In migratory birds, high mobility may reduce population structure through increased dispersal and enable adaptive responses to environmental change, whereas rigid migratory routines predict low dispersal, increased geographic structure, and limited flexibility to respond to change. We used nextRAD sequencing of 14,318 single-nucleotide polymorphisms to explore the population genetics and phylogeographic history of the bar-tailed godwit, Limosa lapponica , a migratory shorebird with six recognized subspecies and known for making the longest non-stop flights of any landbird. Using scenario-testing in an Approximate Bayesian Computation framework, we infer that bar-tailed godwits existed in three main lineages at the Last Glacial Maximum (LGM), when much of their present-day Arctic and sub-Arctic breeding range persisted in a large, unglaciated Siberian-Beringian refugium. Subsequently, population structure developed at both longitudinal extremes: in the east, a genetic cline exists across latitude in the Alaska breeding range of subspecies L. l. baueri ; in the west, one lineage diversified into three extant subspecies L. l. lapponica , taymyrensis , and yamalensis , the former two of which migrate through previously glaciated western Europe. We also detected unrecognized population structure among bar-tailed godwits wintering in Europe, wherein a significant proportion of purported lapponica individuals were in fact taymyrensis , necessitating a re-assessment of the migrations, ecology, and population estimates for these subspecies. In the global range of this long-distance migrant, we found evidence of both (1) fidelity to rigid behavioral routines promoting fine-scale geographic population structure (in the east), and (2) flexibility to colonize recently available migratory flyways and non-breeding areas (in the west).### Competing Interest StatementThe authors have declared no competing interest.

In migratory animals, high mobility may reduce population structure through increased dispersal and enable adaptive responses to environmental change, whereas rigid migratory routines predict low dispersal, increased structure, and limited flexibility to respond to change. We explore the global population structure and phylogeographic history of the bar-tailed godwit, Limosa lapponica, a migratory shorebird known for making the longest non-stop flights of any landbird. Using nextRAD sequencing of 14,318 single-nucleotide polymorphisms and scenario-testing in an Approximate Bayesian Computation framework, we infer that bar-tailed godwits existed in two main lineages at the last glacial maximum, when much of their present-day breeding range persisted in a vast, unglaciated Siberian-Beringian refugium, followed by admixture of these lineages in the eastern Palearctic. Subsequently, population structure developed at both longitudinal extremes: in the east, a genetic cline exists across latitude in the Alaska breeding range of subspecies L. l. baueri; in the west, one lineage diversified into three extant subspecies L. l. lapponica, taymyrensis, and yamalensis, the former two of which migrate through previously glaciated western Europe. In the global range of this long-distance migrant, we found evidence of both (1) fidelity to rigid behavioural routines promoting fine-scale geographic population structure (in the east) and (2) flexibility to colonise recently available migratory flyways and non-breeding areas (in the west). Our results suggest that cultural traditions in highly mobile vertebrates can override the expected effects of high dispersal ability on population structure, and provide insights for the evolution and flexibility of some of the world's longest migrations.

The Pacific Basin, by virtue of its vastness and its complex aeroscape, provides unique opportunities to address questions about the behavioral and physiological capabilities and mechanisms through which birds can complete spectacular flights. No longer is the Pacific seen just as a formidable barrier between terrestrial habitats in the north and the south, but rather as a gateway for specialized species, such as shorebirds, to make a living on hemispherically distributed seasonal resources. This recent change in perspective is dramatic, and the research that underpins it has presented new opportunities to learn about phenomena that often challenge a sense of normal. Ancient Polynesians were aware of the seasonal passage of shorebirds and other landbirds over the Pacific Ocean, incorporating these observations into their navigational “tool kit” as they explored and colonized the Pacific. Some ten centuries later, systematic visual observations and tracking technology have revealed much about movement of these shorebirds, especially the enormity of their individual nonstop flights. This invites a broad suite of questions, often requiring comparative studies with bird migration across other ocean basins, or across continents. For example, how do birds manage many days of nonstop exercise apparently without sleep? What mechanisms explain birds acting as if they possess a Global Positioning System? How do such extreme migrations evolve? Through advances in both theory and tracking technology, biologists are poised to greatly expand the horizons of movement ecology as we know it. In this integrative review, we present a series of intriguing questions about trans-Pacific migrant shorebirds and summarize recent advances in knowledge about migratory behavior operating at temporal scales ranging from immediate decisions during a single flight, to adaptive learning throughout a lifetime, to evolutionary development of migratory pathways. Recent advances in this realm should stimulate future research across the globe and across a broad array of disciplines.

In seasonal environments subject to climate change, organisms typically show phenological changes. As these changes are usually stronger in organisms at lower trophic levels than those at higher trophic levels, mismatches between consumers and their prey may occur during the consumers' reproduction period. While in some species a trophic mismatch induces reductions in offspring growth, this is not always the case. This variation may be caused by the relative strength of the mismatch, or by mitigating factors like increased temperature-reducing energetic costs. We investigated the response of chick growth rate to arthropod abundance and temperature for six populations of ecologically similar shorebirds breeding in the Arctic and sub-Arctic (four subspecies of Red Knot Calidris canutus, Great Knot C. tenuirostris and Surfbird C. virgata). In general, chicks experienced growth benefits (measured as a condition index) when hatching before the seasonal peak in arthropod abundance, and growth reductions when hatching after the peak. The moment in the season at which growth reductions occurred varied between populations, likely depending on whether food was limiting growth before or after the peak. Higher temperatures led to faster growth on average, but could only compensate for increasing trophic mismatch for the population experiencing the coldest conditions. We did not find changes in the timing of peaks in arthropod availability across the study years, possibly because our series of observations was relatively short; timing of hatching displayed no change over the years either. Our results suggest that a trend in trophic mismatches may not yet be evident; however, we show Arctic-breeding shorebirds to be vulnerable to this phenomenon and vulnerability to depend on seasonal prey dynamics.

In a 1998 paper entitled “Guts don’t fly: small digestive organs in obese bar-tailed godwits,” Piersma and Gill (1998) showed that the digestive organs were tiny and the fat loads huge in individuals suspected of embarking on a non-stop flight from Alaska to New Zealand. It was suggested that prior to migratory departure, these godwits would shrink the digestive organs used during fuel deposition and boost the size and capacity of exercise organs to optimize flight performance. Here we document the verity of the proposed physiomorphic changes by comparing organ sizes and body composition of bar-tailed godwits Limosa lapponica baueri collected in modesty midway during their fueling period (mid-September; fueling, n = 7) with the previously published data for godwits that had just departed on their trans-Pacific flight (October 19; flying, n = 9). Mean total body masses for the two groups were nearly identical, but nearly half of the body mass of fueling godwits consisted of water, while fat constituted over half of total body mass of flying godwits. The two groups also differed in their fat-free mass components. The heart and flight muscles were heavier in fueling godwits, but these body components constituted a relatively greater fraction of the fat-free mass in flying godwits. In contrast, organs related to digestion and homeostasis were heavier in fueling godwits, and most of these organ groups were also relatively larger in fueling godwits compared to flying godwits. These results reflect the functional importance of organ and muscle groups related to energy acquisition in fueling godwits and the consequences of flight-related exertion in flying godwits. The extreme physiomorphic changes apparently occurred over a short time window (≤1 month). We conclude that the inferences made on the basis of the 1998 paper were correct. The cues and stimuli which moderate these changes remain to be studied.

Birds that conduct long-distance migrations exhibit varied patterns of consistency in migratory timing and site use. Understanding variation in these traits among populations can help uncover mechanisms driving migratory behaviors and identify potential population threats. Whimbrels (Numenius phaeopus) are long-distance migratory shorebirds with a Holarctic breeding distribution, and recent studies have documented population-specific migrations that vary in duration (short to long) and frequency of stops (none/few to multiple). Factors driving these population-specific differences are unclear. We studied the migration ecology of Whimbrels breeding in Alaska, USA, using satellite transmitters deployed from 2006 to 2010 and tracked through 2015. Whimbrels moved entirely within the Pacific Americas Flyway, and some conducted nonstop flights that exceeded seven days across similar to 8700 km. Birds dispersed across numerous sites throughout the flyway, often using agriculture or aquaculture habitats. Whimbrels generally exhibited fidelity to breeding and non-breeding sites, but typically only exhibited fidelity to staging sites used prior to long, nonstop migratory flights. The duration of migration for Whimbrels at more southern non-breeding locations was longer than for those at more northern non-breeding sites, and birds at more southern sites also terminated southbound migration later and initiated northbound migration earlier than birds at more northern sites. Alaska-breeding Whimbrels exhibited greater variation in migratory behaviors than those in other populations in the species' range. We attribute this within-population diversity to the extensive breadth of non-breeding distributions (similar to 70 degrees latitude across similar to 8600 km), a range that naturally shaped individual responses to unequal migratory demands.

In June and July 2018 and July 2019 we surveyed birds on St. Matthew and Hall islands, isolated in the central Bering Sea. Our surveys were focused on the McKay’s Bunting (Plectrophenax hyperboreus), Rock Sandpiper (Calidris ptilocnemis ptilocnemis), and Pelagic Cormorant (Phalacrocorax pelagicus) but encompassed all birds and yielded 13 species and four subspecies new to the islands’ avifaunal list, and new details and documentation of breeding for eight species. Especially notable discoveries include a mixed pair of Bewick’s (Cygnus columbianus bewickii) and Whistling (C. c. columbianus) Swans, a mixed pair of the Glaucous-winged (Larus glaucescens) and an Asian subspecies of the Herring Gull (L. argentatus vegae), and discovery of a colony of ~100 pairs of the Red-legged Kittiwake (Rissa brevirostris). Also, contrary to many previous expeditions, we found the Gray-crowned Rosy-Finch (Leucosticte tephrocotis umbrina) to be common and conclude that it may be resident. The St. Matthew Island group, including St. Matthew, Hall, and Pinnacle islands, is located in the central Bering Sea (60° 24ʹ N, 172° 42ʹ W; Figure 1), and is part of the Alaska Maritime National Wildlife Refuge (see Winker et al. 2002 for physical descriptions of the islands and details of their habitats).

ABSTRACT The study of avian movement has detailed a spectrum of strategies for the timing and use of sites throughout the annual cycle, from near randomness to complete consistency. New tracking devices now permit the repeated tracking of individual animals throughout the annual cycle, detailing previously unappreciated levels of variation within migratory systems. Godwits (genus Limosa) have featured prominently in studies of avian migration, but information derived from repeated tracking of individuals is limited. The Marbled Godwit subspecies Limosa fedoa beringiae breeds on the central Alaska Peninsula, and little is known about basic aspects of its migration ecology, including the repeatability with which this population times its annual migratory movements or uses migratory and nonbreeding sites. To address these questions, we equipped 9 Marbled Godwits breeding at a site near Ugashik, Alaska, with solar-powered satellite transmitters. We tracked individuals from July, 2008 to October, 2015 and obtained repeat migratory tracks from 5 of these birds. Individuals exhibited high fidelity to breeding, nonbreeding, and migratory stopover sites across years, but in contrast to congeners that conduct consistently timed, long, nonstop migrations, beringiae Marbled Godwits exhibited low levels of individual- or population-level repeatability in the timing of migratory movements. Their relatively short migrations may enable the integration of local environmental cues, potentially facilitating individual flexibility in the timing of annual migratory movements. Curiously, if local cues ultimately drive the timing of Marbled Godwit migratory movements, the population's relatively constrained distribution during both the breeding and nonbreeding season should serve to synchronize birds if they are responding to similar cues. That our sample of Marbled Godwits nonetheless exhibited within- and between-year variation in the timing of their migratory movements suggests a complex integration of annually variable internal and external cues.

Rock Sandpipers Calidris ptilocnemis have the most northerly non‐breeding distribution of any shorebird in the Pacific Basin (upper Cook Inlet, Alaska; 61°N, 151°W). In terms of freezing temperatures, persistent winds and pervasive ice, this site is the harshest used by shorebirds during winter. We integrated physiological, metabolic, behavioural and environmental aspects of the non‐breeding ecology of Rock Sandpipers at the northern extent of their range to determine the relative importance of these factors in facilitating their unique non‐breeding ecology. Not surprisingly, estimated daily energetic demands were greatest during January, the coldest period of winter. These estimates were greatest for foraging birds, and exceeded basal metabolic rates by a factor of 6.5, a scope of increase that approaches the maximum sustained rate of energetic output by shorebirds during periods of migration, but far exceeds these periods in duration. We assessed the quality of their primary prey, the bivalve Macoma balthica , to determine the daily foraging duration required by Rock Sandpipers to satisfy such energetic demands. Based on size‐specific estimates of M. balthica quality, Rock Sandpipers require over 13 h/day of foraging time in upper Cook Inlet in January, even when feeding on the highest quality prey. This range approaches the average daily duration of mudflat availability in this region ( c . 18 h), a maximum value that annually decreases due to the accumulation of shore‐fast ice. Rock Sandpipers are likely to maximize access to foraging sites by following the exposure of ice‐free mudflats across the upper Cook Inlet region and by selecting smaller, higher quality M. balthica to minimize foraging times. Ultimately, this unusual non‐breeding ecology relies on the high quality of their prey resources. Compared with other sites across their range, M. balthica from upper Cook Inlet have relatively light shells, potentially the result of the region's depauperate invertebrate predator community. Given the delicate balance between environmental and prey conditions that currently make Cook Inlet a viable wintering area for Rock Sandpipers, small variations in these variables may affect the suitability of the site in the future.

Between 2004 and 2008, biologists conducted an inventory of breeding birds during May-June primarily in montane areas (>100 m above sea level) of Aniakchak National Monument and Preserve (Aniakchak NMP), Katmai National Park and Preserve (Katmai NPP), and Lake Clark National Park and Preserve (Lake Clark NPP) in southwestern Alaska. Observers conducted 1,021 point counts along 169 transects within 63 10-km X 10-km plots that were randomly selected and stratified by ecological subsection. We created hierarchical N-mixture models to estimate detection probability and abundance for 15 species, including 12 passerines, 2 galliforms, and 1 shorebird. We first modeled detection probability relative to observer, date within season, and proportion of dense vegetation cover around the point, then modeled abundance as a function of land cover composition (proportion of seven coarse-scale land cover types) within 300 m of the survey point. Land cover relationships varied widely among species but most showed selection for low to tall shrubs (0.2-5 m tall) and an avoidance of alpine and dwarf shrub-herbaceous cover types. After adjusting for species not observed, we estimated a minimum of 107 +/- 9 species bred in the areas surveyed within the three parks combined. Species richness was negatively associated with elevation and associated land cover types. At comparable levels of survey effort (n = 721 birds detected), species richness was greatest in Lake Clark NPP (75 +/- 12 species), lowest in Aniakchak NMP (45 +/- 6 species), and intermediate at Katmai NPP (59 +/- 10 species). Species richness was similar at equivalent survey effort (n = 973 birds detected) within the Lime Hills, Alaska Range, and Alaska Peninsula ecoregions (68 +/- 8; 79 +/- 11; 67 +/- 11, respectively). Species composition was similar across all three parks and across the three major ecoregions (Alaska Range, Alaska Peninsula, Lime Hills) that encompass them. Our results provide baseline estimates of relative abundance and models of abundance and species richness relative to land cover that can be used to assess future changes in avian distribution. Additionally, these subarctic montane parks may serve as signals of landscape change and barometers for the assessment of population and distributional changes as a result of warming temperatures and changing precipitation patterns.

The fourth report of the Alaska Checklist Committee outlines 22 species and two subspecies added to the Checklist of Alaska Birds during the five years 2013–2017, resulting in a net total of 521 species and 119 additional subspecies of birds recognized as occurring or having occurred naturally in Alaska through 2017. During the period 2013–2017, 22 species and two subspecies were added to the Checklist of Alaska Birds, resulting in a total of 521 species and 119 additional subspecies recognized by the Alaska Checklist Committee (AKCLC) at the beginning of 2018 as occurring or having occurred naturally in Alaska. Organized in 2000, the committee comprises six voting members (above) and a nonvoting secretary (Withrow). We post a new edition of the Checklist of Alaska Birds early in each new year at the University of Alaska Museum’s website, where the 24th edition (dated 1 January 2018) can be found at . The current Checklist of Alaska Birds is founded on the “Inventory of the species and subspecies of Alaska birds” (Gibson and Kessel 1997) and incorporates additions, deletions, status changes, taxonomic and nomenclatural changes, etc., reflected in earlier AKCLC reports (Gibson et al. 2003, 2008, 2013), in the second edition of the “Inventory” (Gibson and Withrow 2015), and on information we report here for the first time. Volume 49, Number 3, 2018

Arctic summers are brief, and there has been strong selection for migratory birds to arrive in Arctic nesting areas as early as possible to time breeding with peak food availability and complete reproduction. The timing of emergence of nesting habitat in spring is, however, extremely variable in the Arctic, and few long-term studies have examined the ability of avian migrants to track spring conditions to assure optimal nesting. Such studies require long-term migration monitoring under variable spring habitat conditions. These conditions were met during our long-term (1977–2008) study of the timing of arrival of shorebirds to their nesting grounds on the central Yukon–Kuskokwim (Y-K) Delta, in western Alaska. Over this period, the timing of arrival on the nesting grounds of 12 species of shorebirds varied significantly, with the Black-bellied Plover (Pluvialis squatarola) generally arriving first (mean arrival of 4 May), the Red Phalarope (Phalaropus fulicarius) usually arriving last (mean arrival of 20 May). The Western Sandpiper (Calidris mauri), Dunlin (Calidris alpina), and Red-necked Phalarope (Phalaropus lobatus), the most common breeding shorebird species we studied, all arrived about the same time each year (7–9 May). From year to year, first arrival of all species varied significantly by more than 2 weeks, but there was no long-term trend in arrival times over the length of our study. Shorebird arrival was highly correlated with the timing of the break-up of ice on the Kashunuk River, which in turn was correlated with decreasing snow cover and increasing ambient temperature. The date of break-up of river ice also varied by year but did not advance significantly during our study. After arriving on the breeding grounds, Arctic-nesting shorebirds rely on local food resources, which on the Y-K Delta they achieve by timing their arrival to coincide with availability of snow-free habitat. Temperatures along the terminal portions of the spring migration route were significantly correlated with both conditions on the breeding ground and the timing of shorebirds’ arrival there, which suggests that shorebirds may use environmental cues during spring migration to regulate its pace. Ours is one of the few multi-decadal studies to evaluate the responses of a suite of migrant species to annual variation in conditions in their Arctic breeding habitat. Shorebirds’ adaptations to variable conditions on the Y-K Delta are probably rooted in frequent changes to the landscape since the last glacial maximum. Such inherent flexibility may serve them well under future scenarios associated with a changing climate.

At what phenotypic level do closely related subspecies that live in different environments differ with respect to food detection, ingestion, and processing? This question motivated an experimental study on rock sandpipers (Calidris ptilocnemis). The species’ nonbreeding range spans 20 degrees of latitude, the extremes of which are inhabited by two subspecies: Calidris p. ptilocnemis that winters primarily in upper Cook Inlet, Alaska (61°N), and C. p. tschuktschorum that overlaps slightly with C. p. ptilocnemis but whose range extends much farther south (∼40°N). In view of the strongly contrasting energetic demands of their distinct nonbreeding distributions, we conducted experiments to assess the behavioural, physiological, and sensory aspects of foraging, and we used the bivalve Macoma balthica for all trials. Ptilocnemis consumed a wider range of prey sizes, had higher maximum rates of energy intake, processed shell waste at higher maximum rates, and handled prey more quickly. Notably, however, the two subspecies did not differ in their abilities to find buried prey. The subspecies were similar in size and had equally sized gizzards, but the more northern ptilocnemis individuals were 10–14% heavier than their same-sex tschuktschorum counterparts. The higher body mass in ptilocnemis likely resulted from hypertrophy of digestive organs (e.g. intestine, liver) related to digestion and nutrient assimilation. Given the previously established equality of the two subspecies’ metabolic capacities, we propose that the high-latitude nonbreeding range of ptilocnemis rock sandpipers is primarily facilitated by digestive (i.e. physiological) aspects of their foraging ecology rather than behavioural or sensory aspects.

In 1918, the esteemed North American naturalist Joseph Grinnell, in describing the state of knowledge of the Wandering Tattler Tringa incana Gmelin, 1789, mentioned “... it will probably be a matter of but a few years before full knowledge of its nesting habits is obtained” (Grinnell et al. 1918). Almost a century later the breeding behavior of this species arguably remained the most poorly known of any shorebird in North America, owing largely to a suite of peculiar natural history traits that have challenged biologists since the species was first described.

Migratory birds occupy geographically and ecologically disparate areas during their annual cycle with conditions on breeding and non-breeding grounds playing separate and important roles in population dynamics. We used data from nuclear microsatellite and mitochondrial DNA control region loci to assess the breeding and non-breeding spatial genetic structure of a transoceanic migrant shorebird, the bristle-thighed curlew. We found spatial variance in the distribution of allelic and haplotypic frequencies between the curlew’s two breeding areas in Alaska but did not observe this spatial structure throughout its non-breeding range on low-lying tropical and subtropical islands in the Central Pacific (Oceania). This suggests that the two breeding populations do not spatially segregate during the non-breeding season. Lack of migratory connectivity is likely attributable to the species’ behavior, as bristle-thighed curlews exhibit differential timing of migration and some individuals move among islands during non-breeding months. Given the detrimental impact of many past and current human activities on island ecosystems, admixture of breeding populations in Oceania may render the bristle-thighed curlew less vulnerable to perturbations there, as neither breeding population will be disproportionally affected by local habitat losses or by stochastic events. Furthermore, lack of migratory connectivity may enable bristle-thighed curlews to respond to changing island ecosystems by altering their non-breeding distribution. However, availability of suitable non-breeding habitat for curlews in Oceania is increasingly limited on both low-lying and high islands by habitat loss, sea level rise, and invasive mammalian predators that pose a threat to flightless and flight-compromised curlews during the molting period.

Satellite telemetry has become a common technique to investigate avian life-histories, but whether such tagging will affect fitness is a critical unknown. In this study, we evaluate multi-year effects of implanted transmitters on migratory timing and reproductive performance in shorebirds. Shorebirds increasingly are recognized as good models in ecology and evolution. That many of them are of conservation concern adds to the research responsibilities. In May 2009, we captured 56 female Black-tailed Godwits Limosa limosa limosa during late incubation in The Netherlands. Of these, 15 birds were equipped with 26-g satellite transmitters with a percutaneous antenna (7.8 % ± 0.2 SD of body mass), surgically implanted in the coelom. We compared immediate nest survival, timing of migration, subsequent nest site fidelity and reproductive behaviour including egg laying with those of the remaining birds, a comparison group of 41 females. We found no effects on immediate nest survival. Fledging success and subsequent southward and northward migration patterns of the implanted birds conformed to the expectations, and arrival time on the breeding grounds in 2010–2012 did not differ from the comparison group. Compared with the comparison group, in the year after implantation, implanted birds were equally faithful to the nest site and showed equal territorial behaviour, but a paucity of behaviours indicating nests or clutches. In the 3 years after implantation, the yearly apparent survival of implanted birds was 16 % points lower. Despite intense searching, we found only three eggs of two implanted birds; all were deformed. A similarly deformed egg was reported in a similarly implanted Whimbrel Numenius phaeopus returning to breed in central Alaska. The presence in the body cavity of an object slightly smaller than a normal egg may thus lead to egg malformation and, likely, reduced egg viability. That the use of implanted satellite transmitters in these large shorebirds reduced nesting propensity and might also lead to fertility losses argues against the use of implanted transmitters for studies on breeding biology, and for a careful evaluation of the methodology in studies of migration.

The annual 29 000 km long migration of the bar-tailed godwit, Limosa lapponica baueri, around the Pacific Ocean traverses what is arguably the most complex and seasonally structured atmospheric setting on Earth. Faced with marked variation in wind regimes and storm conditions across oceanic migration corridors, individuals must make critical decisions about when and where to fly during nonstop flights of a week's duration or longer. At a minimum, their decisions will affect wind profitability and thus reduce energetic costs of migration; in the extreme, poor decisions or unpredictable weather events will risk survival. We used satellite telemetry to track the annual migration of 24 bar-tailed godwits and analysed their flight performance relative to wind conditions during three major migration legs between nonbreeding grounds in New Zealand and breeding grounds in Alaska. Because flight altitudes of birds en route were unknown, we modelled flight efficiency at six geopotential heights across each migratory segment. Birds selected departure dates when atmospheric conditions conferred the greatest wind assistance both at departure and throughout their flights. This behaviour suggests that there exists a cognitive mechanism, heretofore unknown among migratory birds, that allows godwits to assess changes in weather conditions that are linked (i.e. teleconnected) across widely separated atmospheric regions. Godwits also showed adaptive flexibility in their response not only to cues related to seasonal changes in macrometeorology, such as spatial shifting of storm tracks and temporal periods of cyclogenesis, but also to cues associated with stochastic events, especially at departure sites. Godwits showed limits to their response behaviours, however, especially relative to rapidly developing stochastic events while en route. We found that flight efficiency depended significantly upon altitude and hypothesize that godwits exhibit further adaptive flexibility by varying flight altitude en route to optimize flight efficiency. The Association for the Study of Animal Behaviour. Published by Elsevier Ltd.