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BackgroundMigrants have been hypothesised to use different migration strategies between seasons: a time-minimization strategy during their pre-breeding migration towards the breeding grounds and an energy-minimization strategy during their post-breeding migration towards the wintering grounds. Besides season, we propose body size as a key factor in shaping migratory behaviour. Specifically, given that body size is expected to correlate negatively with maximum migration speed and that large birds tend to use more time to complete their annual life-history events (such as moult, breeding and migration), we hypothesise that large-sized species are time stressed all year round. Consequently, large birds are not only likely to adopt a time-minimization strategy during pre-breeding migration, but also during post-breeding migration, to guarantee a timely arrival at both the non-breeding (i.e. wintering) and breeding grounds.MethodsWe tested this idea using individual tracks across six long-distance migratory shorebird species (family Scolopacidae) along the East Asian-Australasian Flyway varying in size from 50 g to 750 g lean body mass. Migration performance was compared between pre- and post-breeding migration using four quantifiable migratory behaviours that serve to distinguish between a time- and energy-minimization strategy, including migration speed, number of staging sites, total migration distance and step length from one site to the next.ResultsDuring pre- and post-breeding migration, the shorebirds generally covered similar distances, but they tended to migrate faster, used fewer staging sites, and tended to use longer step lengths during pre-breeding migration. These seasonal differences are consistent with the prediction that a time-minimization strategy is used during pre-breeding migration, whereas an energy-minimization strategy is used during post-breeding migration. However, there was also a tendency for the seasonal difference in migration speed to progressively disappear with an increase in body size, supporting our hypothesis that larger species tend to use time-minimization strategies during both pre- and post-breeding migration.ConclusionsOur study highlights that body size plays an important role in shaping migratory behaviour. Larger migratory bird species are potentially time constrained during not only the pre- but also the post-breeding migration. Conservation of their habitats during both seasons may thus be crucial for averting further population declines.Electronic supplementary materialThe online version of this article (10.1186/s40462-017-0114-0) contains supplementary material, which is available to authorized users.

BackgroundGeolocators are useful for tracking movements of long-distance migrants, but potential negative effects on birds have not been well studied. We tested for effects of geolocators (0.8–2.0 g total, representing 0.1–3.9 % of mean body mass) on 16 species of migratory shorebirds, including five species with 2–4 subspecies each for a total of 23 study taxa. Study species spanned a range of body sizes (26–1091 g) and eight genera, and were tagged at 23 breeding and eight nonbreeding sites. We compared breeding performance and return rates of birds with geolocators to control groups while controlling for potential confounding variables.ResultsWe detected negative effects of tags for three small-bodied species. Geolocators reduced annual return rates for two of 23 taxa: by 63 % for semipalmated sandpipers and by 43 % for the arcticola subspecies of dunlin. High resighting effort for geolocator birds could have masked additional negative effects. Geolocators were more likely to negatively affect return rates if the total mass of geolocators and color markers was 2.5–5.8 % of body mass than if tags were 0.3–2.3 % of body mass. Carrying a geolocator reduced nest success by 42 % for semipalmated sandpipers and tripled the probability of partial clutch failure in semipalmated and western sandpipers. Geolocators mounted perpendicular to the leg on a flag had stronger negative effects on nest success than geolocators mounted parallel to the leg on a band. However, parallel-band geolocators were more likely to reduce return rates and cause injuries to the leg. No effects of geolocators were found on breeding movements or changes in body mass. Among-site variation in geolocator effect size was high, suggesting that local factors were important.ConclusionsNegative effects of geolocators occurred only for three of the smallest species in our dataset, but were substantial when present. Future studies could mitigate impacts of tags by reducing protruding parts and minimizing use of additional markers. Investigators could maximize recovery of tags by strategically deploying geolocators on males, previously marked individuals, and successful breeders, though targeting subsets of a population could bias the resulting migratory movement data in some species.Electronic supplementary materialThe online version of this article (doi:10.1186/s40462-016-0077-6) contains supplementary material, which is available to authorized users.

The behavioural rhythms of organisms are thought to be under strong selection, influenced by the rhythmicity of the environment1–4. Such behavioural rhythms are well studied in isolated individuals under laboratory conditions1,5, but free-living individuals have to temporally synchronize their activities with those of others, including potential mates, competitors, prey and predators6–10. Individuals can temporally segregate their daily activities (e.g. prey avoiding predators, subordinates avoiding dominants) or synchronize their activities (e.g. group foraging, communal defence, pairs reproducing or caring for offspring)6–9,11. The behavioural rhythms that emerge from such social synchronization and the underlying evolutionary and ecological drivers that shape them remain poorly understood5–7,9. Here, we address this in the context of biparental care, a particularly sensitive phase of social synchronization12 where pair members potentially compromise their individual rhythms. Using data from 729 nests of 91 populations of 32 biparentally-incubating shorebird species, where parents synchronize to achieve continuous coverage of developing eggs, we report remarkable within– and between-species diversity in incubation rhythms. Between species, the median length of one parent’s incubation bout varied from 1 – 19 hours, while period length–the time in which a parent’s probability to incubate cycles once between its highest and lowest value – varied from 6 – 43 hours. The length of incubation bouts was unrelated to variables reflecting energetic demands, but species relying on crypsis (the ability to avoid detection by other animals) had longer incubation bouts than those that are readily visible or actively protect their nest against predators. Rhythms entrainable to the 24-h light-dark cycle were less prevalent at high latitudes and absent in 18 species. Our results indicate that even under similar environmental conditions and despite 24-h environmental cues, social synchronization can generate far more diverse behavioural rhythms than expected from studies of individuals in captivity5–7,9. The risk of predation, not the risk of starvation, may be a key factor underlying the diversity in these rhythms.

Summary 1.The accidental or deliberate release of alien species may be very disruptive to native biota, principally through competition or predation. Naturalized populations of mute swans Cygnus olor in western Europe and North America have overgrazed native aquatic vegetation, competed with other waterbirds, and damaged arable and fodder crops. Numbers may be controlled by destroying or oiling a proportion of eggs in each clutch to prevent hatching (clutch reduction). 2. A difference equation model was used to examine the effectiveness of clutch reduction on a mute swan population in the Wylye Valley, Wiltshire, UK. Model parameters were derived mainly from a long-term study of individually marked birds. Survival and emigration were parameterized as a combined function. The model focused on the non-breeding subpopulation, considered to have a negative impact on local fisheries by overgrazing aquatic macrophytes. 3. The model was also parameterized for another swan population in the West Midlands, UK. This population was characterized by rapid growth in a much larger area, compared with the smaller, comparatively stable population in the Wylye Valley. There were insufficient data available to parameterize accurately the model for other areas. 4. Annual survival rates were high in both populations, ranging between 68% and 73% for juveniles, first-years and non-breeding adults, and between 72% and 90% for breeding adults. Immigration was an important factor in the dynamics of both populations. 5. The effects of different levels of clutch reduction on the Wylye Valley mute swan population were simulated. Reducing clutches to two eggs per clutch lowered nonbreeding numbers by 30% over a 10-year period. Total destruction of all eggs in each clutch stabilized the non-breeding subpopulation but did not eradicate it. The effects of clutch reduction were offset by high survival rates and immigration. 6. In the West Midlands, the simulated restriction of clutches to two eggs stabilized the nonbreeding subpopulation but did not affect breeding numbers. Total destruction of all eggs in each clutch markedly reduced the non-breeding subpopulation. However, immigration was underestimated in this model and may have further diluted the effects of total egg destruction. 7. Synthesis and applications . Clutch reduction is labour intensive, requires persistence to be effective, and its effects may vary between populations depending on immigration rates. An evaluation of the consequences of clutch reduction in advance of implementation in the field is therefore highly desirable. The population model described in this study provides the means to do this. This study has demonstrated that control of breeding output did not alleviate a localized conflict, in this case overgrazing by swans in rivers. Deterrent measures and habitat management at the site of conflict may be more effective. The model permits the exploration of a range of demographic manipulations to determine optimum population management regimes before they are implemented.

Migration is a common phenomenon across many animal taxa. Understanding how migration scales with body size across species is fundamental in the development of migration theory and in making size-related predictions. Although aerodynamic theory and ecophysiological scaling laws have assisted greatly in generating such predictions, their verifications have been limited by a lack of empirical data across a range of body sizes. The recent development of ultra-light tracking devices and its rapid application to migration now allows us to put theory to the test. We used tracking data of seven closely related migratory sandpiper species (family Scolopacidae) along the East Asian-Australasian Flyway to compare their migratory behaviour when migrating towards the breeding grounds as a function of size (50-750 g). We found a marked decline in migration speed (migration distance divided by total migration duration, including time at stopover sites and in flight) with size. Departure date from the non-breeding (i.e. wintering) ground and arrival date at the breeding ground also scaled negatively with size. Total migration duration, migration distance, total staging duration (the number of days staying at stopover sites plus days preparing, i.e. fuelling, prior to initial migration) and step length (distance covered within one migratory leg) were not significantly related with size. Correction for phylogeny showed consistent results for all variables. Besides improving our fundamental understanding of inter-specific variation in migration behaviours, the finding of a clear scaling with size in migration speed and migration timing highlight differential size related capabilities and constraints of migrants. Migratory birds, including sandpipers, are declining on a global scale and particularly along the East Asian-Australasian Flyway. This notion of size-dependency in migratory traits may have a bearing on their vulnerability to specific environmental disturbances along their flyways.

Background In-flight conditions are hypothesized to influence the timing and success of long-distance migration. Wind assistance and thermal uplift are thought to reduce the energetic costs of flight, humidity, air pressure and temperature may affect the migrants’ water balance, and clouds may impede navigation. Recent advances in animal-borne long-distance tracking enable evaluating the importance of these factors in determining animals’ flight altitude. Methods Here we determine the effects of wind, humidity, temperature, cloud cover, and altitude (as proxy for climbing costs and air pressure) on flight altitude selection of two long-distance migratory shorebirds, far eastern curlew (Numenius madagascariensis) and whimbrel (Numenius phaeopus). To reveal the predominant drivers of flight altitude selection during migration we compared the atmospheric conditions at the altitude the birds were found flying with conditions elsewhere in the air column using conditional logistic mixed effect models. Results Our results demonstrate that despite occasional high-altitude migrations (up to 5550 m above ground level), our study species typically forego flying at high altitudes, limiting climbing costs and potentially alleviating water loss and facilitating navigation. While mainly preferring migrating at low altitude, notably in combination with low air temperature, the birds also preferred flying with wind support to likely reduce flight costs. They avoided clouds, perhaps to help navigation or to reduce the risks from adverse weather. Conclusions We conclude that the primary determinant of avian migrant’s flight altitude selection is a preference for low altitude, with wind support as an important secondary factor. Our approach and findings can assist in predicting climate change effects on migration and in mitigating bird strikes with air traffic, wind farms, power lines, and other human-made structures.

Background: Conserving migratory birds is challenging due to their reliance on multiple distant sites at different stages of their annual life cycle. The concept of "flyway", which refers to all areas covered by the breeding, nonbreeding, and migrating of birds, provides a framework for international cooperation for conservation. In the same flyway, however, the migratory activities of the same species can differ substantially between seasons and populations. Clarifying the seasonal and population differences in migration is helpful for understanding migration ecology and for identifying conservation gaps. Methods: Using satellite-tracking we tracked the migration of Whimbrels (Numenius phaeopus variegatus) from nonbreeding sites at Moreton Bay (MB) and Roebuck Bay (RB) in Australia in the East Asian-Australasian Flyway. Mantel tests were used to analyze the strength of migration connectivity between the nonbreeding and breeding sites of MB and RB populations. Welch's t test was used to compare the migration activities between the two populations and between northward and southward migration. Results: During northward migration, migration distance and duration were longer for the MB population than for the RB population. The distance and duration of the first leg flight during northward migration were longer for the MB population than for the RB population, suggesting that MB individuals deposited more fuel before departing from nonbreeding sites to support their longer nonstop flight. The RB population exhibited weaker migration connectivity (breeding sites dispersing over a range of 60 longitudes) than the MB population (breeding sites concentrating in a range of 5 longitudes in Far Eastern Russia). Compared with MB population, RB population was more dependent on the stopover sites in the Yellow Sea and the coastal regions in China, where tidal habitat has suffered dramatic loss. However, RB population increased while MB population decreased over the past decades, suggesting that loss of tidal habitat at stopover sites had less impact on the Whimbrel populations, which can use diverse habitat types. Different trends between the populations might be due to the different degrees of hunting pressure in their breeding grounds. Conclusions: This study highlights that conservation measures can be improved by understanding the full annual life cycle of movements of multiple populations of Whimbrels and probably other migratory birds.