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Most processes within organisms, and most interactions between organisms and their environment, have distinct time profiles. The temporal coordination of such processes is crucial across levels of biological organization, but disciplines differ widely in their approaches to study timing. Such differences are accentuated between ecologists, who are centrally concerned with a holistic view of an organism in relation to its external environment, and chronobiologists, who emphasize internal timekeeping within an organism and the mechanisms of its adjustment to the environment. We argue that ecological and chronobiological perspectives are complementary, and that studies at the intersection will enable both fields to jointly overcome obstacles that currently hinder progress. However, to achieve this integration, we first have to cross some conceptual barriers, clarifying prohibitively inaccessible terminologies. We critically assess main assumptions and concepts in either field, as well as their common interests. Both approaches intersect in their need to understand the extent and regulation of temporal plasticity, and in the concept of ‘chronotype’, i.e. the characteristic temporal properties of individuals which are the targets of natural and sexual selection. We then highlight promising developments, point out open questions, acknowledge difficulties and propose directions for further integration of ecological and chronobiological perspectives through Wild Clock research. This article is part of the themed issue ‘Wild Clocks: integrating chronobiology and ecology to understand timekeeping in free-living animals’.

Marine organisms adapt to complex temporal environments that include daily, tidal, semi-lunar, lunar and seasonal cycles. However, our understanding of marine biological rhythms and their underlying molecular basis is mainly confined to a few model organisms in rather simplistic laboratory settings. Here, we use new empirical data and recent examples of marine biorhythms to highlight how field ecologists and laboratory chronobiologists can complement each other's efforts. First, with continuous tracking of intertidal shorebirds in the field, we reveal individual differences in tidal and circadian foraging rhythms. Second, we demonstrate that shorebird species that spend 8–10 months in tidal environments rarely maintain such tidal or circadian rhythms during breeding, likely because of other, more pertinent, temporally structured, local ecological pressures such as predation or social environment. Finally, we use examples of initial findings from invertebrates (arthropods and polychaete worms) that are being developed as model species to study the molecular bases of lunar-related rhythms. These examples indicate that canonical circadian clock genes (i.e. the homologous clock genes identified in many higher organisms) may not be involved in lunar/tidal phenotypes. Together, our results and the examples we describe emphasize that linking field and laboratory studies is likely to generate a better ecological appreciation of lunar-related rhythms in the wild. This article is part of the themed issue ‘Wild clocks: integrating chronobiology and ecology to understand timekeeping in free-living animals’.

Reductions in body size are increasingly being identified as a response to climate warming. Here we present evidence for a case of such body shrinkage, potentially due to malnutrition in early life. We show that an avian long-distance migrant (red knot, Calidris canutus canutus), which is experiencing globally unrivaled warming rates at its high-Arctic breeding grounds, produces smaller offspring with shorter bills during summers with early snowmelt. This has consequences half a world away at their tropical wintering grounds, where shorter-billed individuals have reduced survival rates. This is associated with these molluscivores eating fewer deeply buried bivalve prey and more shallowly buried seagrass rhizomes. We suggest that seasonal migrants can experience reduced fitness at one end of their range as a result of a changing climate at the other end.

Upon settlement after a pelagic larval phase, brown shrimp Crangon crangon depend on intertidal flats. During low as well as high tide the young brown shrimp play roles as predators of meiofauna and as prey for fish and birds. Unlike the biology of the commercially important adults, knowledge on these juveniles remains sketchy. Here we provide an analysis of 35 years (1984–2018) of brown shrimp monitoring in May–June on intertidal flats in the westernmost Dutch Wadden Sea. Intertidal shrimp densities were sampled bi-weekly at three stations during low tide, using sampling corers. We show that over this 35-year period the appearance of shrimp on mudflats advanced by 12 days (− 0.34 days yr−1). Simultaneously, densities on 7 May increased by more than 2.4 times, from 28 shrimp m−2 in 1984 to 69 shrimp m−2 in 2018. Across years, mean shrimp length decreased from 12.6 to 10.7 mm, but length in early May did not change. The advancement in settlement and the increasing shrimp densities correlated with increases in the seawater temperatures in April more than during earlier times of the year. We propose four interpretations of these changes: (1) shrimp settle on the mudflat when they reach a certain ‘threshold’ length, (2) settlement of shrimp is controlled by a critical period of ‘threshold’ temperature sensitivity, (3) timing of shrimp settlement is a response to food availability on mudflats or (4) a direct response to inferred predation pressure. The different interpretations will lead to different scenarios of change in a warming world.

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 analysed seasonal apparent 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 knot Calidris canutus piersmai, great knot Calidris tenuirostris, bar‐tailed godwit Limosa 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 2012 annual apparent 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.

For almost a century, the term ‘flyways’ has been used to order relations over time and space. It has been used to coordinate scientific research and communication as well as monitoring and management efforts for waterbird conservation. In this article, we revisit the concept of ‘boundary object’ (Star and Griesemer 1989) to investigate how this term ‘flyways’ has been central to common efforts while also having multiple meanings for the actors it connects. The article discusses both contemporary and historical achievements of the term by analysing its underlying knowledge infrastructure. We account for the complex assemblages of social, material, natural, and technical systems that shape how the term ‘flyway’ has been functioning as a boundary object and how this has changed over time. By discussing how the term ‘flyways’ as a boundary object and its underlying knowledge infrastructure shape each other, we empower the actors to define, visualise, communicate, and imagine flyways in more purposeful ways. Our analysis contributes to the literature on boundary objects and knowledge infrastructures by expanding their original definitions, arguing for a co-productive relation between them.

Ground‐nesting shorebirds must balance the need for acoustic communication at the nest with the constant threat posed by predators. Although it may seem likely that their calls are adapted to minimize detection by predators, little is known about how these birds communicate at the nest or whether they employ cryptic strategies to avoid predation. Using passive acoustic devices and software to analyse extensive acoustic data, we quantified and categorised the calls of black‐tailed godwits Limosa limosa limosa recorded throughout the whole incubation at eight nests at a dairy farm in the Netherlands in March–June 2021. While incubating, godwits frequently use five main call types, with distinct diurnal patterns and high variation in the number of calls between breeding pairs. Birds used two quiet calls, one for communication at the nest and a second without an easily suggested meaning. Three loud calls were presumably used for predator alert, territory establishment, and long‐range communication. Interestingly, although nests were close to each other and exposed to the same aerial predators, the involvement of incubating birds in predator alert calling consistently differed. Furthermore, we described the relationship between the number of predator alert calls and the probability of a godwit flying off the nest. Our findings show that incubating godwits predominantly use loud vocalizations during the day, with only a few calls at night, which were more frequent on nights with a full moon. These descriptive findings for a single godwit community should now be expanded to other contexts, experimental situations, and shorebird species.

Nearly 20% of all bird species migrate between breeding and nonbreeding sites annually. Their migrations include storied feats of endurance and physiology, from non-stop trans-Pacific crossings to flights at the cruising altitudes of jetliners. Despite intense interest in these performances, there remains great uncertainty about which factors most directly influence bird behaviour during migratory flights. We used GPS trackers that measure an individual's altitude and wingbeat frequency to track the migration of black-tailed godwits ( Limosa limosa ) and identify the abiotic factors influencing their in-flight migratory behaviour. We found that godwits flew at altitudes above 5000 m during 21% of all migratory flights, and reached maximum flight altitudes of nearly 6000 m. The partial pressure of oxygen at these altitudes is less than 50% of that at sea level, yet these extremely high flights occurred in the absence of topographical barriers. Instead, they were associated with high air temperatures at lower altitudes and increasing wind support at higher altitudes. Our results therefore suggest that wind, temperature and topography all play a role in determining migratory behaviour, but that their relative importance is context dependent. Extremely high-altitude flights may thus not be especially rare, but they may only occur in very specific environmental contexts.

With its focus on wetlands, the Ramsar Convention provides the clearest global agreement helping the conservation of migratory waterbirds. Two specific criteria (5 and 6) support the scientific basis for sites to achieve Ramsar recognition based on waterbird counts, while criterion 4, on species and ecological communities, also plays a role. Other international conventions and agreements follow these criteria. We identify several reasons why the listing thus established can only “catch” the absolute minimum wetland network for the conservation of migratory waterbirds. We argue that individual tracking and modern observational tools allow to better delineate the areas needed to effectively give migratory waterbird populations full life cycle protection. The sophisticated techniques to measure population characteristics now available should be used to modernize the guidance for the application of Criteria 4 and 6 of the Ramsar Convention for waterbirds, based on (i) time spent in a site throughout migration; (ii) critical (“untouchable”) sites; (iii) robustness of designated site network including buffer areas; (iv) full life cycle information—including early life phases; and (v) refuges used on‐and‐off during migration in emergency situations. In these enhanced ways, migratory waterbirds can enact their roles as effective sentinels of the ecological state of the world.

Long-distance migration, and the study of the migrants who undertake these journeys, has fascinated generations of biologists. However, many aspects of the annual cycles of these migrants remain a mystery as do many of the driving forces behind the evolution and maintenance of the migrations themselves. In this article we discuss nutritional, energetic, temporal and disease-risk bottlenecks in the annual cycle of long-distance migrants, taking a sandpiper, the red knot Calidris canutus, as a focal species. Red knots have six recognized subspecies each with different migratory routes, well-known patterns of connectivity and contrasting annual cycles. The diversity of red knot annual cycles allows us to discuss the existence and the effects of bottlenecks in a comparative framework. We examine the evidence for bottlenecks focusing on the quality of breeding plumage and the timing of moult as indicators in the six subspecies. In terms of breeding plumage coloration, quality and timing of prealternate body moult (from non-breeding into breeding plumage), the longest migrating knot subspecies, Calidris canutus rogersi and Calidris canutus rufa, show the greatest impact of bottlenecking. The same is true in terms of prebasic body moult (from breeding into non-breeding plumage) which in case of both C. c. rogersi and C. c. rufa overlaps with southward migration and may even commence in the breeding grounds. To close our discussion of bottlenecks in long-distance migrants, we make predictions about how migrants might be impacted via physiological 'trade-offs' throughout the annual cycle, using investment in immune function as an example. We also predict how bottlenecks may affect the distribution of mortality throughout the annual cycle. We hope that this framework will be applicable to other species and types of migrants, thus expanding the comparative database for the future evaluation of seasonal selection pressures and the evolution of annual cycles in long-distance migrants. Furthermore, we hope that this synthesis of recent advancements in the knowledge of red knot annual cycles will prove useful in the ongoing attempts to model annual cycles in migratory birds.