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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.

Migration during spring is usually faster than during autumn because of competition for breeding territories. In some cases, however, the costs and benefits associated with the environment can lead to slower spring migration, but examples are quite rare. We compared seasonal migration strategies of the endangered Baltic population of the dunlin Calidris alpina schinzii using light‐level geolocator data from 26 individuals breeding in Finland. Autumn migration was faster, with individuals showing a ‘jump’ and ‘skipping’ migration strategy characterised by fewer stationary periods, shorter total stopping time and faster flight. Spring migration was slower, with individuals using a ‘skipping’ strategy. The duration of migration was longer for early departing birds during spring but not during autumn suggesting that early spring migrants are prevented from arriving to the breeding areas or that fueling conditions are worse on the stopover sites for early arriving individuals. Dunlins showed high migratory connectivity. All individuals had one long staging at the Wadden Sea in the autumn after which half of the individuals flew 4500 km non‐stop to Banc d’Arguin, Mauritania. The other half stopped briefly on the Atlantic coast on their way to Mauritania. One bird wintered on the coast of Portugal. Nine individuals that carried geolocators for two years were site faithful to their final non‐breeding sites. Based on the strategies during the non‐breeding period we identified, Baltic dunlin may be especially vulnerable to rapid environmental changes at the staging and non‐breeding areas. Consequently, the preservation of the identified non‐breeding areas is important for their conservation.

Animals should cue on information that predicts reproductive success. After failure of an initial reproductive attempt, decisions on whether or not to initiate a second reproductive attempt may be affected by individual experience and social information. If the prospects of breeding success are poor, long-lived animals in particular should not invest in current reproductive success (CRS) in case it generates costs to future reproductive success (FRS). In birds, predation risk experienced during breeding may provide a cue for renesting success. Species having a high FRS potential should be flexible and take predation risk into account in their renesting decisions. We tested this prediction using breeding data of a long-lived wader, the southern dunlin Calidris alpina schinzii. As predicted, dunlin cued on predation risk information acquired from direct experience of nest failure due to predation and ambient nest predation risk. While the overall renesting rate was low (34.5%), the early season renesting rate was high but declined with season, indicating probable temporal changes in the costs and benefits of renesting. We develop a conceptual cost-benefit model to describe the effects of the phase and the length of breeding season on predation risk responses in renesting. We suggest that species investing in FRS should not continue breeding in short breeding seasons in response to predation risk but without time constraints, their response should be similar to species investing in CRS, e.g. within-season dispersal and increased nest concealment.