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Although invasive non-native species can adversely affect biodiversity in many ways, predation of native species by non-native species on islands can be severely damaging. Results of numerous studies document non-native birds preying on birds on islands, but our understanding of the number and type of species affected has been limited by the lack of a global review of these impacts. I identified the non-native bird species that have been recorded preying on birds, the locations where this predation occurred, and the bird species affected. Because the impacts of non-native birds can be particularly severe on small islands, I then identified the islands

Average annual temperatures in the Arctic increased by 2–3 °C during the second half of the twentieth century. Because shorebirds initiate northward migration to Arctic nesting sites based on cues at distant wintering grounds, climate-driven changes in the phenology of Arctic invertebrates may lead to a mismatch between the nutritional demands of shorebirds and the invertebrate prey essential for egg formation and subsequent chick survival. To explore the environmental drivers affecting invertebrate availability, we modeled the biomass of invertebrates captured in modified Malaise-pitfall traps over three summers at eight Arctic Shorebird Demographics Network sites as a function of accumulated degree-days and other weather variables. To assess climate-driven changes in invertebrate phenology, we used data from the nearest long-term weather stations to hindcast invertebrate availability over 63 summers, 1950–2012. Our results confirmed the importance of both accumulated and daily temperatures as predictors of invertebrate availability while also showing that wind speed negatively affected invertebrate availability at the majority of sites. Additionally, our results suggest that seasonal prey availability for Arctic shorebirds is occurring earlier and that the potential for trophic mismatch is greatest at the northernmost sites, where hindcast invertebrate phenology advanced by approximately 1–2.5 days per decade. Phenological mismatch could have long-term population-level effects on shorebird species that are unable to adjust their breeding schedules to the increasingly earlier invertebrate phenologies.

In this paper, we extend our understanding of the migration of Black-tailed Godwits (Limosa limosa limosa) by describing: (1) the orientation and geographic locations of individual migratory routes and (2) the spatial distribution of godwits across seasons and years. We accomplish this using satellite-tracking data from 36 adult godwits breeding in the 200-ha Haanmeer polder in The Netherlands, from 2015 to 2018. During both southward and northward migration, godwits used a narrow migratory corridor along which most individuals made stops within a network of sites, especially the Bay of Biscay, France and Doñana, Spain. Most sites were used consistently by the same individuals across years. However, sites in Morocco were used during northward migration by 75% of individuals, but not revisited by the same individual across years. After southward migration, a small proportion (15%) of godwits spent the entire non-breeding period north of the Sahara, but most (85%) crossed the Sahara and spent at least part of the non-breeding season among seven coastal sites in West Africa and one site in the Inner Niger Delta. Although site-use patterns varied among individuals, individuals showed high site fidelity and were consistent in the number of sites they used from year to year. The considerable differences in the spatial distribution of individuals that breed within a kilometre of one another raise questions about the causes and consequences of individual migratory differences. We discuss that full annual cycle tracking of juveniles from birth to adulthood is needed to understand the source of these individual differences. Our results on the spatial distribution of godwits throughout their annual cycle lay an important foundation of information that can be used to help conserve this declining species.

n Focus: Reneerkens, J., Versluijs, T. S. L., Piersma, T., Alves, J. A., Boorman, M., Corse, C., ... Lok, T. (2020). Low fitness at low latitudes: wintering in the tropics in-creases migratory delays and mortality rates in an Arctic breeding shorebird. Journal of Animal Ecology, 89, 691–703. A central question in migratory ecology has been to understand the fitness consequences of individual variation in migration distance among different species and populations. Reneerkens et al. (2020) investigated the demographic consequences of long-distance migration for Sanderlings Calidris alba, an Arctic-breeding species of sandpiper. Their study population has a remark-able geographic distribution with a breeding range that is concentrated in north-east Greenland and Ellesmere Island, Canada but a nonbreeding range that extends across 85° of latitude from Scotland to Namibia. The authors report on unexpected patterns of latitudinal variation in three demographic parameters: timing of passage on northward migration, probability of juvenile migration and apparent survival of adults. Sanderlings travelling 1,800–2,800 km to settle at north temperate sites dur-ing the nonbreeding season had earlier passage dates, and also higher probabilities of migration and apparent survival. In contrast, birds travelling 6,000–7,800 km to equato-rial sites experienced later passage dates, delayed maturity and lower apparent survival. However, if Sanderlings migrated even farther and flew over 11,000 km to nonbreed-ing sites in Namibia, then their performance was restored to early passage dates and higher survival. Movement tracks from birds tagged with geolocators showed that birds wintering in Namibia make nonstop flights of 7,500 km that bypass West Africa during northward migration. Thus, all lines of evidence suggest that Sanderlings face adversity when spending the nonbreeding season at equatorial latitudes. Moreover, the central finding that components of fitness can have nonlinear relationships with migration dis-tance is a novel discovery that leads to many additional questions. The new findings have broader implications for theoretical models of migration, and for understanding how different patterns of movements may arise or be maintained in migratory species

Background Despite an increasing number of surveys and a growing interest in birdwatching, the population and distribution of Asian Dowitcher (Limnodromus semipalmatus), a species endemic to the East Asian–Australasian and Central Asian Flyways, remains poorly understood, and published information about the species is largely outdated. In boreal spring 2019, over 22,432 Asian Dowitchers were recorded in a coastal wetland at Lianyungang, Jiangsu Province, China, constituting 97.5% of its estimated global population. Methods In 2019 and 2020, we conducted field surveys at Lianyungang to determine the numbers of Asian Dowitchers using the area during both southward and northward migrations. We also assessed the distribution and abundance of Asian Dowitchers elsewhere along the China coast by searching literature and consulting expert opinion. Results The coastal wetlands of Lianyungang are the most important stopover site for Asian Dowitchers during both northward and southward migrations; they supported over 90% of the estimated global population during northward migration in two consecutive years (May 2019 and 2020). This area also supported at least 15.83% and 28.42% (or 30.74% and 53.51% using modelled estimates) of the global population during southward migration in 2019 and 2020 respectively. Coastal wetlands in the west and north of Bohai Bay also have been important stopover sites for the species since the 1990s. Although comprehensive, long-term monitoring data are lacking, available evidence suggests that the population of the species may have declined. Conclusions The high concentration of Asian Dowitchers at Lianyungang during migration means the species is highly susceptible to human disturbances and natural stochastic events. The coastal wetlands of Lianyungang should be protected and potentially qualify for inclusion in China’s forthcoming nomination for World Heritage listing of Migratory Bird Sanctuaries along the Coast of Yellow Sea-Bohai Gulf of China (Phase II) in 2023. Additional research is needed to understand Asian Dowitchers’ distribution and ecology, as well as why such a high proportion of their population rely on the Lianyungang coast.

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.

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.

Sea level rise (SLR) and disturbances from increased storm activity are expected to diminish coastal ecosystems available to nesting species by removing habitat and inundating nests during incubation. We updated the United States Geological Survey's (USGS) Coastal Vulnerability Index, which provides a qualitative and relative assessment of a coastal area's vulnerability to erosion and shoreline retreat as a function of SLR and other factors, for the South Atlantic Bight. We considered a eustatic SLR projection of 14 cm by 2030. We linked long-term survey data for 3 sea turtle species, 3 shorebird species, and 5 seabird species to future coastal erosion vulnerability to SLR to understand effects of future SLR on nesting habitats. Over 2,000 km (43%) of the South Atlantic Bight coastline is projected to have an increase in coastal erosion vulnerability by the 2030s, with respect to its present vulnerability. Future vulnerability of SLR-induced erosion along the South Atlantic Bight is spatially variable, and the 11 coastal study species also varied in their use of coastal habitats with high future coastal vulnerability to SLR. For example, only 23% of high-density nesting habitat for the brown pelican (Pelecanus occidentalis) is expected to be at increased vulnerability to SLR, whereas >70% of the high-nesting density habitat for 2 seabird species (gull-billed tern [Gelochelidon nilotica], sandwich tern [Thalasseus sandvicensis]) is predicted to have higher future coastal erosion vulnerability by 2030. We provide predictions for the level of susceptibility of the study species to erosion from future SLR, which is the first step in managing coastal species for the changing environmental conditions associated with climate change and SLR. (c) 2019 The Authors. Journal of Wildlife Management Published by Wiley Periodicals, Inc. on behalf of The Wildlife Society. USFWS; South Atlantic Landscape Conservation Cooperative The findings and conclusions in this article are those of the authors and do not necessarily represent the views of the U.S. Fish and Wildlife Service. Sea turtle data were provided by the Florida Fish and Wildlife Conservation Commission Statewide Nesting Beach Survey, Canaveral National Seashore, University of Central Florida Marine Turtle Research Group, Archie Carr National Wildlife Refuge survey, Georgia Department of Natural Resources Sea Turtle Conservation Program, South Carolina Department of Natural Resources Marine Turtle Conservation Program, and North Carolina Wildlife Resources Commission Sea Turtle Project. D. Griffin provided data and information for South Carolina sea turtles. Seabird and shorebird nesting and occupancy data were provided by the Florida Fish and Wildlife Conservation Commission, Georgia Department of Natural Resources, South Carolina Department of Natural Resources, and North Carolina Wildlife Resources Commission. J. Brush provided the Florida seabird and shorebird data and background information. We sincerely thank the individual permit holders and volunteers who collected the sea turtle data for Florida, Georgia, North Carolina, and South Carolina, and the Air Force, 45th Space Wing. We appreciate the contributions from S. Pierre and M. Auman in organizing the biological data. We also thank S. Maddock and W. Golder for wintering piping plover data. The authors thank S. Dellinger and J. Heisey for their contributions to the wave gauge data analyses. We thank S. Hagan and D. Passeri for their contribution to the shoreline change analysis. Funding for Florida's sea turtle monitoring program has been provided by grants to the Florida Wildlife Commission under Project E -7 from the USFWS and by proceeds from sales of the Florida marine turtle license plates. This work was funded by the South Atlantic Landscape Conservation Cooperative.

Background Anthropogenic disturbance can negatively affect an animal’s energy budget by evoking movement responses. Existing research focuses mainly on immediate displacement as a disturbance effect, since this can be easily observed in the field. However, effects on movement over longer timescales are poorly examined and it is largely unknown if and to what extent they reflect immediate responses. Longer-term responses could for example be larger than immediate responses if birds, after disturbance, return to the original location and thereby travel twice the immediate disturbed distance. Methods We combined GPS tracking data with observational data to quantify the effects of anthropogenic (air force and walkers) and non-anthropogenic disturbances on distances travelled by roosting Eurasian Oystercatchers (Haematopus ostralegus) during the non-breeding season. We compared immediate displacement after a disturbance with distance travelled during the entire high tide period (longer-term response), while accounting for environmental factors. Additionally, we calculated energy expenditure due to disturbance based on observed disturbance frequencies. Results Disturbance resulted in an immediate displacement response of ~ 200 m (median). Air force disturbances tended to yield larger immediate responses than walker and, especially, than non-anthropogenic disturbances. Longer-term responses and immediate responses were approximately similar, suggesting that, over longer timescales, spatial disturbance effects in the study area remain confined to immediate effects. However, disturbances were infrequent (0.17 disturbances per bird per hour) and most disturbances were of natural origin (62%). Consequently, anthropogenic disturbance of roosting oystercatchers in the study area on average costs 0.08% of the daily energy expenditure. Conclusions Our results suggest that immediate spatial responses to disturbance can be a useful proxy for spatial responses over longer timescales. Over the non-exhaustive range of conditions investigated, energetic consequences of spatial disturbance responses for an oystercatcher in the study area are marginal due to low disturbance levels.

Most birds incubate their eggs, which requires time and energy at the expense of other activities. Birds generally have two incubation strategies: biparental where both mates cooperate in incubating eggs, and uniparental where a single parent incubates. In harsh and unpredictable environments, incubation is challenging due to high energetic demands and variable resource availability. We studied the relationships between the incubation behaviour of sandpipers (genus Calidris) and two environmental variables: temperature and a proxy of primary productivity (i.e. NDVI). We investigated how these relationships vary between incubation strategies and across species among strategies. We also studied how the relationship between current temperature and incubation behaviour varies with previous day's temperature. We monitored the incubation behaviour of nine sandpiper species using thermologgers at 15 arctic sites between 2016 and 2019. We also used thermologgers to record the ground surface temperature at conspecific nest sites and extracted NDVI values from a remote sensing product. We found no relationship between either environmental variables and biparental incubation behaviour. Conversely, as ground-surface temperature increased, uniparental species decreased total duration of recesses (TDR) and mean duration of recesses (MDR), but increased number of recesses (NR). Moreover, small species showed stronger relationships with ground-surface temperature than large species. When all uniparental species were combined, an increase in NDVI was correlated with higher mean duration, total duration and number of recesses, but relationships varied widely across species. Finally, some uniparental species showed a lag effect with a higher nest attentiveness after a warm day while more recesses occurred after a cold day than was predicted based on current temperatures. We demonstrate the complex interplay between shorebird incubation strategies, incubation behaviour, and environmental conditions. Understanding how species respond to changes in their environment during incubation helps predict their future reproductive success.

Several factors affect the flight altitude of migratory birds , such as topography, ambient temperature, wind conditions, air humidity, predation avoidance, landmark orientation, and avoiding over-heating from direct sunlight. 1 , 2 , 3 , 4 , 5 , 6 Recent tracking of migratory birds over long distances has shown that migrants change flight altitude more commonly and dramatically than previously thought. 4 , 5 , 6 , 7 , 8 The reasons behind these altitude changes are not well understood. In theirseasonal migrations between Sweden and sub-Saharan Africa, great snipes Gallinago media make non-stop flights of 4,000–7,000 km, lasting 60–90 h. 9 , 10 Activity and air pressure data from multisensor dataloggers showed that great snipes repeatedly changed altitudes around dawn and dusk, between average cruising heights about 2,000 m (above sea level) at night and around 4,000 m during daytime. Frequency and autocorrelation analyses corroborated a conspicuous diel cycle in flight altitude. Most birds regularly flew at 6,000 m and one bird reached 8,700 m, possibly the highest altitude ever recorded for an identified migrating bird. The diel altitude changes took place independently of climate zone, topography, and habitat overflown. Ambient temperature, wind condition, and humidity have no important diel variation at the high altitudes chosen by great snipes. Instead, improved view for orientation by landmarks, predator avoidance , and not least, seeking cold altitudes at day to counteract heating from direct sunlight are the most plausible explanations for the diel altitude cycle. Together with similar recent findings for a small songbird , 6 the great snipes’ altitudinal performance sheds new light on the complexity and challenges of migratory flights.

Acoustic analysis does not support the elevation of Central and Western Canary Islands subspecies of the Eurasian Stone-curlew Burhinus oedicnemus distinctus to full species. To verify whether the vocal repertoires of Eurasian Stone-curlew subspecies oedicnemus and distinctus show biologically significant quantitative and qualitative differences. We carried out an acoustic analysis of some of the most frequently used call types of Eurasian Stone-curlew, recorded in Italy and in the Canary Islands. Additionally, we used playback experiments to test the response of the two subspecies to calls of their own and the other subspecies. The vocal repertoires of the individuals belonging to the two subspecies were rather similar, but the quantitative analysis of acoustic parameters showed some differences between the populations. In particular, the three most used call types showed higher frequency and faster utterance rhythm for distinctus than oedicnemus. Playback experiments indicated that individuals from the nominate subspecies responded in the same way to the playback of calls of individuals belonging to both subspecies. Acoustic analysis supports the distinctiveness of Eurasian Stone-curlew populations from Central and Western Canary Islands, thus confirming the available morphological and genetic data. These results, however, do not suggest that distinctus should be elevated to a full species.

Factors affecting stopover duration in birds are still much discussed. Field studies report contradictory evidence regarding the effect of fuel reserves on bird decision to depart from a stopover site, while the majority of laboratory tests performed on passerines reported a positive relationship between body conditions and migratory restlessness. In recent years, a few studies integrating laboratory and field investigations suggested that the amount of migratory restlessness could be a proxy of the individual willingness to depart from a stopover site in night-migrating passerines, but similar studies are lacking for other groups of birds. In this paper the factors affecting stopover length in the Wood sandpiper Tringa glareola (Charadriiformes Scolopacidae) were studied by integrating field observations and laboratory tests during spring migration. Our aims were to investigate (1) the influence of body conditions on migratory restlessness and (2) the influence of body conditions and of the amount of migratory restlessness on stopover duration, after controlling for meteorological conditions. Contrary to our expectations, we did not find any relationship between body condition and migratory restlessness nor stopover duration, while for the 1st time in a non-passerine species, evidence was produced that the amount of migratory restlessness is significantly related to the likelihood of leaving the stopover area. Our results emphasize the relevance of the studies integrating field and laboratory tests to understand the physiological and ecological factors affecting migratory behaviour in different bird species. Body conditions seem to be unrelated both to migratory restlessness and stopover duration. Migratory restlessness predicts the likelihood of leaving the stopover site. The quantification of migratory restlessness seems to be a useful tool for studying the effect of intrinsic and extrinsic factors on stopover behaviour in shorebirds.

Understanding relationships between habitat and ecological processes often depends on the spatial scale at which the landscape is measured. A common objective in studies of avian nesting ecology is to link habitat characteristics with nest predation for generating potential conservation and management strategies aimed at increasing avian reproductive success. Yet scale dependence likely plays a considerable role in these assessments, particularly when multiple species of nest predators are present. We identified nest predators of snowy plovers, a ground‐nesting shorebird, and modeled predator‐specific hazard rates of nest mortality in relation to the landscape at multiple spatial scales (200 m, 500 m, 1 km, 2.5 km, and 5 km) and at the nest site. We used a Bayesian latent indicator approach for weighting the influence of habitat across spatial scales and to gain insight into the hierarchical nature of how specific predators responded to the landscape when preying upon plover nests. We did not identify a common spatial scale that best predicted nest predation by all predators, though our results indicated that unfragmented playa and marsh habitats at large spatial scales reduced nest predation by all nest predators. Land cover features including grass, shrubs, human development, and open water elicited variable responses depending on the predator group, and predator groups responded at different spatial scales. Further, the spatial configuration of nests (nest dispersion) influenced nest predation by common ravens Corvus corax and foxes (kit fox Vulpes macrotis and red fox Vulpes vulpes), suggesting that certain predators develop a search image for nests. Our results illustrate that clarifying complex relationships between nest predation and habitat characteristics requires the incorporation of predator identity and scale dependence. Thus, management and conservation plans aiming to increase avian productivity by altering or restoring habitats will likely benefit from consideration of predator‐specific patterns at multiple spatial scales.

SummaryThe Spoon-billed Sandpiper Calidris pygmaea is a ‘Critically Endangered’ migratory shorebird. The species faces an array of threats in its non-breeding range, making conservation intervention essential. However, conservation efforts are reliant on identifying the species’ key stopover and wintering sites. Using Maximum Entropy models, we predicted Spoon-billed Sandpiper distribution across the non-breeding range, using data from recent field surveys and satellite tracking. Model outputs suggest only a limited number of stopover sites are suitable for migrating birds, with sites in the Yellow Sea and on the Jiangsu coast in China highlighted as particularly important. All the previously known core wintering sites were identified by the model including the Ganges-Brahmaputra Delta, Nan Thar Island and the Gulf of Mottama. In addition, the model highlighted sites subsequently found to be occupied, and pinpointed potential new sites meriting investigation, notably on Borneo and Sulawesi, and in parts of India and the Philippines. A comparison between the areas identified as most likely to be occupied and protected areas showed that very few locations are covered by conservation designations. Known sites must be managed for conservation as a priority, and potential new sites should be surveyed as soon as is feasible to assess occupancy status. Site protection should take place in concert with conservation interventions including habitat management, discouraging hunting, and fostering alternative livelihoods.

Publisher's version (útgefin grein), For many migratory species, migration can represent a significant part of the annual cycle and the strategies used to move between the breeding and non-breeding areas vary considerably. Weather conditions are important during migration, particularly wind and temperature, and can play a crucial role in the timing of events during the annual cycle of migratory birds. When timing of specific events is important, for example spring arrival and laying dates, the effects of weather on the previous migration might lead to important fitness consequences, as it may alter migration speed. During spring, Icelandic whimbrels Numenius phaeopus islandicus display two main migratory behaviors: a direct flight from the wintering to the breeding sites (direct migration), or, more commonly, two flights with one stopover (stopover migration). We investigated how wind conditions, temperature and spring departure date may drive individuals to adopt either migratory behavior. Interestingly, we found no differences in wind support during migratory flights, in temperature closer to Iceland or on crosswinds experienced in the region before reaching the main stopover areas. However, when individuals undertook a direct flight, departure date from the wintering sites was on average later, but this was not explained by wind patterns over a period of 7 days prior to departure. In addition, we explored the variation at the individual level for three birds that changed migratory behavior between years. The differences in all variables for these individuals reflected the variation observed at the population level. Overall, in such long migrations, it seems advantageous to perform a shorter flight to a stopover area, from where the weather conditions in the breeding areas may be assessed and avoid the risk of facing stochastic inclement weather prior to breeding, while synchronizing time of arrival with conspecifics. In contrast, direct flights seem more common when individuals are time pressed., This work was funded by RANNIS (Grant Nos: 130412-052 and 152470-052), the University of Iceland Research Fund, FCT/MCTES through national funds to CESAM (UIDP/50017/2020 + UIDB/50017/2020), CC (PD/BD/113534/2015), and JA (SFRH/BPD/91527/2012) and ProPolar.

Publiher's version (útgefin grein), Ground-nesting species are vulnerable to a wide range of predators and often experience very high levels of nest predation. Strategies to reduce nest vulnerability can include concealing nests in vegetation and/or nesting in locations in which nests and eggs are camouflaged and less easy for predators to locate. These strategies could have important implications for the distribution of ground-nesting species and the success rates of nests in areas with differing vegetation structure. However, the factors influencing the success of nest concealment and camouflage strategies in ground-nesting species are complex. Here we explore the effects of local vegetation structure and extent of nest concealment on nest predation rates in a range of ground-nesting, sympatric wader species with differing nest concealment strategies (open-nest species: Oystercatcher Haematopus ostralegus, Golden Plover Pluvialis apricaria and Whimbrel Numenius phaeopus; concealed-nest species: Black-tailed Godwit Limosa limosa, Redshank Tringa totanus and Snipe Gallinago gallinago) in south Iceland, in landscapes that comprise substantial variability in vegetation structure at a range of scales. We monitored 469 nests of these six wader species in 2015 and 2016 and ~40% of these nests were predated. Nest predation rates were similar for open-nest and concealed-nest species and did not vary with vegetation structure in the surrounding landscape, but nest-concealing species were ~10% more likely to have nests predated when they were poorly concealed, and the frequency of poorly concealed nests was higher in colder conditions at the start of the breeding season. For concealed-nest species, the reduced capacity to hide nests in colder conditions is likely to reflect low rates of vegetation growth in such conditions. The ongoing trend for warmer springs at subarctic latitudes could result in more rapid vegetation growth, with consequent increases in the success rates of early nests of concealed-nest species. Temperature-related effects on nest concealment from predators could thus be an important mechanism through which climate change affecting vegetation could have population-level impacts on breeding birds at higher latitudes., This work was supported by University of Iceland Research Fund, an Icelandic Research Council (Rannís) Grant (number 152470‐052), an NERC Grant (number NE/M012549/1), and FCT/MCTES to CESAM (UID/AMB/50017/2019) and individual grants to C.C. (PD/BD/113534/2015) and J.A.A. (SFRH/BPD/91527/2012) through national funds and ProPolar. The authors would like to thank Harry Ewing, Aldís Pálsdóttir, Catriona Morrison, Olivia Hicks, Mags Ramsey and Lilja Jóhannesdóttir for their assistance in the field, all iScreamers for fruitful discussions and the landowners who allowed access to their land for this study. We would like to thank Jeroen Reneerkens and three anonymous reviewers for their helpful comments on the manuscript.

Edital Universal Grant number 440722/2014-4/Brazilian Research Council (CNPq) Programa de Apoio a N?cleos Emergentes - Centro de Piscicultura do IFPA Campus Bragan?a/Funda??o Amaz?nia Paraense de Amparo ? Pesquisa (FAPESPA) Canada-Brazil Awards - Joint Research Projects (/The Canadian Bureau for International Education (CBIE) Instituto Brasileiro de Neuroci?ncias (IBNnet)/Financiadora de Estudos e Projetos (FINEP) Natural Sciences and Engineering Research Council of Canada Instituto Federal de Educa??o Ci?ncia e Tecnologia do Par? - Campus Bragan?a. Laborat?rio de Biologia Molecular e Neuroecologia. Bragan?a, PA, Brazil. Universidade Federal do Par?. Instituto de Ci?ncias Biol?gicas. Hospital Universit?rio Jo?o de Barros Barreto. Laborat?rio de Investiga??es em Neurodegenera??o e Infec??o. Bel?m, PA, Brazil. Universidade Federal do Par?. Instituto de Ci?ncias Biol?gicas. Hospital Universit?rio Jo?o de Barros Barreto. Laborat?rio de Investiga??es em Neurodegenera??o e Infec??o. Bel?m, PA, Brazil. Instituto Federal de Educa??o Ci?ncia e Tecnologia do Par? - Campus Bragan?a. Laborat?rio de Biologia Molecular e Neuroecologia. Bragan?a, PA, Brazil. Universidade Federal do Par?. Museu Paraense Em?lio Goeldi. Curso de P?s-Gradua??o em Zoologia. Bel?m, PA, Brazil. Instituto Federal de Educa??o Ci?ncia e Tecnologia do Par? - Campus Bragan?a. Laborat?rio de Biologia Molecular e Neuroecologia. Bragan?a, PA, Brazil. Universidade Federal do Par?. Instituto de Ci?ncias Biol?gicas. Hospital Universit?rio Jo?o de Barros Barreto. Laborat?rio de Investiga??es em Neurodegenera??o e Infec??o. Bel?m, PA, Brazil. Instituto Federal de Educa??o Ci?ncia e Tecnologia do Par? - Campus Bragan?a. Laborat?rio de Biologia Molecular e Neuroecologia. Bragan?a, PA, Brazil. Universidade Federal do Par?. Instituto de Estudos Costeiros. Bragan?a, PA, Brazil. Universidade Federal do Par?. Instituto de Estudos Costeiros. Bragan?a, PA, Brazil. Instituto Federal de Educa??o Ci?ncia e Tecnologia do Par? - Campus Bragan?a. Laborat?rio de Biologia Molecular e Neuroecologia. Bragan?a, PA, Brazil. Universidade Federal do Par?. Instituto de Ci?ncias Biol?gicas. Hospital Universit?rio Jo?o de Barros Barreto. Laborat?rio de Investiga??es em Neurodegenera??o e Infec??o. Bel?m, PA, Brazil / Minist?rio da Sa?de. Secretaria de Vigil?ncia em Sa?de. Instituto Evandro Chagas. Laborat?rio de Miscroscopia Eletr?nica. Ananindeua, PA, Brasil. Instituto Federal de Educa??o Ci?ncia e Tecnologia do Par? - Campus Bragan?a. Laborat?rio de Biologia Molecular e Neuroecologia. Bragan?a, PA, Brazil. University of Western Ontario. Department of Psychology, Advanced Facility for Avian Research. London, ON, Canada. Universidade Federal do Par?. Instituto de Ci?ncias Biol?gicas. Hospital Universit?rio Jo?o de Barros Barreto. Laborat?rio de Investiga??es em Neurodegenera??o e Infec??o. Bel?m, PA, Brazil. Instituto Federal de Educa??o Ci?ncia e Tecnologia do Par? - Campus Bragan?a. Laborat?rio de Biologia Molecular e Neuroecologia. Bragan?a, PA, Brazil. Semipalmated sandpiper (Calidris pusilla) migration to the Southern Hemisphere includes a 5-day non-stop flight over the Atlantic Ocean, whereas semipalmated plover (Charadrius semipalmatus) migration, to the same area, is largely over land, with stopovers for feeding and rest. We compared the number and 3D morphology of hippocampal astrocytes of Ch. semipalmatus before and after autumnal migration with those of C. pusilla to test the hypothesis that the contrasting migratory flights of these species could differentially shape hippocampal astrocyte number and morphology. We captured individuals from both species in the Bay of Fundy (Canada) and in the coastal region of Bragan?a (Brazil) and processed their brains for selective GFAP immunolabeling of astrocytes. Hierarchical cluster analysis of astrocyte morphological features distinguished two families of morphological phenotypes, named type I and type II, which were differentially affected after migratory flights. Stereological counts of hippocampal astrocytes demonstrated that the number of astrocytes decreased significantly in C. pusilla, but did not change in Ch. semipalmatus. In addition, C. pusilla and Ch. semipalmatus hippocampal astrocyte morphological features were differentially affected after autumnal migration. We evaluated whether astrocyte morphometric variables were influenced by phylogenetic differences between C. pusilla and Ch. semipalmatus, using phylogenetically independent contrast approach, and phylogenetic trees generated by nuclear and mitochondrial markers. Our findings suggest that phylogenetic differences do not explain the results and that contrasting long-distance migratory flights shape plasticity of type I and type II astrocytes in different ways, which may imply distinct physiological roles for these cells.

Environmental changes can propagate through food webs in complex ways via trophic cascades. In the North American Arctic, hyperabundant populations of geese are causing significant habitat change and the resulting trophic cascades are known to impact plant and invertebrate communities. However, the potential impacts on other tundra‐nesting birds are not fully understood. Here, we evaluate the impacts of light geese (Snow Geese, Chen caerulescens, and Ross’ Goose, Chen rossii) populations on other tundra‐nesting birds, using count data collected during bird surveys conducted at varying distances from light goose colonies across the Canadian Arctic. From a dataset of 920 12–16 ha plots distributed across the Canadian Arctic, we identified 527 that were within 200 km of the mapped extent of known light goose colonies. After accounting for regional variation, we demonstrate that densities of most Cover‐Nesting Shorebirds and passerines are depressed in the vicinity of light goose colonies. We suggest that these trends are the results of the combined effects of goose‐induced changes in habitat and predator–prey interactions. These direct and indirect effects of light geese could be contributing to the declines experienced by some tundra‐nesting bird populations in parts of their range. Ongoing climate‐related changes to habitat, predators, and arthropods could interact with this effect of geese to further alter the suitability of tundra habitats for Arctic‐breeding birds, in additive or even unanticipated ways.

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.

Because migration is highly time-constrained and migration timing varies among individuals, the responses of migrants to food shortage at a refuelling site could differ between individuals that arrive early and late at the site. To test this hypothesis, we compared the stopover decision, in terms of occurrence and length of stay (LOS), of radiotagged Great Knots Calidris tenuirostris before (2012) and after (2015) a dramatic decline in food supply at a critical spring final pre-breeding refuelling site in the northern Yellow Sea. The probability of occurrence at the refuelling site was consistent between the two years, whereas the average LOS significantly shortened in the year of food shortage in late-arriving individuals. This suggests migration timing intensifies the influence of food shortage in late-arriving individuals, which might be more sensitive and vulnerable to food shortage at refuelling sites compared with early-arriving individuals.

Each May for over three decades, avian influenza A viruses (IAVs) have been isolated from shorebirds and gulls (order Charadriiformes) at Delaware Bay (DE Bay), USA, which is a critical stopover site for shorebirds on their spring migration to arctic breeding grounds. At DE Bay, most isolates have been recovered from ruddy turnstones (Arenaria interpres), but it is unknown if this species is involved in either the maintenance or movement of these viruses outside of this site. We collected and tested fecal samples from 2823 ruddy turnstones in Florida and Georgia in the southeastern United States during four winter/spring sample periods—2010, 2011, 2012, and 2013—and during the winters of 2014/2015 and 2015/2016. Twenty-five low pathogenicity IAVs were recovered representing five subtypes (H3N4, H3N8, H5N9, H6N1, and H12N2). Many of these subtypes matched those recovered at DE Bay during the previous year or that year’s migratory cycle, suggesting that IAVs present on these southern wintering areas represent a source of virus introduction to DE Bay via migrating ruddy turnstones. Analyses of all IAV gene segments of H5N9 and H6N1 viruses recovered from ruddy turnstones at DE Bay during May 2012 and from the southeast during the spring of 2012 revealed a high level of genetic relatedness at the nucleotide level, suggesting that migrating ruddy turnstones move IAVs from wintering grounds to the DE Bay ecosystem.

BackgroundDeparture decisions and behaviors of migratory birds at stopover sites are expected to maximize fitness by trade-offs among avoiding predators, optimizing refueling (energy) capacity, and matching other life-history events. We predict that species with different body sizes and migratory destinations will exhibit different behaviors when departing from the same stopover site. We also predict that with strong time constraint at the final pre-breeding stopover site, departure decisions may be less sensitive to exogenous factors, such as wind condition, compared to other stopover or nonbreeding sites.MethodsWe recorded migratory departures of four shorebird species, i.e. Eurasian Curlew (Numenius arquata), Bar-tailed Godwit (Limosa lapponica), Great Knot (Calidris tenuirostris), and Grey Plover (Pluvialis squatarola), at Yalujiang Estuary Wetlands in China, a final pre-breeding stopover site in the northern Yellow Sea, from 2011 to 2014. We compared flock sizes, departure time and departure directions between species, and investigated the effects of tide and weather conditions (rain and ground wind speed and direction) on the departure decision of shorebirds.ResultsWe found that larger species departed in smaller flocks and were more variable in daily departure time. Departure trajectory of all four species appeared to be influenced by coastal topography. With the east–west coastline and intertidal mudflat on the south, birds exhibited westward or eastward deflection from the shortest migratory routes. Bar-tailed Godwit was the only species that deviated to the southeast and did not climb over the land. Birds avoided departure during precipitation, while their departure was not related to ground wind benefit or tidal condition.ConclusionsBody size among species, which influences their vulnerability to predators, might be important in shaping shorebird departure strategies. Diverse departure directions could be the result of different wind use tactics in climbing stage. Narrow optimal time window of breeding might lead to reduced flexibility in departure date at a final pre-breeding site. Both endogenous and exogenous are important in shaping departure behaviors and decisions.

The Least Sandpiper Calidris minutillais a Nearctic migrant shorebird that breeds in the North American taiga and subarctic tundra biomes and mainly spends the nonbreeding season in the southern United States, Central America, the Caribbean, and northern South America. There are only two extralimital records of this species for Argentina with photographic evidence: one came from La Pampa province and other from Córdoba province.We observed a single Least Sandpiperin the banks of Arroyo Chico (37 o 26´8"S, 57o 11´43.49"W), a small stream that crosses Medaland Ranch. A White-rumped Sandpiper Calidris fuscicollis accompanied the Least. Approximately 50 meters from them, also on the banks of Arroyo Chico, there was a mixed flock of resting shorebirds comprising 63 Hudsonian Godwits Limosa haemastica, 11 American Golden Plovers,six Black-necked Stilts Himantopus mexicanus, two Stilt Sandpipers Calidris himantopus, two White-rumped Sandpipers, and two Lesser Yellowlegs Tringa flavipes; there were also three Neotropic Cormorants Phalacrocorax brasilianus. The Least Sandpiper was feeding by probing in the mud, but quickly flew as we approached to take a photograph. It was easily identified by its smaller size compared to the White-rumped and by its yellowishgreen short legs with longish toes. Fil: Martínez Curci, Natalia Soledad. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Pretelli, Matías Guillermo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Cavalli, Matilde. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Isacch, Juan Pablo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Loredo, M. Ángeles. Municipalidad de Villa Gesell; Argentina

While the principal use of light-recording geolocators is to determine geographical locations of migratory birds, supplementary wetness data have been used to refine estimates of minimum flight duration, on the assumption that a wet logger indicates the bird is on the ground. We provide a test of this assumption, by comparing wetness values against directly observed migratory departures of logger-equipped Bar-tailed Godwits Limosa lapponica and Red Knots Calidris canutus from the Manawatu River Estuary, New Zealand. Loggers recorded wetness every 10 min (Biotrack MK4093 and MK5093) or every one or four hours (Migrate Technology C65K). We retrieved loggers from 41 godwits from 2008– 2014 and from seven Red Knots in 2013–2014 for which we had corresponding departure information; in total there were 51 departures of godwits and seven of knots that we could match to actual departure times (this included multiple years for some godwits). Overall, 10-min wetness data were very accurate for both godwits and knots (median estimated departure times were 14 min and one min later than true departure, respectively), as were the 60-min and 240-min loggers on godwits if corrected by the wet counts that are recorded within measurement intervals (medians of 16 min earlier and 8 min earlier, respectively). These longer-interval loggers were still reasonably accurate without this adjustment (medians of 37 min and 74 min later, respectively). There was substantial variation between individuals and logger types, with 10-min loggers going dry up to 148 min (godwit) or 55 min (knot) earlier than true departure, while the 60-min and 240-min loggers recorded wetness up to 142 min or 124 min later than true departure (or 195 min or 232 min later, if unadjusted). Some of this variation simply reflects the interval over which wetness is recorded, but bird behaviour and/or logger performance must play a role in some cases (e.g. the logger going dry before departure or remaining wet after departure). Given observed bird behaviour upon arrival after migration (feeding on wet tidal flats), the wetness recording of geolocators is likely to give an accurate estimate of migratory flight duration, at least for species that frequent wet, particularly marine, habitats.

We investigated habitat preference of Common Sandpipers as part of a monitoring program in the Őrség National Park, Hungary. Field observations were conducted during the summers between 2008 and 2012 along a 47-km long section of the River Rába. During the observations we recorded the number and location of birds on the river bank. We divided the studied area into 1 km long sections and measured the proportion of the visually distinguishable habitat types (water, low gravel and sand bank, vegetation and degraded area) from a digitalized map. Furthermore, we recorded the number of the low banks and the number of bends of the river within each section, as well as the sections’ distance from the closest hydroelectric power plants and human settlements. In 2012 we also performed a detailed habitat mapping, recording the proportion of the vegetation types along the river bank and the number of fishing spots, embankment strengthenings and gravel banks. We tested the correlations between these habitat variables and number of birds present in the river sections. Our results show that Common Sandpipers were observed more frequently in locations which have (1) larger number and area of low gravel and sand banks, (2) less dense vegetation, and (3) lower proportion of degraded habitats. These findings can be taken into account in the conservation management of River Rába

In expanding populations, individuals may increasingly be forced to use sites of relatively low quality. This process, named the "buffer effect," was previously described for the Black-tailed Godwit (Limosa limosa islandica) in its use of nonbreeding sites in Great Britain and of breeding areas in Iceland. On the basis of diet analyses from droppings and stable isotopes, we describe a new case for the expanding French wintering population of the Black-tailed Godwit, an expansion accompanied by a drastic change in feeding strategy. In the 1990s, Black-tailed Godwits started using intertidal mudflats at Ile de Re, where they eat the rhizomes of seagrass (Zostera noltii) rather than the customary shellfish (Macoma balthica) eaten at both the preferred (initial) site (Aiguillon Bay) and the area occupied last (Yves to Marennes-Oleron bays). Individually color-marked godwits appeared faithful to both diet type and site, suggesting a cost of a change of strategy. This represents a first case of rhizome-feeding in shorebirds, and it exemplifies a case of carnivorous birds occupying a new site shifting to herbivory.

The major histocompatibility complex (MHC) includes highly polymorphic gene families encoding proteins crucial to the vertebrate acquired immune system. Classical MHC class I (MHCI) genes code for molecules expressed on the surfaces of most nucleated cells and are associated with defense against intracellular pathogens, such as viruses. These genes have been studied in a few wild bird species, but have not been studied in long-distance migrating shorebirds. Red Knots Calidris canutus are medium-sized, monogamous sandpipers with migratory routes that span the globe. Understanding how such long-distancemigrants protect themselves from disease has gained new relevance since the emergence of avian-borne diseases, including intracellular pathogens recognized by MHCI molecules, such as avian influenza. In this study, we characterized MHCI genes in knots and found 36 alleles in eight individuals and evidence for six putatively functional and expressed MHCI genes in a single bird. We also found evidence for recombination and for positive selection at putative peptide binding sites in exons 2 and 3. These results suggest surprisingly high MHC diversity in knots, given their demographic history. This may be a result of selection from diverse pathogens encountered by shorebirds throughout their annual migrations.

The spatial distribution of foraging animals at a given time simultaneously depends on (1) exogenous environmental variables such as resource availability and abiotic habitat characteristics, and (2) the endogenous variable social aggregation made up of the opposing mechanisms of conspecific attraction and repulsion. We developed an exogenous environment-social aggregation model to analyse the spatial distributions of six abundant shorebird species in the Dutch Wadden Sea at resolutions of 150 x 150, 200 x 200 and 250 x 250 m. We used these resolutions to check the robustness of the estimates to the modifiable areal unit problem. We estimated the model parameters by spatial autoregression. This approach enables, among others, estimation of the direct and indirect effects of an exogenous environmental variable on animal density. The former is given by the regression coefficient and the latter, which is due to the amplification of the direct effect by social aggregation, by the spatial multiplier. At all resolution levels and for all species, the explanatory power of social aggregation, measured by Nagelkerke R-2, was larger than the combined contribution of the exogenous environmental variables food availability, silt content and mudflat elevation. Social aggregation was stronger for dunlin, Calidris alpina, red knot, Calidris canutus, and curlew, Numenius arquata, than for oystercatcher, Haematopus ostralegus, grey plover, Pluvialis squatarola, and bar-tailed godwit, Limosa lapponica. The total impacts (that is, direct effect plus all indirect impacts) of the exogenous environmental predictors tended to exceed substantially the direct effects (which tend to be the only ones examined in studies on foraging distributions). (C) 2012 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

The effect of habitat management is commonly evaluated by measuring population growth, which does not distinguish changes in reproductive success from changes in survival or the effects of immigration or emigration. Management has rarely been evaluated considering complete life cycle of the target organisms, including also possible negative impacts from management. We evaluated the effectiveness of cattle grazing in the restoration of coastal meadows as a breeding habitat for small and medium-sized ground-nesting birds by examining the size and demography of a southern dunlin (Calidris alpina schinzii) breeding population. Using a stochastic renesting model that includes within-season variation in breeding parameters, we evaluated the effect of grazing time and stocking rates on reproduction. The census data indicated that the population was stable when nest trampling was prevented, but detailed demographic models showed that the population on managed meadows was a sink that persisted by attracting immigrants. Even small reductions in reproductive success caused by trampling were detrimental to long-term viability. We suggest that the best management strategy is to postpone grazing to after the 19th of June, which is about three weeks later than what is optimal from the farmer's point of view. The differing results from the two evaluation approaches warn against planning and evaluating management only based on census population size and highlight the need to consider target-specific life history characteristics and demography. Even though grazing management is crucial for creating and maintaining suitable habitats, we found that it was insufficient in maintaining a viable population without additional measures that increase nest success. In the presently studied case and in populations with similar breeding cycles, impacts from nest trampling can be avoided by starting grazing when about 70% of the breeding season has past.

The Greenlandic and west-central Siberian breeding populations of Sanderlings Calidris alba are separated by ca. 2000 km during the breeding season, but mix in Europe to some extent during migration. However, the number of Siberian Sanderlings that spend the non- breeding season along the East Atlantic Flyway (extending from western Europe to South Africa), if any, is unknown. Although both populations are considered part of the nominate subspecies C. a. alba based on morphology, population structure in Sanderlings has yet to be described with molecular methods. We examined genetic differenti- ation at the mtDNA control region (CR) and seven microsatellite loci between Greenland- and Siberia-breed- ing Sanderlings in order to: (1) develop a diagnostic tool for assessing the breeding origin of Sanderlings along the East Atlantic Flyway, and (2) provide a comparison with the co-distributed and ecologically similar Red Knot, in which CR differentiation of geographically analogous populations (C. canutus islandica and C. c. canutus) has indicated isolation of lineages near the time of the last glacial maximum. By contrast, we found only weak dif- ferentiation between the Sanderling breeding populations at the CR, and no differentiation at microsatellite loci. These results suggest that the assignment of breeding ori- gin of Sanderlings on Afro-European flyways will not be possible with simple and inexpensive genetic methods, and imply that Sanderlings and Red Knots have very different post-glacial phylogeographic histories.

International audience; Trace elements concentrations (Ag, As, Cd, Co, Cr, Cu, Fe, Hg, Mn, Ni, Pb, Se, Zn) were investigated in the liver, kidneys, muscle and feathers of 31 black-tailed godwits (Limosa limosa) accidentally killed during catches by mist net in the Pertuis Charentais, Atlantic coast of France. Analyses of carbon and nitrogen stable isotope ratios were carried out in liver, muscle and feathers in order to elucidate dietary patterns and to determine whether differences in diet explained the variation in elemental uptake. This study also aimed to have a preliminary assessment of sub-lethal effects triggered by trace elements through the investigation of gene expressions by quantitative real-time PCR, antioxidant enzyme activities (catalase, superoxide dismutase, glutathione peroxidase), and metallothionein (MT) levels. The results showed that Cr and Ni concentrations in tissues of adults were lower than in juveniles in part because adults may have eliminated these trace elements through moulting. Except for Cd and Ni, trace element concentrations were negatively correlated to the body mass of godwits. Ag, As, Hg and Se concentrations were positively linked with the trophic position of birds. The diet could be considered as a fundamental route of exposure for these elements demonstrating therefore the qualitative linkage between dietary habits of godwits and their contaminant concentrations. Our results strongly suggest that even though trace elements concentrations were mostly below toxicity threshold level, the elevated concentrations of As, Ag, Cd, Cu, Fe and Se may however trigger sub-lethal effects. Trace elements appear to enhance expression of genes involved in oxidative stress defence, which indicates the production of reactive oxygen species. Moreover, birds with the highest concentrations appeared to have an increased mitochondrial metabolism suggesting that the fight against trace element toxicity requires additional energetic needs notably to produce detoxification mechanisms such as metallothioneins.

In this paper, I argue that to fully grasp the generation and maintenance of variation in the migratory phenotypes of (shore-)birds we need to expand our scientific search image and include developmental processes and non-genetic pathways of inheritance in the explanatory frameworks. Traditionally, studies of micro-evolution of migratory phenotypes were restricted to comparative studies on migratory versus non-migratory taxa, and artificial selection and heritability experiments on quantitative behavioural traits related to migration. Such studies had a focus on the genetic axis of inheritance and were restricted to songbirds. In avian groups such as the shorebird families Scolopacidae and Charadriidae, all but a few island species are migrants, which precludes comparative studies at the species level. Like other taxa, shorebirds have geographically separate breeding populations (either or not recognized as subspecies on the basis of morphological differences) which differentiate with respect to the length, general direction and timing of migration, including the use of fuelling at staging sites and the timing of moult. However, their breeding systems preclude artificial selection and heritability experiments on quantitative traits. This would seem to limit the prospects of evolutionary analysis until one realizes that the speed of evolutionary innovation in shorebird migratory life-histories may be so fast as to necessitate other avenues of explanation and investigation. According to our best current estimates based on mitochondrial gene sequence variation, in Red Knots Calidris canutus considerable phenotypic variation has evolved since the Last Glacial Maximum ca. 20,000 years ago, to the extent that six subspecies are currently recognized. This would be too short a time for the origin of the qualitatively and quantitatively distinct and non-overlapping traits to be explained by random point mutations followed by natural selection, although we cannot dismiss the possibility of previously unexpressed (standing) genetic variation followed by selection. I argue that, to understand the flyway evolution of such shorebirds in the 'extended' evolutionary framework, we need to give due attention to developmental versatility and broad-sense epigenetic evolutionary mechanisms. This means that experimental studies at the phenotypic level are now necessary. This could involve a combination of observational studies in our rapidly changing world, common garden experiments, and even experiments involving global-scale displacements of particular migratory phenotypes at different phases of development. I provide suggestions on how such experiments could be carried out.

TOC Summary: A lineage unique to Australia has been identified., We investigated carriage of avian influenza viruses by wild birds in Australia, 2005–2008, to assess the risks to poultry industries and human health. We collected 21,858 (7,357 cloacal, 14,501 fecal) samples and detected 300 viruses, representing a detection rate of ≈1.4%. Rates were highest in autumn (March–May) and differed substantially between bird types, areas, and years. We typed 107 avian influenza viruses and identified 19 H5, 8 H7, and 16 H9 (40% of typed viruses). All were of low pathogenicity. These viruses formed clearly different phylogenetic clades to lineages from Eurasia or North America, suggesting the potential existence of Australian lineages. H7 viruses were similar to highly pathogenic H7 strains that caused outbreaks in poultry in Australia. Several periods of increased detection rates (numbers or subtypes of viruses) were identified. This study demonstrates the need for ongoing surveillance to detect emerging pathogenic strains and facilitate prevention of outbreaks.

Each year, hundreds of thousands of shorebirds use Delaware Bay, on the northeast coast of the United States, as a final stopover before migration to breeding areas. The bay provides them with abundant Horseshoe Crab (Limulus polyphemus) eggs, which they use to gain the fat stores necessary for continued migration and subsequent survival on the breeding grounds. However, abundant food attracts dense mixed-species flocks, which may facilitate pathogen transmission, and migration itself may suppress immune defense. Despite the potential importance of disease risk and immune function during migration, little is known about how immune function changes during stopover in migratory shorebirds. To examine this, we measured constitutive immune function in Red Knots (Calidris canutus aura) during stopover in Delaware Bay. We found lower total leukocyte, lymphocyte, and monocyte concentrations, complement-mediated lysis, and haptoglobin activity in birds recovering protein after migration than in birds storing fat to fuel subsequent flight. We discuss two possible reasons for this result. First, fueling birds may have an increased rate of infection or may be bolstering immune defense in response to high antigen exposure. Second, recovering birds may be immuno-compromised because of the physical strain of migratory flight or as a result of adaptive tradeoffs between immune function and migration. Received IS January 2009, accepted 26 August 2009.

In response to environmental change, species have been observed altering their migratory behaviours. Few studies, however, have been able to determine whether these alterations resulted from inherited, plastic or flexible changes. Here, we present a unique observation of a rapid population-level shift in migratory routes—over 300 km from Spain to Portugal—by continental black-tailed godwits Limosa limosa limosa . This shift did not result from adult godwits changing staging sites, as adult site use was highly consistent. Rather, the shift resulted from young godwits predominantly using Portugal over Spain. We found no differences in reproductive success or survival among individuals using either staging site, indicating that the shift resulted from developmental plasticity rather than natural selection. Our results therefore suggest that new migratory routes can develop within a generation and that young individuals may be the agents of such rapid changes.

Subspecies Calidris canutus islandica of the Red Knot breeds on the arctic tundra of northeastern Canada and northern Greenland and winters along the coasts of northwestern Europe. During northward migration, it stops over in either Iceland or northern Norway. It has been assumed that it does the same during southward migration. Using ratios of stable carbon isotopes (delta(13)C) in whole blood, blood cells, and plasma, we investigated evidence for a stopover in Iceland en route from the breeding grounds to the Dutch Wadden Sea. With the expected diet (shellfish) and stopover duration at Iceland (12-15 days, maximum 17 days) and the turnover rates of blood cells (15.1 days) and plasma (6.0 days), Red Knots that stopped in Iceland should arrive with a blood (cell) delta(13)C midway between a tundra (-24.7 parts per thousand) and a marine value (-14.0 parts per thousand) and a plasma delta(13)C approaching the marine value (-15.3 parts per thousand). However, many adults arriving at the Wadden Sea had delta(13)C ratios in blood (cells) and plasma below these levels, and some arrived with clear tundra signals in blood cells, suggesting that they skipped Iceland during southward migration. Surprisingly, available data suggest this also to be true for juveniles during their first southward migration. The delta(13)C signature of second-year birds confirmed that they oversummered in the Wadden Sea. Our findings contradict the largely untested idea that juvenile shorebirds make more stopovers than adults as well as the idea that the migration between the Nearctic and Europe is necessarily a two-leg process.

Afro-Siberian Red Knots Calidris canutus canutus use the western Dutch Wadden Sea as a refuelling area during southward migration from Taimyr to West Africa. Here we document the decline of their food stocks in this area, based on a yearly large-scale benthic mapping effort, from 1996 to 2005. For each benthic sampling position, intake rate (mg/s, ash-free dry mass) was predicted by an optimal diet model based on digestive rate maximization. Over the ten years, when accounting for a threshold value to meet energetic fuelling demands, subspecies canutus lost 86% of its suitable foraging area. Over this period, the proportion of probable canutus in mist-net catches in July-August declined relative to overwintering islandica Knots. This suggests that canutus dropped even more in numbers than islandica, for which we showed earlier a food-explained decline in numbers. We discuss the possible causality between a decline in the quality of intertidal mudflats in the Dutch Wadden Sea and population declines of Knots in the West-African wintering quarters.

Every year, Brazil receives large flocks of nearly 40 migratory shorebirds species. The accumulation of body fat and nutrients during the summer is necessary for the long return flights of these birds and is fundamental for feather moulting and the change of their plumage. We present here an examination of the relationship between body mass and plumage change in Arenaria interpres (Linnaeus, 1758), one of those birds, over time during its wintering period on the Brazilian coast. We analyzed information collected at five traditional stopover sites along the Brazilian coast, between 1997 and 2007. During the month of September, individuals with intermediate or breeding plumage had smaller body masses as compared to other months. From October to December, adult individuals were only observed with eclipse plumage and had average body masses of approximately 100 g. In March, individuals with intermediate, eclipse and breeding plumages were recorded, but their average body mass remained at approximately 100 g. In April and May the numbers of individuals with breeding or intermediate plumage increased, and they showed significant increases in body mass at a rate of approximately 1.5 and 2.3 g per day, in the north-northeastern and south coast, respectively, leading to an average mass of 124 and 143g in these months. That is suggested to be the departure mass of A. interpres in the Brazilian north-northeastern and south coast, respectively, when starting the migration to the breeding sites.

The Red Knot (Calidris canutus) is a long-distance migrant breeding on tundra in the high Arctic and wintering along temperate and tropical coasts. Preflight fueling rate is a major determinant of successful migration, yet individual fueling rates are impossible to determine because Red Knots cannot be recaptured easily. These problems can be overcome by estimating changes in body mass from plasma metabolites. Plasma metabolites are, however, sensitive to stress and time since last meal, limiting studies to situations where birds can be bled almost immediately after capture. Such sampling is almost impossible in the field, where Red Knots are often captured with mist nets in darkness. This study on captive Red Knots investigates whether plasma metabolites obtained from blood samples taken up to 3 hr after capture can be used to predict individual long-term (weekly) body-mass changes during the natural spring preflight fueling period. Triglyceride decreased and beta-hydroxybuty rate increased with time since capture, and these changes varied with time since start of the spring fueling period. beta-Hydroxybutyrate and uric acid were correlated with weekly body-mass change, but triglyceride was not. Triglyceride was correlated with overall body mass. Weekly body-mass change was best predicted with a model including all metabolites and body mass. Time of blood sampling (immediately or 3 hr after capture) did not affect the accuracy of the predictions. The predictions were not accurate enough to allow comparisons of individuals; they should be used only to compare groups.

Four Great Knots (Calidris tenuirostris) were kept for six years in a constant-temperature indoor aviary. For two of those six years, they were kept under photoperiodic conditions that mimicked natural changes in daylength for wild birds, followed by four years under a constant photoperiod (light:dark cycle 12:12 h). Under cyclical "natural" photoperiods, three of the four birds maintained cycles of body mass and contour and flight-feather molt somewhat comparable to that of free-living birds, though the multiple mass peaks characteristic of northward migration were replaced by a single period of high body mass; the mass peaks for southward migration appeared to be absent. Contour-feather molts between nonbreeding and breeding plumages were delayed, and the period of wing molt was longer than in free-living birds. Under constant photoperiods, clear circannual phenotype cycles were maintained. The length of the period with elevated body mass tripled but was partly compensated by a shortening of the duration of wing molt (which never coincided with high body masses). Nevertheless, total cycle lengths were > 13 months. Perhaps most interestingly, under constant photoperiod, there was evidence that two components of what is normally considered an integrated phenotypic event, the prebasic molts of contour and wing feathers, were desynchronized. This suggests that the underlying organizational structure of traits is modular to some extent. Such modularity would increase the flexibility and versatility of the cyclic phenotype in evolutionary contexts.

Heavy physical work can result in physiological stress and suppressed immune function. Accordingly, long-distance migrant birds that fly for thousands of km within days can be expected to show immunosuppression, and hence be more vulnerable to infections en route. The red knot Calidris canutus Linnaeus is a long-distance migrant shorebird. We flew red knots the equivalent of 1500 km over 6 days in a wind tunnel. The humoral and cell-mediated immune responses of the flyers were compared to those of non-flying controls. Humoral immunity was measured as antibody production against injected diphtheria and tetanus antigens, and cell-mediated response as phytohemagglutinin-induced wing-web swelling. Blood corticosterone levels, which may modulate immune function, were measured in parallel. The long flights had no detectable effects on humoral or cell-mediated immune responses, or on corticosterone levels. Thus, flight performance per se may not be particularly stressful or immunosuppressive in red knots. Some birds assigned as flyers refused to fly for extended periods. Before flights started, these non-flyers had significantly lower antibody responses against tetanus than the birds that carried out the full flight program. This suggests that only birds in good physical condition may be willing to take on heavy exercise. We conclude that these long-distance migrants appear well adapted to the work load induced by long flights, enabling them to cope with long flight distances without increased stress levels and suppression of immunity. Whether this also applies in the wild, where the migrating birds may face adverse weather and food conditions, remains to be investigated.

Birds living in seasonal environments change physiology and behavior in correspondence to temporally changing environmental supplies, demands and opportunities. We recently reported that the chemical composition of uropygial gland secretions of sandpipers (Scolopacidae, order Charadriformes) changes during the breeding season from mixtures of monoesters to diesters, which fulfill specific functions related to incubation. A proper temporal match between the expression of diester preen waxes and incubation requires a flexible organization of the trait. Here we analyze the possible degrees of flexibility with reference to the functionality of better-understood molt and body mass cycles of free-living and captive red knots (Calidris canutus). The relative flexibility of seasonal cycles in preen wax composition was examined by two experimental perturbations: (1) giving birds restricted access to food and (2) monitoring them long-term under a constant photoperiodic regime. We found that wax type cannot change instantaneously, but that changing the type of wax is under similar organizational time constraints as the replacement of feathers. Just as molt and mass cycles, the seasonal rhythm of diester secretion appeared to be under endogenous control: most birds placed in a constant photoperiod still maintained seasonally changing preen waxes. Diester preen wax secretion was synchronized with the peak in body mass in spring, but became less well expressed under constant photoperiodic conditions and when food availability was limited.

Effective conservation of endangered species requires a solid understanding of the demographic causes of population change. Bird populations breeding on agricultural grasslands have declined because their preferred habitat of herb-rich meadows has been replaced by grassland monocultures. The timing of agricultural activities in these monocultural grasslands is critical, as they often coincide with the nesting phase of breeding birds. Here, we aim to identify the effect of habitat management and targeted nest protection on nest survival of Black-tailed Godwits Limosa limosa in the Netherlands, a population that has shown a 70% reduction in breeding population size since the 1970s. To protect nests in monocultures from destruction, farmers are paid to either delay mowing or leave a patch of unmown grass around the nest, a patch which in practice varied in size. In herb-rich meadows, which are typically managed for bird conservation purposes, mowing occurs after hatching. Nest survival declined as the season advanced, more steeply on monocultures than on meadows. Targeted nest protection was only partially successful, as nest predation was considerably higher on mown grassland monocultures with small unmown patches around the nest than in mown monocultures with large unmown patches and in unmown fields. Increased predator densities over the years have been suggested as an important cause of the trend towards lower nest survival, but here we show that nest survival was higher on herb-rich meadows than on monocultures, and similar to the 1980s. It thus seems that increased predator densities are an increased threat during the egg stage only if habitat quality is low. High-quality habitat in the form of herb-rich meadows therefore provides a degree of protection against predators.

Lay Summary It is believed that a bird’s energetic reserves determine when and for how long it incubates its eggs. We challenge this view for species where both parents incubate. We experimentally reduced the energetic demands of incubation by heating and insulating the nest. These treatments had no major effect on the length of incubation bouts., Incubation is energetically demanding, but it is debated whether these demands constrain incubation-scheduling (i.e., the length, constancy, and timing of incubation bouts) in cases where both parents incubate. Using 2 methods, we experimentally reduced the energetic demands of incubation in the semipalmated sandpiper, a biparental shorebird breeding in the harsh conditions of the high Arctic. First, we decreased the demands of incubation for 1 parent only by exchanging 1 of the 4 eggs for an artificial egg that heated up when the focal bird incubated. Second, we reanalyzed the data from the only published experimental study that has explicitly tested energetic constraints on incubation-scheduling in a biparentally incubating species (Cresswell et al. 2003). In this experiment, the energetic demands of incubation were decreased for both parents by insulating the nest cup. We expected that the treated birds, in both experiments, would change the length of their incubation bouts, if biparental incubation-scheduling is energetically constrained. However, we found no evidence that heating or insulation of the nest affected the length of incubation bouts: the combined effect of both experiments was an increase in bout length of 3.6min (95% CI: −33 to 40), which is equivalent to a 0.5% increase in the length of the average incubation bout. These results demonstrate that the observed biparental incubation-scheduling in semipalmated sandpipers is not primarily driven by energetic constraints and therefore by the state of the incubating bird, implying that we still do not understand the factors driving biparental incubation-scheduling.

Extreme weather events have the potential to alter both short- and long-term population dynamics as well as community- and ecosystem-level function. Such events are rare and stochastic, making it difficult to fully document how organisms respond to them and predict the repercussions of similar events in the future. To improve our understanding of the mechanisms by which short-term events can incur long-term consequences, we documented the behavioural responses and fitness consequences for a long-distance migratory bird, the continental black-tailed godwit Limosa limosa limosa, resulting from a spring snowstorm and three-week period of record low temperatures. The event caused measurable responses at three spatial scales - continental, regional and local - including migratory delays (+19 days), reverse migrations (>90 km), elevated metabolic costs (+8·8% maintenance metabolic rate) and increased foraging rates (+37%). There were few long-term fitness consequences, however, and subsequent breeding seasons instead witnessed high levels of reproductive success and little evidence of carry-over effects. This suggests that populations with continued access to food, behavioural flexibility and time to dissipate the costs of the event can likely withstand the consequences of an extreme weather event. For populations constrained in one of these respects, though, extreme events may entail extreme ecological consequences.

Mechanistic insights and predictive understanding of the spatial distributions of foragers are typically derived by fitting either field measurements on intake rates and food abundance, or observations from controlled experiments, to functional response models. It has remained unclear, however, whether and why one approach should be favoured above the other, as direct comparative studies are rare. The field measurements required to parameterize either single or multi-species functional response models are relatively easy to obtain, except at sites with low food densities and at places with high food densities, as the former will be avoided and the second will be rare. Also, in foragers facing a digestive bottleneck, intake rates (calculated over total time) will be constant over a wide range of food densities. In addition, interference effects may depress intake rates further. All of this hinders the appropriate estimation of parameters such as the 'instantaneous area of discovery' and the handling time, using a type II functional response model also known as 'Holling's disc equation'. Here we compare field- and controlled experimental measurements of intake rate as a function of food abundance in female bar-tailed godwits Limosa lapponica feeding on lugworms Arenicola marina. We show that a fit of the type II functional response model to field measurements predicts lower intake rates (about 2·5 times), longer handling times (about 4 times) and lower 'instantaneous areas of discovery' (about 30-70 times), compared with measurements from controlled experimental conditions. In agreement with the assumptions of Holling's disc equation, under controlled experimental settings both the instantaneous area of discovery and the handling time remained constant with an increase in food density. The field data, however, would lead us to conclude that although handling time remains constant, the instantaneous area of discovery decreased with increasing prey densities. This will result into highly underestimated sensory capacities when using field data. Our results demonstrate that the elucidation of the fundamental mechanisms behind prey detection and prey processing capacities of a species necessitates measurements of functional response functions under the whole range of prey densities on solitary feeding individuals, which is only possible under controlled conditions. Field measurements yield 'consistency tests' of the distributional patterns in a specific ecological context.

To cope with changes in the environment, organisms not only show behavioural but also phenotypic adjustments. This is well established for the digestive tract. Here we present a first case of birds adjusting their flight machinery in response to predation risk. In an indoor experiment, ruddy turnstones Arenaria interpres were subjected to an unpredictable daily appearance of either a raptor or a small gull (as a control). Ruddy turnstones experiencing threat induced by a flying raptor model, longer than after similar passage by the gull model, refrained from feeding after this disturbance. Pectoral muscle mass, but not lean mass, responded in a course of a few days to changes in the perceived threat of predation. Pectoral muscle mass increased after raptor scares. Taking the small increases in body mass into account, pectoral muscle mass was 3.6% higher than aerodynamically predicted for constant flight performance. This demonstrates that perceived risk factors may directly affect organ size.

Most populations of migrant shorebirds around the world are in serious decline, suggesting that vital condition-dependent rates such as fecundity and annual survival are being affected globally. A striking example is the red knot (Calidris canutus rufa) population wintering in Tierra del Fuego, which undertakes marathon 30 000 km hemispheric migrations annually. In spring, migrant birds forage voraciously on horseshoe crab eggs in Delaware Bay in the eastern USA before departing to breed in Arctic polar deserts. From 1997 to 2002 an increasing proportion of knots failed to reach threshold departure masses of 180-200 g, possibly because of later arrival in the Bay and food shortage from concurrent over-harvesting of crabs. Reduced nutrient storage, especially in late-arriving birds, possibly combined with reduced sizes of intestine and liver during refuelling, had severe fitness consequences for adult survival and recruitment of young in 2000-2002. From 1997 to 2002 known survivors in Delaware Bay were heavier at initial capture than birds never seen again, annual survival of adults decreased by 37% between May 2000 and May 2001, and the number of second-year birds in wintering flocks declined by 47%. Population size in Tierra del Fuego declined alarmingly from 51 000 to 27 000 in 2000-2002, seriously threatening the viability of this subspecies. Demographic modelling predicts imminent endangerment and an increased risk of extinction of the subspecies without urgent risk-averse management.