad. This is beneficial in that it allows Semipalmated Sandpipers to respond to a changing food base, but quality of food is also important. PUFAs are important in providing the energy required by migrating birds and may play a role in migratory preparation. We assessed fatty acid (FA) concentrations and proportions in three common food items. We found that polychaetes should adequately meet the needs of migrating sandpipers in terms of FA composition. Concentrations of FAs in biofilm were low, but proportionally, n–3 PUFAs were well represented, particularly in biofilm collected in Shepody Bay, where it forms a substantial part of the sandpiper diet. Therefore, provided that birds can consume a sufficient volume of biofilm, it is also probably a suitable source of essential FAs. Our results suggest that Semipalmated Sandpipers in the Bay of Fundy can meet their FA needs with a variety of dietary options.

River deltas along Alaska’s Beaufort Sea coast are used by hatch-year semipalmated sandpipers (Calidris pusilla) after leaving their terrestrial natal sites, but the drivers of their use of these stopover sites on the first “hop” of fall migration are unknown. We quantified sandpiper temporal distribution and abundance as related to food resources at three river deltas during the beginning of their fall migration (post-breeding period) to compare the habitat quality among these deltas. We conducted population counts, sampled invertebrates, and captured birds to collect blood samples from individuals for triglyceride and stable isotope analyses to determine fattening rates and diet. Patterns of sandpiper and invertebrate abundance were complex and varied among deltas and within seasons. River deltas were used by sandpipers from late July to late August, and peak sandpiper counts ranged from 1000 to 4000 individuals, of which 98% were hatch-year semipalmated sandpipers. Isotopic signatures from blood plasma samples indicated that birds switched from a diet of upland tundra to delta invertebrate taxa as the migration season progressed, suggesting a dependence on delta invertebrates. Despite differences in diet among deltas, we found no differences in fattening rates of juvenile sandpipers as indicated by triglyceride levels. The number of sandpipers was positively associated with abundance of Amphipoda and Oligochaeta at the Jago and Okpilak-Hulahula deltas; an isotopic mixing model indicated that sandpipers consumed Amphipoda and Oligochaeta at Jago, mostly Chironomidae at Okpilak-Hulahula and Spionidae at Canning. Regardless of the difference in sandpiper diets at the Beaufort Sea deltas, their similar fattening rates throughout the season indicate that all of these stopover sites provide a critical food resource for hatch-year sandpipers beginning their first migration.

Semipalmated Sandpiper (Calidris pusilla) populations have undergone significant declines at core nonbreeding sites in northeastern South America. Breeding populations have also declined in the eastern North American Arctic, but appear to be stable or increasing in the central and western Arctic. To identify vulnerable populations and sites, we documented the migratory connectivity of Semipalmated Sandpipers using light-level geolocators, deploying 250 at 8 Arctic sites across the species' breeding range from 2011 to 2015, plus 87 at a single wintering site in northeastern Brazil in 2013 and 2014. We recovered 59 units and resighted 7 more (26% return rate) on the breeding grounds, but none at the nonbreeding site. We recovered only ∼3% of units deployed in 2013 at eastern Arctic breeding sites, but recovered 33% of those deployed in 2015. Overall, birds with geolocators were 57% as likely to return as those carrying alphanumeric flags. Stopover durations at prairie sites (mean: 8.7 days southbou...

To examine how habitat use by sandpipers (Calidris spp.; Baird's sandpipers, dunlin, least sandpipers, pectoral sandpipers, semipalmated sandpipers, stilt sandpipers, and white-rumped sandpipers) varies across a broad suite of environmental conditions, we conducted surveys at wetlands throughout the spring migratory period in 2013 and 2014 in 2 important stopover regions: the Rainwater Basin (RWB) in Nebraska, USA, and the Prairie Pothole Region (PPR) in South Dakota, USA. Because providing adequate energetic resources for migratory birds is a high priority for wetland management, we also measured invertebrate abundance at managed wetlands in the RWB to determine how food abundance influences the occupancy and abundance of sandpipers on wetlands throughout the migratory period. To quantify habitat use, we surveyed wetlands every 7–10 days in both regions and visually estimated wetland attributes. Our results indicate that invertebrate abundance predicted occupancy, but not abundance, of sandpipers at wetlands in the RWB. The wetland vegetation characteristics that predict sandpiper occupancy are similar in both regions, but wetlands in the PPR support a higher abundance of sandpipers than wetlands in the RWB. Our results suggest that sandpipers make stopover decisions that balance local and regional wetland conditions. Managers should maintain the cues (i.e., mudflat) and ecological conditions beyond invertebrate abundance that predict sandpiper habitat use to successfully provide resources for sandpipers during migratory stopover if that is a goal of wetland management. © 2017 The Wildlife Society.

We compiled a >50‐year record of morphometrics for semipalmated sandpipers (Calidris pusilla), a shorebird species with a Nearctic breeding distribution and intercontinental migration to South America. Our data included >57,000 individuals captured 1972–2015 at five breeding locations and three major stopover sites, plus 139 museum specimens collected in earlier decades. Wing length increased by ca. 1.5 mm (>1%) prior to 1980, followed by a decrease of 3.85 mm (nearly 4%) over the subsequent 35 years. This can account for previously reported changes in metrics at a migratory stopover site from 1985 to 2006. Wing length decreased at a rate of 1,098 darwins, or 0.176 haldanes, within the ranges of other field studies of phenotypic change. Bill length, in contrast, showed no consistent change over the full period of our study. Decreased body size as a universal response of animal populations to climate warming, and several other potential mechanisms, are unable to account for the increasing and decreasing wing length pattern observed. We propose that the post‐WWII near‐extirpation of falcon populations and their post‐1973 recovery driven by the widespread use and subsequent limitation on DDT in North America selected initially for greater flight efficiency and latterly for greater agility. This predation danger hypothesis accounts for many features of the morphometric data and deserves further investigation in this and other species.

1.Animals adjust to seasonal challenges in physical, behavioural and spatial ways. Such adjustments are commonly associated with diet changes that often can be characterised isotopically.2.We introduce the ‘double diet switch model’, with which the occurrence and timing of two subsequent diet switches of an individual animal can be traced with a single sample assayed for stable isotopes. We demonstrate the model for Sanderling, Calidris alba, a small shorebird that migrates from the Nearctic tundra breeding grounds to the intertidal flats of the Wadden Sea; during this migration some birds may stage in the North Atlantic areas.3.The ‘double diet switch model’ successfully predicted the occurrence and timing of two diet switches in 59 Sanderlings captured in the Wadden Sea in July–September. Excluding birds that likely had over-summered at North Atlantic staging areas, the model predicted that Sanderlings departed from the Arctic on 13 July (range: 9–17 July), had a staging duration of 18·6 days in the North Atlantic, and arrived in the Wadden Sea on 1 August (31 July–1 August).The estimated mean Arctic departure dates coincided with the mean hatching date, suggesting that many individuals failed to produce young or left the care to a partner. Estimated mean arrival date matched the main arrival period in the Wadden Sea obtained from observation data. In this study we did not use lipid-free tissues, which may bias model predictions. After correcting for lipid components, the estimated departure date was 11 days later and the staging duration 8·5 days shorter, while arrival date was similar.4.The ‘double diet switch model’ successfully identified the occurrence and timing of two subsequent diet switches. The ‘double diet switch model’ will not only apply to switches between three isotopic levels (as in the case study on Sanderling) but also to scenarios where the second switch reverses to the initial isotopic level. Due to this general applicability, the model can be adapted to a wide range of taxa and situations. Foreseeable applications include changes in habitat and food type, ontogenetic development or drastic phenotypic changes such as the metamorphosis in insects and amphibians

The louse genus Carduiceps Clay & Meinertzhagen, 1939 is widely distributed on sandpipers and stints (Calidrinae). The current taxonomy includes three species on the Calidrinae (Carduiceps meinertzhageni, Carduiceps scalaris, Carduiceps zonarius) and four species on noncalidrine hosts. We estimated a phylogeny of four of the seven species of Carduiceps (the three mentioned above and Carduiceps fulvofasciatus) from 13 of the 29 hosts based on three mitochondrial loci, and evaluated the relative importance of flyway differentiation (same host species has different lice along different flyways) and flyway homogenization (different host species have the same lice along the same flyway). We found no evidence for either process. Instead, the present, morphology-based, taxonomy of the genus corresponds exactly to the gene-based phylogeny, with all four included species monophyletic. Carduiceps zonarius is found both to inhabit a wider range of hosts than wing lice of the genus Lunaceps occurring on the same group of birds, and to occur on Calidris sandpipers of all sizes, both of which are unexpected for a body louse. The previously proposed family Esthiopteridae is found to be monophyletic with good support. The concatenated dataset suggests that the pigeon louse genus Columbicola may be closely related to the auk and diver louse genus Craspedonirmus. These two genera share some morphological characters with Carduiceps, but no support was obtained for grouping these three genera together. Based on mitochondrial data alone, the relationships among genera within this proposed family cannot be properly assessed, but some previously suggested relationships within this proposed family are confirmed.

Peregrine falcons (Falco peregrinus) have undergone a steady hemisphere-wide recovery since the ban on DDT in 1973, resulting in an ongoing increase in the level of danger posed for migrant birds, such as Arctic-breeding sandpipers. We anticipate that in response migrant semipalmated sandpipers (Calidris pusilla) have adjusted migratory behaviour, including a shift in stopover site usage towards locations offering greater safety from falcon predation.We assessed semipalmated sandpiper stopover usage within the Atlantic Canada Shorebird Survey dataset. Based on 3,030 surveys (totalling ∼32M birds) made during southward migration, 1974 - 2017, at 198 stopover locations, we assessed the spatial distribution of site usage in each year (with a ‘priority matching distribution’ index, PMD) in relation to the size (intertidal area) and safety (proportion of a site’s intertidal area further than 150m of the shoreline) of each location. The PMD index value is > 1 when usage is concentrated at dangerous locations, 1.0 when usage matches location size, and < 1 when usage is concentrated at safer locations.A large majority of migrants were found at the safest sites in all years, however our analysis of the PMD demonstrated that the fraction using safer sites increased over time. In 1974, 80% of birds were found at the safest 20% of the sites, while in 2017, this had increased to 97%. A sensitivity analysis shows that the shift was made specifically towards safer (and not just larger) sites. The shift as measured by a PMD index decline cannot be accounted for by possible biases inherent in the data set. We conclude that the data support the prediction that increasing predator danger has induced a shift by southbound migrant semipalmated sandpipers to safer sites.

Louse flies are blood-sucking ectoparasites of birds and mammals of veterinary and medical importance. These flies damage host skin and serve as vectors for blood-associated pathogens. Little is known about their phenology, host selection, and prevalence in wild birds. Our study provided a broad-scale assessment of louse flies parasitizing Charadrii and Scolopaci shorebirds during their migration through Central Europe. We collected louse flies from 3,129 specimens in 22 species of shorebirds at a major migratory stopover site in central Poland in 2011–17. In total, 31 Ornithomya avicularia and 15 Ornithomya chloropus specimens occurred on five wader species: Common Snipe (Gallinago gallinago), Ruff (Calidris pugnax), Dunlin (Calidris alpina), Curlew Sandpiper (Calidris ferruginea), and Wood Sandpiper (Tringa glareola). Prevalences ranged from 0.23% to 4.62%. No individual birds were infested by both species of fly, and no flies were found on an additional 17 wader species. The rarer of the two flies, O. chloropus, occurred on all infested bird species, whereas O. avicularia occurred only on Common Snipe and Curlew Sandpiper. In comparison with the other four infested wader species, Common Snipe (n=1,194) was more frequently infested with O. avicularia than with O. chloropus. Our study shows that the prevalence of louse flies on shorebirds is much lower than on other avian taxa. Ecologic and evolutionary mechanisms leading to the relative rarity of louse flies on shorebirds remain to be understood.

Publisher's version (útgefin grein), Evolutionary theories of seasonal migration generally assume that the costs of longer migrations are balanced by benefits at the non-breeding destinations. We tested, and rejected, the null hypothesis of equal survival and timing of spring migration for High Arctic breeding sanderling Calidris alba using six and eight winter destinations between 55°N and 25°S, respectively. Annual apparent survival was considerably lower for adult birds wintering in tropical West Africa (Mauritania: 0.74 and Ghana: 0.75) than in three European sites (0.84, 0.84 and 0.87) and in subtropical Namibia (0.85). Moreover, compared with adults, second calendar-year sanderlings in the tropics, but not in Europe, often refrained from migrating north during the first possible breeding season. During northward migration, tropical-wintering sanderlings occurred at their final staging site in Iceland 5–15 days later than birds wintering further north or south. Namibia-wintering sanderlings tracked with solar geolocators only staged in West Africa during southward migration. The low annual survival, the later age of first northward migration and the later passage through Iceland during northward migration of tropical-wintering sanderlings, in addition to the skipping of this area during northward but not southward migration by Namibia-wintering sanderlings, all suggest they face issues during the late non-breeding season in West Africa. Migrating sanderlings defy long distances but may end up in winter areas with poor fitness prospects. We suggest that ecological conditions in tropical West Africa make the fuelling prior to northward departure problematic., Annual expeditions to Mauritania were organized by NIOZ, and we especially thank Maarten Brugge, Anne Dekinga, Jutta Leyrer and Bernard Spaans for their contributions. The Parc National du Banc d'Arguin granted research permits and facilitated access. J.R. and T.S.L.V. thank Aarhus University for logistical support at Zackenberg. Benoît Sittler organized expeditions to Karupelv Valley. The Farlington Ringing Group provided cannon‐net equipment. This work was supported by two grants from the Netherlands Polar Programme (851.40.072 and 866.15.207) of the Netherlands Organisation for Scientific research (NWO) and from the Metawad project awarded by Waddenfonds (WF209925) to JR and TP. The measurements in Mauritania had their beginnings in the Prins Bernhard Cultuurfondsprijs to TP. JR and TP also received INTERACT grants for Transnational Access from the European Community's Seventh Framework Programme (grant agreement No262693). JR received a generous donation from World Wildlife Fund Netherlands. JAA was supported by FCT (SFRH/BPD/91527/2012). OG and Loïc Bollache were supported by the French Polar Institute (IPEV; program ‘1036 Interactions') and TL by a Veni grant (no. 016.Veni.192.245) from NWO. The authors declare no conflict of interest. This study is based on the efforts of more than 2,000 observers reporting colour‐ringed sanderlings. We especially thank Guðmundur Örn Benediktsson, John Bowler, Ruth Croger, Anne de Potier, Benjamin Gnep, Kim Fischer, Kirsten Grond, Eileen Hughes, Hilger Lemke, Pedro Lourenço, Andy Johnson, Pierre Leon, Jelle Loonstra, Sebastien Nedellec, Afonso Rocha, Brian Rogers, Ron Summers, Jan van Dijk and Hein Verkade. Anneke Bol, Marco van der Velde and Yvonne Verkuil molecularly sexed the majority of birds, Maria Teixeira and Jérôme Moreau sexed eight individuals. Ron Porter created flags for geolocator attachment. Eldar Rakhimberdiev answered questions concerning FLightR and Allert Bijleveld, Jesse Conklin, Rosemarie Kentie, Thomas Oudman, Janne Ouwehand, Emma Penning, Eldar Rakhimberdiev, Brett Sandercock, Ron Summers, Yvonne Verkuil and two reviewers critically commented on drafts. Benjamin Gnep created Figure.