Summary: There is increasing concern about the world's animal migrations. With many land‐use and climatological changes occurring simultaneously, pinning down the causes of large‐scale conservation problems requires sophisticated and data‐intensive approaches. Declining shorebird numbers along the East Asian–Australasian Flyway, in combination with data on habitat loss along the Yellow Sea (where these birds refuel during long‐distance migrations), indicate a flyway under threat. If habitat loss at staging areas indeed leads to flyway‐wide bird losses, we would predict that: (i) decreases in survival only occur during the season that birds use the Yellow Sea, and (ii) decreases in survival occur in migrants that share a reliance on the vanishing intertidal flats along the Yellow Sea, even if ecologically distinct and using different breeding grounds. Monitored from 2006–2013, we analysedseasonalapparent survival patterns of three shorebird species with non‐overlapping Arctic breeding areas and considerable differences in foraging ecology, but a shared use of both north‐west Australian non‐breeding grounds and the Yellow Sea coasts to refuel during northward and southward migrations (red knotCalidris canutus piersmai, great knotCalidris tenuirostris,bar‐tailed godwitLimosa lapponica menzbieri). Distinguishing two three‐month non‐breeding periods and a six‐month migration and breeding period, and analysing survival of the three species and the three seasons in a single model, we statistically evaluated differences at both the species and season levels. Whereas apparent survival remained high in north‐west Australia, during the time away from the non‐breeding grounds survival in all three species began to decline in 2011, having lost 20 percentage points by 2012. By 2012annualapparent survival had become as low as 0·71 in bar‐tailed godwits, 0·68 in great knots and 0·67 in red knots. In a separate analysis for red knots, no mortality occurred during the migration from Australia to China. In the summers of low summer survival, weather conditions were benign in the Arctic breeding areas. We argue that rapid seashore habitat loss in the Yellow Sea is the most likely explanation of reduced summer survival, with dire (but uncertain) forecasts for the future of these flyway populations. This interpretation is consistent with recent findings of declining shorebird numbers at seemingly intact southern non‐breeding sites. Policy implications. Due to established economic interests, governments are usually reluctant to act for conservation, unless unambiguous evidence for particular cause–effect chains is apparent. This study adds to an increasing body of evidence that habitat loss along the Yellow Sea shores explains the widespread declines in shorebird numbers along the East Asian–Australasian Flyway and threatens the long‐term prospects of several long‐distance migrating species. To halt further losses, the clearance of coastal intertidal habitat must stop now.

Satellite‐based technologies that track individual animal movements enable the mapping of their spatial and temporal patterns of occurrence. This is particularly useful in poorly studied or remote regions where there is a need for the rapid gathering of relevant ecological knowledge to inform management actions. One such region is East Asia, where many intertidal habitats are being degraded at unprecedented rates and shorebird populations relying on these habitats show rapid declines. We examine the utility of satellite tracking to accelerate the identification of coastal sites of conservation importance in the East Asian–Australasian Flyway. In 2015–2017, we used solar‐powered satellite transmitters to track the migration of 32 great knots (Calidris tenuirostris), an “Endangered” shorebird species widely distributed in the Flyway and fully dependent on intertidal habitats for foraging during the non‐breeding season. From the great knot tracks, a total of 92 stopping sites along the Flyway were identified. Surprisingly, 63% of these sites were not known as important shorebird sites before our study; in fact, every one of the tracked individuals used sites that were previously unrecognized. Site knowledge from on‐ground studies in the Flyway is most complete for the Yellow Sea and generally lacking for Southeast Asia, Southern China and Eastern Russia. Synthesis and applications. Satellite tracking highlighted coastal habitats that are potentially important for shorebirds but lack ecological information and conservation recognition, such as those in Southern China and Southeast Asia. At the same time, the distributional data of tracked individuals can direct on‐ground surveys at the lesser known sites to collect information on bird numbers and habitat characteristics. To recognize and subsequently protect valuable coastal habitats, filling knowledge gaps by integrating bird tracking with ground‐based methods should be prioritized.

Summary: Many shorebird populations are in decline along the East Asian-Australasian Flyway. The rapid loss of coastal wetlands in the Yellow Sea, which provide critical stop-over sites during migration, is believed to be the cause of the alarming trends. The Yalu Jiang coastal wetland, a protected area in the north Yellow Sea, supports the largest known migratory staging populations of Bar-tailed GodwitsLimosa lapponica(menzbieriandbauerisubspecies) and Great KnotsCalidris tenuirostris. Monitoring of the macrozoobenthos food for these shorebirds from 2011 to 2016 showed declines of over 99% in the densities of the bivalvePotamocorbula laevis, the major food here for both Bar-tailed Godwits and Great Knots. The loss of the bivalve might be caused by any combination of, but not limited to: (1) change in hydrological conditions and sediment composition due to nearby port construction, (2) run-off of agrochemicals from the extensive shoreline sea cucumber farms, and (3) parasitic infection. Surprisingly, the numbers of birds using the Yalu Jiang coastal wetland remained stable during the study period, except for the subspecies of Bar-tailed GodwitL. l. menzbieri, which exhibited a 91% decline in peak numbers. The lack of an overall decline in the number of bird days in Great Knots and in the peak numbers ofL. l. baueri, also given the published simultaneous decreases in their annual survival, implies a lack of alternative habitats that birds could relocate to. This study highlights that food declines at staging sites could be an overlooked but important factor causing population declines of shorebirds along the Flyway. Maintaining the quality of protected staging sites is as important in shorebird conservation as is the safeguarding of staging sites from land claim. Meanwhile, it calls for immediate action to restore the food base for these beleaguered migrant shorebirds at Yalu Jiang coastal wetland.

Negative density-dependence is generally studied within a single trophic level, thereby neglecting its effect on higher trophic levels. The ‘functional response’ couples a predator's intake rate to prey density. Most widespread is a type II functional response, where intake rate increases asymptotically with prey density; this predicts the highest predator densities at the highest prey densities. In one of the most stringent tests of this generality to date, we measured density and quality of bivalve prey (edible cocklesCerastoderma edule) across 50 km² of mudflat, and simultaneously, with a novel time-of-arrival methodology, tracked their avian predators (red knotsCalidris canutus). Because of negative density-dependence in the individual quality of cockles, the predicted energy intake rates of red knots declined at high prey densities (a type IV, rather than a type II functional response). Resource-selection modelling revealed that red knots indeed selected areas of intermediate cockle densities where energy intake rates were maximized given their phenotype-specific digestive constraints (as indicated by gizzard mass). Because negative density-dependence is common, we question the current consensus and suggest that predators commonly maximize their energy intake rates at intermediate prey densities. Prey density alone may thus poorly predict intake rates, carrying capacity and spatial distributions of predators.

Organisms cope with environmental stressors by behavioral, morphological, and physiological adjustments. Documentation of such adjustments in the wild provides information on the response space in nature and the extent to which behavioral and bodily adjustments lead to appropriate performance effects. Here we studied the morphological and digestive adjustments in a staging population of migrating Great Knots Calidris tenuirostris in response to stark declines in food abundance and quality at the Yalu Jiang estuarine wetland (northern Yellow Sea, China). At Yalu Jiang, from 2011 to 2017 the densities of intertidal mollusks, the food of Great Knots, declined 15‐fold. The staple prey of Great Knots shifted from the relatively soft‐shelled bivalve Potamocorbula laevis in 2011–2012 to harder‐shelled mollusks such as the gastropod Umbonium thomasi in 2016–2017. The crushing of the mollusks in the gizzard would require a threefold to 11‐fold increase in break force. This was partially resolved by a 15% increase in gizzard mass which would yield a 32% increase in shell processing capacity. The consumption of harder‐shelled mollusks was also accompanied by reliance on regurgitates to excrete unbreakable parts of prey, rather than the usual intestinal voidance of shell fragments as feces. Despite the changes in digestive morphology and strategy, there was still an 85% reduction in intake rate in 2016–2017 compared with 2011–2012. With these morphological and digestive adjustments, the Great Knots remaining faithful to the staging site to a certain extent buffered the disadvantageous effects of dramatic food declines. However, compensation was not complete. Locally, birds will have had to extend foraging time and use a greater daily foraging range. This study offers a perspective on how individual animals may mitigate the effects of environmental change by morphological and digestive strategies and the limits to the response space of long‐distance migrating shorebirds in the wild.