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Although many recent tracking studies have uncovered considerable variation in the migratory routines of birds,1,2 the source of this variation is surprisingly poorly discussed.3 We hypothesize that a wealth of possible factors, including factors other than genetics, translate into these variable outcomes. To demonstrate how factors that are not inherited can shape migratory routine during development, we performed a translocation and delayed-release experiment with juvenile, hand-raised black-tailed godwits Limosa limosa limosa that were carefully matched for ancestral background, releasing siblings 1,000 km apart. Translocated juveniles adopted the spatiotemporal pattern of migration that is habitual for the population at the release location rather than the origin. This leads to the rejection of the hypothesis that the migration of inexperienced birds is only shaped by pre-release factors, including genes, maternal material in the eggs, and a common environment from hatching to fledging. Instead, these findings are consistent with inexperienced migrants also developing their knowledge and capacities through contextual individual learning,4 the precise nature of which now warrants study. The fact that hand-raised godwits, depending on the context in which they are released, can be induced to show different migratory routines, wintering sites, and breeding locations from their siblings indicates that processes during development offer the substrate for rapid adaptation of long-distance migration.

Until recently, Limosa limosa melanuroides was thought to be the only subspecies of Black-tailed Godwit in the East Asian–Australasian Flyway. For this reason, all previous occurrences and counts of Black-tailed Godwits in the flyway have been assigned to melanuroides. However, a larger-bodied subspecies, bohaii, has recently been discovered in the flyway. As a result, the occurrence of Black-tailed Godwits in the flyway needs to be reconsidered such that the specific distribution of each subspecies becomes known. To this end, we developed a simple discriminant function to assign individuals to subspecies based on their bill and wing length. Cross-validation with individuals known to be bohaii or melanuroides, based on molecular analysis, showed the developed function to be 97.7% accurate. When applied to measurements of godwits captured at 22 sites across 9 countries in East–Southeast Asia and Australia, we found that bohaii and melanuroides occurred at most sites and overlapped in their distribution from Kamchatka to Australia. We examined photos from all along the flyway to verify this surprising result, confirming that both subspecies co-occur in most locations. Based on these results, we hypothesise that bohaii and melanuroides from the west of their breeding ranges mostly migrate over Chinese mainland. Birds of both subspecies from the east of their ranges are expected to migrate along the Pacific Ocean. We encourage ringing groups in East–Southeast Asia and Australia to use this simple method to keep adding knowledge about Black-tailed Godwits in the East Asian–Australasian Flyway.

Migration: The large variation in the migration of godwits is mostly the result of consistent differences between individuals. These differences are large compared to other migratory bird species. The survival of godwits during migration is also higher compared to other bird species. The mortality that does occur on migration is mostly the result of adverse wind conditions during northward Sahara crossings.The considerable variation in the migration of godwits might therefore result from an absence of a consistent selection pressure during migration. The consistent differences among individuals develop during multiple years of life. A translocation experiment to Poland with naïve juveniles from The Netherlands showed that the development of their migratory direction and wintering location was not merely inherited. Polish individuals are less consistent in their migration than those from The Netherlands. This might be because there is less canalization through social factors in the lower density Polish population. Population: Godwits are monogamous, but do sometimes lay eggs in the nests of other godwits. Individuals lay similarly sized eggs every year. Our traditional observation methods often fail to capture the first nest and the renests of individual godwits. This observational bias has led to biased estimates in the past. Even though the sex ratio at hatch is equal, higher survival of male chicks and adults leads to a surplus of males. Sexual dimorphism develops during the chick phase in godwits and needs to be accounted for when evaluating the condition of chicks.

Longitudinal tracking studies have revealed consistent differences in the migration patterns of individuals from the same populations. The sources or processes causing this individual variation are largely unresolved. As a result, it is mostly unknown how much, how fast and when animals can adjust their migrations to changing environments. We studied the ontogeny of migration in a long-distance migratory shorebird, the black-tailed godwit Limosa limosa limosa, a species known to exhibit marked individuality in the migratory routines of adults. By observing how and when these individual differences arise, we aimed to elucidate whether individual differences in migratory behaviour are inherited or emerge as a result of developmental plasticity. We simultaneously tracked juvenile and adult godwits from the same breeding area on their south- and northward migrations. To determine how and when individual differences begin to arise, we related juvenile migration routes, timing and mortality rates to hatch date and hatch year. Then, we compared adult and juvenile migration patterns to identify potential age-dependent differences. In juveniles, the timing of their first southward departure was related to hatch date. However, their subsequent migration routes, orientation, destination, migratory duration and likelihood of mortality were unrelated to the year or timing of migration, or their sex. Juveniles left the Netherlands after all tracked adults. They then flew non-stop to West Africa more often and incurred higher mortality rates than adults. Some juveniles also took routes and visited stopover sites far outside the well-documented adult migratory corridor. Such juveniles, however, were not more likely to die. We found that juveniles exhibited different migratory patterns than adults, but no evidence that these behaviours are under natural selection. We thus eliminate the possibility that the individual differences observed among adult godwits are present at hatch o

1. Insufficient reproduction as a consequence of predation on eggs and chicks is a major determinant of population decline in ground-nesting birds, including waders. For many populations, there is an urgent need to maintain breeding populations at key sites, and conservation practitioners need to find viable management solutions to reduce predation. 2. One tool available to the practitioner is fences that exclude key predators from areas containing breeding birds. Temporary electric fencing is an increasingly popular predator exclusion intervention, but such fences have costs associated with purchase and the time needed to erect and maintain them. Their effectiveness and optimal application are also frequently questioned. 3. We evaluate the use of temporary ditch-side four-strand electric fences in lowland grasslands in two countries, The Netherlands and England, in areas containing high densities of breeding waders. 4. In both countries and in all years, godwit and lapwing nest survival was significantly higher within areas enclosed by ditch-side electric fences. Brood survival, assessed for godwits in The Netherlands, was also higher within fenced areas in all years. This demonstrates that using temporary electric fences to enclose ground-nesting birds can be an effective tool for improving breeding productivity. 5. In our study, closely managed electric fences were effective at excluding red foxes Vulpes vulpes, but not avian and other mammalian predators. The positive effect that electric fencing had on nest and brood survival therefore likely results from a reduction in the total number of visits by mammalian predators, and especially visits by foxes. 6. Although it requires a substantial time investment throughout the period of use, our temporary electric fence design provides flexibility compared to other fence designs when it comes to enclosing different areas within a season and between years, as the targets for protection change or as land and flood manage

Habitat loss and shifts associated with climate change threaten global biodiversity, with impacts likely to be most pronounced at high latitudes. With the disappearance of the tundra breeding habitats, migratory shorebirds that breed at these high latitudes are likely to be even more vulnerable to climate change than those in temperate regions. We examined this idea using new distributional information on two subspecies of Black-tailed Godwits Limosa limosa in Asia: the northerly, bog-breeding L. l. bohaii and the more southerly, steppe-breeding L. l. melanuroides. Based on breeding locations of tagged and molecularly assayed birds, we modelled the current breeding distributions of the two subspecies with species distribution models, tested those models for robustness and then used them to predict climatically suitable breeding ranges in 2070 according to bioclimatic variables and different climate change scenarios. Our models were robust and showed that climate change is expected to push bohaii into the northern rim of the Eurasian continent. Melanuroides is also expected to shift northward, stopping in the Yablonovyy and Stanovoy Ranges, and breeding elevation is expected to increase. Climatically suitable breeding habitat ranges would shrink to 16% and 11% of the currently estimated ranges of bohaii and melanuroides, respectively. Overall, this study provides the first predictions for the future distributions of two little-known Black-tailed Godwit subspecies and highlights the importance of factoring in shifts in bird distribution when designing climate-proof conservation strategies.

The endangered continental Black-tailed Godwit (Limosa limosa limosa) is a migratory ground-nesting wader breeding in a wide variety of open, wet habitats across Europe. Conservation research has concentrated on the causes of population decline, but we know surprisingly little about whether any resources limit local breeding populations and if so, whether these are resources for the adults or the chicks. We collected data from 63 key breeding sites in five countries across Europe to test whether, after correcting for differences in surveyed areas, the size of Godwit breeding populations was related to environmental variables (vegetation biomass, soil moisture) or food resources for adult birds (soil invertebrates) or chicks (vegetation dwelling arthropods) measured during different times of the reproductive cycle. We found the number of Godwit territories to be positively related to arthropod abundance during the chick-hatching period. We found additional, weaker support for a positive relation between Godwit territory numbers and the abundance of soil-dwelling invertebrates (mostly earthworms) at clutch laying, but not at chick-hatching. These relationships were observed across countries, while we found little support for relationships within countries, possibly due to the smaller range in conditions that exist within countries. Both vegetation growth and soil moisture weren’t related to Godwit territory numbers. Our results suggest that food abundance for chicks, and to a lesser extent adult birds, are key factors determining the size of local Godwit breeding populations. Conservation management aiming to enhance local Godwit populations should therefore consider the impacts of management strategies on the arthropod prey of chicks.

Grassland breeding waders have been steadily declining across Europe. Recent studies indicating a dramatic decline in grassland invertebrates' abundance and biomass, the key food of most grassland wader chicks, suggest a likely driver of the demise of waders. While agricultural intensification is generally inferred as the main cause for arthropod decline there is surprisingly little information on the relationship between land use intensity and total arthropod abundance in grasslands. Here, we explored those relationships across several key wader breeding habitats by surveying ground-active, aerial and soil-dwelling invertebrate communities in five European countries that range from natural undisturbed bogs to intensively managed grasslands. Using maximum vegetation growth and soil moisture content we investigated how they shape the size of the invertebrate community within and across different countries. We found predominantly positive relationships between grassland invertebrate abundance, biomass and body weight with increasing vegetation growth and soil moisture. Maximum vegetation growth was strongly positively related to ground-active invertebrate abundance and biomass and abundance of soil dwelling invertebrates (mainly earthworms). Body weight of aerial invertebrates furthermore increased with increasing maximum vegetation growth. Our results provide little support for the hypothesis that agricultural practices associated with intensification of grassland management result in an abundance decline of invertebrate prey for wader chicks. Conservation practices aiming to enhance wader chick survival require a careful balancing act between maintaining habitat productivity to secure high prey abundance, and keeping productivity low enough to maintain open swards that do not need to be cut before chicks have fledged.

Long-term population studies can identify changes in population dynamics over time. However, to realize meaningful conclusions, these studies rely on accurate measurements of individual traits and population characteristics. Here, we evaluate the accuracy of the observational methods used to measure reproductive traits in individually marked black-tailed godwits (Limosa limosa limosa). By comparing estimates from traditional methods with data obtained from light-level geolocators, we provide an accurate estimate of the likelihood of renesting in godwits and the repeatability of the lay dates of first clutches. From 2012 to 2018, we used periods of shading recorded on the light-level geolocators carried by 68 individual godwits to document their nesting behaviour. We then compared these estimates to those simultaneously obtained by our long-term observational study. We found that among recaptured geolocator-carrying godwits, all birds renested after a failed first clutch, regardless of the date of nest loss or the number of days already spent incubating. We also found that 43% of these godwits laid a second replacement clutch after a failed first replacement, and that 21% of these godwits renested after a hatched first clutch. However, the observational study correctly identified only 3% of the replacement clutches produced by geolocator-carrying individuals and designated as first clutches a number of nests that were actually replacement clutches. Additionally, on the basis of the observational study, the repeatability of lay date was 0.24 (95% CI 0.17–0.31), whereas it was 0.54 (95% CI 0.28–0.75) using geolocator-carrying individuals. We use examples from our own and other godwit studies to illustrate how the biases in our observational study discovered here may have affected the outcome of demographic estimates, individual-level comparisons, and the design, implementation and evaluation of conservation practices. These examples emphasize the importance of impro

The miniaturization of tracking devices is now rapidly increasing our knowledge on the spatiotemporal organization of seasonal migration. So far, most studies aimed at understanding within- and between-individual variation in migratory routines focus on single populations. This has also been the case for continental Black-tailed Godwits Limosa l. limosa (hereafter Godwits), with most work carried out on individuals from the Dutch breeding population, migrating in relatively large numbers in the westernmost part of the range. Here we report the migratory timing and routes of four adult individuals of the same subspecies from the low-density population in eastern Poland and compare this with previously published data on Godwits breeding in The Netherlands. During northward migration, the birds from Poland departed and arrived later from their wintering and breeding grounds. However, on southward migration the Polish breeding Godwits departed earlier, but arrived one month later than the Dutch birds on their wintering grounds in sub-Saharan Africa. Despite the small sample size of tracked birds from Poland, we find a significantly higher between-individual variation in timing during southward migration in Polish Godwits as compared to the Dutch Godwits. Furthermore, not only did migratory routes differ, but the few Polish Godwits tracked showed a higher level of between- and within-individual variation in route choice during both southward and northward migration. To explain this remarkable discrepancy, we propose that the properties of transmission of social information may be different between Godwits from a high-density population (i.e. the one in The Netherlands) and a low-density population (in Poland) and that this leads to different levels of canalization. To examine this hypothesis, future studies should not only follow individuals from an early age onwards, but also quantify and experimentally manipulate their social environments during migration.

As is the case for most avian species, there is considerable variation in the egg size of Continental Black-tailed Godwits Limosa l. limosa breeding in The Netherlands. It is interesting that egg size has costs and benefits yet varies considerably at the population level. To better understand this variation in egg size, we tested its relationship to a suite of individual and environmental factors. We found that egg size can decrease up to 2.8% throughout a breeding season and that egg size increases with clutch size by 1.4% with each additional egg in the clutch. Female body mass and body size explained 5% of the total variation in egg size observed across the population. Furthermore, females wintering south of the Sahara laid 3% smaller eggs than those wintering north of the Sahara. We also found that egg size increases with age, which may indicate age-related differences in the endogenous and/or exogenous conditions of females. The variation in egg size was, however, mostly the result of consistent differences among individuals across years (repeatability = 0.60). A comparison of daughters with mothers suggested that most of this individual repeatability reflects heritable variation (heritability = 0.64). The actual individual traits that underlie this heritable variation among individuals remain mostly undetermined. Smaller eggs did have a slightly lower chance of hatching, but we found no relationship between egg size and chick survival. Finally, nest and chick survival were strongly correlated with lay date. Thus, in Black-tailed Godwits, lay date may actually reflect a female's endogenous and/or exogenous condition at the moment of egg-laying. This finding may be general across birds, since food supplementation experiments usually result in advanced laying and larger clutch sizes rather than in larger eggs.

Variation in migratory behavior is the result of different individual strategies and fluctuations in individual performances. A first step toward understanding these differences in migratory behavior among individuals is, therefore, to assess the relative contributions of inter- and intra-individual differences to this variation. We did this using light-level geolocators deployed on the breeding grounds to follow continental black-tailed godwits (Limosa limosa limosa) throughout their south- and northward migrations over multiple years. Based on repeated tracks from 36 individuals, we found two general patterns in godwit migratory behavior: First, migratory timing in black-tailed godwits varies mostly because individual godwits migrate at different times of the year. Second, individuals also exhibit considerable variation in timing within their respective migratory windows. Although the absolute amount of inter-individual variation in timing decreased over the course of northward migration, individual godwits still arrived at their breeding grounds across a span of more than 5 weeks. These differences in migratory timing among individuals are larger than those currently observed in other migratory bird species and suggest that the selective forces that limit the variation in migratory timing in other species are relaxed or absent in godwits. Furthermore, we could not attribute these individual differences to the sex or wintering location of an individual. We suggest that different developmental trajectories enabled by developmental plasticity likely result in these generally consistent, life-long annual routines. To investigate this possibility and to gain an understanding of the different selection pressures that could be acting during migration and throughout a godwit's life, future studies should track juvenile godwits and other migratory birds from birth to adulthood while also manipulating their spatiotemporal environment during development.

[Capsule] Most Continental Black-tailed Godwits Limosa limosa limosa using the Doñana wetlands during post-breeding migration appear to begin moult before they arrive and suspend moult before they migrate onwards to West Africa., [Aims] We aim to describe the primary moult strategies and patterns in the Continental Black-tailed Godwits using the Doñana wetlands, a major passage and wintering area for waterbirds in southern Spain., [Methods] Individual godwits were captured, marked and their primary moult was scored in Doñana during the non-breeding season (June–March) in 2011 and 2012. Data from resightings of colour-marked godwits and birds equipped with satellite transmitters were used to estimate stopover duration during post-breeding migration (June–September) to determine if godwits move to West Africa before completing their primary moult., [Results] Average primary moult duration was estimated to be 84 days ± 9 se, during 29 June–21 September and did not differ between sexes. Only 2% of individuals were observed with suspended moult. We estimated stopover duration in Doñana to be 13 days ± 2 se before migrating to West Africa., [Conclusions] Most godwits stage for about two weeks in the Doñana wetlands during southward migration, moult their primaries and appear to suspend moult before crossing the Sahara. Others may complete their primary moult in Doñana, or elsewhere in Europe and overwinter in Doñana where increasing numbers of godwits have been detected in recent years. A few individuals may finish the moult in Doñana and migrate to West Africa late in the post-breeding season.