Search

High‐quality staging sites are critical for long‐distance migratory shorebirds to rest and refuel but are under threat from human development, including expansion of wind energy projects. However, predicting migration timing and movements in relation to weather conditions at staging sites can increase our understanding and mitigate effects of wind turbine collisions. Here we assessed northward migration timing and orientation in relation to environmental conditions at an inland staging area in Saskatchewan, Canada, with active and proposed wind energy developments. The area is known to host ~25% of North America's Sanderling Calidris alba population and 16 other Arctic‐breeding migrant shorebird species. We quantified arrival and departure time of day in relation to weather using data from 140 of 237 Sanderlings radiotagged locally and at a southern staging site in the Gulf of Mexico with the Motus Wildlife Tracking System (April–June, 2015–2017). Although Sanderling arrival times were not related to time of day or weather, departures were more likely at sunset in winds blowing towards the northwest at intermediate speeds (

Changes in demographic rates underpin changes in population size, and understanding demographic rates can greatly aid the design and development of strategies to maintain populations in the face of environmental changes. However, acquiring estimates of demographic parameters at relevant spatial scales is difficult. Measures of annual survival rates can be particularly challenging to obtain because large-scale, long-term tracking of individuals is difficult and the resulting data contain many inherent biases. In recent years, advances in both tracking and analytical techniques have meant that, for some taxonomic groups, sufficient numbers of survival estimates are available to allow variation within and among species to be explored. Here we review published estimates of annual adult survival rates in shorebird species across the globe, and construct models to explore the phylogenetic, geographical, seasonal and sex-based variation in survival rates. Models of 295 survival estimates from 56 species show that survival rates calculated from recoveries of dead individuals or from return rates of marked individuals are significantly lower than estimates from mark–recapture models. Survival rates also vary across flyways, largely as a consequence of differences in the genera that have been studied and the analytical methods used, with published estimates from the Americas and from smaller shorebirds (Actitis, Calidris and Charadrius spp.) tending to be underestimated. By incorporating the analytical method used to generate each estimate within a mixed model framework, we provide method-corrected species-specific and genus-specific adult annual survival estimates for 52 species of 15 genera.

10 pages; International audience; Birds exhibit a wide diversity of breeding strategies. During incubation or chick-rearing, parental care can be either uniparental, by either the male or the female, or biparental. Understanding the selective pressures that drive these different strategies represents an exciting challenge for ecologists. In this context, assigning the type of parental care at the nest (e.g. biparental or uniparental incubation strategy) is often a prerequisite to answering questions in evolutionary ecology. The aim of this study was to produce a standardized method unequivocally to assign an incubation strategy to any Sanderling Calidris alba nest found in the field by monitoring nest temperature profiles. Using drops of >3 °C in nest temperature (recorded with thermistors) to distinguish incubation and recess periods, we showed that the number of recesses and the total duration of these recesses from 09:00 to 17:00 h UTC allowed us reliably (99.1% after 24 h and 100% when monitoring the nest for at least 4 days) to assign the incubation strategy at the nest for 21 breeding adults (14 nests). Monitoring nest temperature for at least 24 h is an effective method to assign an incubation strategy without having to re-visit nests, thereby saving time in the field and minimizing both disturbance and related increase in predation risk of clutches. Given the advantages of our method, we suggest that it should be used more widely in studies that aim to document incubation strategies and patterns in regions where ambient temperatures are at least 3 °C below the median nest temperature.

Biparental incubation is a form of cooperation between parents, but it is not conflict-free because parents trade off incubation against other activities (e.g. self-maintenance, mating opportunities). How parents resolve such conflict and achieve cooperation remains unknown. To understand better the potential for conflict, cooperation and the constraints on incubation behaviour, investigation of the parents' behaviour, both during incubation and when they are off incubation-duty, is necessary. Using a combination of automated incubation-monitoring and radiotelemetry we simultaneously investigated the behaviours of both parents in the biparentally incubating Semipalmated Sandpiper Calidris pusilla, a shorebird breeding under continuous daylight in the high Arctic. Here, we describe the off-nest behaviour of 32 off-duty parents from 17 nests. Off-duty parents roamed on average 224 m from their nest, implying that direct communication with the incubating partner is unlikely. On average, off-duty parents spent only 59% of their time feeding. Off-nest distance and behaviour (like previously reported incubation behaviour) differed between the sexes, and varied with time and weather. Males roamed less far from the nest and spent less time feeding than did females. At night, parents stayed closer to the nest and tended to spend less time feeding than during the day. Further exploratory analyses revealed that the time spent feeding increased over the incubation period, and that at night, but not during the day, off-duty parents spent more time feeding under relatively windy conditions. Hence, under energetically stressful conditions, parents may be forced to feed more. Our results suggest that parents are likely to conflict over the favourable feeding times, i.e. over when to incubate (within a day or incubation period). Our study also indicates that Semipalmated Sandpiper parents do not continuously keep track of each other to optimize incubation scheduling and, hence, that the off-duty parent's decision to remain closer to the nest drives the length of incubation bouts.

With only a single extant representative, endemic to the Tuamotu Archipelago, the Polynesian sandpipers (Aechmorhynchus and Prosobonia) may have had a larger distribution in Eastern Polynesia in the past, with four endemic taxa. Although these aberrant sandpipers' membership to the Scolocapidae has been well supported, finding their closest living taxa has proved difficult and the phylogenetic relationships of these taxa have remained unresolved. We present the first molecular analysis of the Polynesian sandpipers, including sampling of the only known specimen of the extinct Prosobonia leucoptera, collected in 1773. Based on mitochondrial and nuclear gene sequence data, the phylogenetic analyses demonstrate that the Polynesian sandpipers were sister taxa and belonged to the clade that included the other sandpipers (Calidris and allies) and turnstones (Arenaria), although without a close relative among extant genera. Divergence time estimates suggested that the lineage leading to Prosobonia diverged from the other extant sandpipers during the Oligocene and that either the Line Islands or the Tuamotu Archipelago were probably the first archipelagos colonized by the Prosobonia lineage. On the basis of these results, we suggest that Aechmorhynchus parvirostris and Prosobonia leucoptera be regarded as related species within the same genus, and thus that the senior name Prosobonia be used for both taxa.

We measured deuterium isotope ratios (δDf) in primary feathers to distinguish first-year from older Pectoral Sandpipers Calidris melanotos captured in Barrow, Alaska, during the breeding season. δDf showed a distinct bimodal distribution, and model-based clustering placed the δDf values into two non-overlapping groups. More negative δDf corresponded to Arctic areas, probably identifying first-year birds with Arctic-grown juvenile feathers retained from the previous year. The more positive values corresponded to lower latitudes, possibly identifying older birds that grew their feathers at non-Arctic latitudes.

Incubation can be as energetically costly as rearing chicks(Williams 1991, Tatner & Bryant 1993, Thomson et al.1998a) and might even limit clutch size (Heaney M Thomson et al. 1998b).In Semipalmated Sandpipers Calidris pusilla, anotherbiparental incubator, parents increased incubation by10% when heat loss from eggs was reduced by insulat-ing the nest (Cresswell et al. 2003). These results weresimilar to those for uniparental incubators (Thomsonet al. 1998a, Bryan & Bryant 1999), suggesting biparen-tal incubating species might be as constrained byenergy availability as uniparental incubators. However,few data exist on the incubation strategies of biparen-tal incubating passerines in response to changes in foodavailability.In this study, we assessed the effect of food supple-mentation on incubation behaviour of the biparentalSilvereye Zosterops lateralis. We provided a food supple-ment to free-living Silvereyes and compared the incuba-tion attentiveness of adults before and after they weresupplemented. Many studies show that food supplemen-tation can increase attentiveness (e.g. Nilsson & Smith1988, Eikenaar et al. 2003, Gorman & Nager 2003,Pearse et al. 2004, Chalfoun & Martin 2007, but seeMoreno 1989, Mackintosh & Briskie 2005), but fewhave determined how individual incubation behaviourchanges in response to short-term food supplementation.We used a repeated-measures approach in which incuba-tion behaviour of individual birds was compared beforeand after they received supplementary food. Our resultsshow that Silvereyes increased their attentiveness andother measures of incubation investment in response tofood supplementation.METHODSStudy site and subjectsThe study was conducted during spring 2000 and 2001 atthe Kowhai Bush Reserve (173 36 ¢E, 42 23 S)near Kaikoura, New Zealand. The Silvereye is a small(11–14 g) passerine that ranges throughout Australia andthe South Pacific. The species is insectivorous, but con-sumes nectar and fruit when available. Sexes are similarin size and colouration and both males and females incu-bate, brood and feed nestlings (Heather & Robertson1996). Nests are spaced 10–30 m apart but adults arenon-territorial and only defend an area c. 1 m around thenest. Females alone build a well-concealed cup nest in thelower canopy in which they lay two to four eggs. Incuba-tion begins once the clutch is complete and nestlingshatch about 11 days later (Heather & Robertson 1996).We measured changes in incubation behaviour withfood supplementation at 10 nests. To avoid pseudorepli-cation, we only sampled nests once and did not samplenests of individuals used in previous experiments. Obser-vations of colour-ringed Silvereyes indicate that theyre-nest near previous breeding attempts and thus we didnot use any nest found in the same general area( 50 m) as a previously studied nest.

We describe the migration, biometrics and moult of Red Knot Calidris canutus canutus in southern Africa and compare them with the biometrics and moult of Calidris canutus islandica in northern Europe to examine possible adaptations to different environments during the non-breeding season. Northward and southward migration of C. c. canutus took place along the coast of Western Europe and there was one recovery in West Africa (Mauritania), suggesting a coastal migration round West Africa rather than migration across the Sahara, as recorded in other waders. Adult Knots in South Africa had no additional fattening in November-January (fat index of 7%), in contrast to C. c. islandica wintering in Britain. This is consistent with the theory that extra fat is required only where food shortages are likely. The bills of canutus were longer than those of islandica but their wings were shorter, confirming the sub-specific assignments and origin of this population. The average duration of primary moult in South Africa was 95 days, shorter than that of other Arctic-breeding waders that moult in South Africa, but longer than of islandica moulting in Scotland (77 days). Mean starting and completion dates were 20 July and 5 October for islandica and 25 October and 28 January for canutus. The timing and duration of primary moult for these two subspecies suggest that waders need to complete moult before the northern winter when food supplies are limited, whilst waders in benign climates face no such pressures. First-year canutus either retained old primaries for much of their first year or had a partial moult of inner or other primaries. Adults departed on northward migration in mid-April, having attained a mean departure mass of c. 190 g (maximum 232 g). The mean fat index at this time was 24% (maximum 29%) and the fat-free flight muscle mass increased. The predicted flight range of 4000 km falls short of the distance to the first likely refuelling site in West Africa, suggesting that birds rely on assistance from favourable winds.

Parental brood attendance patterns vary greatly among shorebird species. For monogamous calidridine species, biparental care with female-first brood departure is most common. It is believed that adult sandpipers balance potential individual survival costs associated with extended parental care against the benefit gained by their brood of prolonged parental care. These costs and benefits are difficult to quantify and factors affecting the termination of parental brood attendance are unclear. We compared clutch size, nesting phenology, and parental attendance patterns of Western Sandpipers Calidris mauri at Nome and Kanaryarmiut, Alaska, sites separated by three degrees of latitude. The sites differed in breeding density and duration of breeding season, but the distribution of clutch sizes did not differ between sites or between nesting attempts. Parental attendance patterns were similar between sites, suggesting that parental attendance is a highly conserved life-history trait in Western Sandpipers. Male Western Sandpipers attended broods longer than females, and the duration of parental attendance decreased at a similar rate for both sexes as the season progressed. Male and female Western Sandpipers undertake differential migrations to their non-breeding grounds, with males typically settling at more northerly locations and females at more southerly sites, a migration pattern shared by certain other monogamous calidridine species. These same species exhibit similar parental brood attendance patterns, suggesting the strong role of overall migration distance in shaping the expression of parental attendance behaviours. A contrast of more geographically disjunct sites coupled with a better understanding of the migratory connectivity between Western Sandpiper breeding and non-breeding populations would elucidate the role of cross-seasonal effects on parental brood attendance decisions.

Predators may influence many aspects of the daily life and seasonal movements of their prey. Here we quantify direct, and evaluate indirect effects of predation by three falcon species (Lanner Falcon Falco biarmicus, Barbary Falcon Falco pelegrinoides and Peregrine Falcon Falco peregrinus) on coastal shorebirds wintering on the Banc d’Arguin, Mauritania, an area hosting approximately 30% of the East Atlantic Flyway population of shorebirds. On the basis of 754 h of observation over five winters, 97 witnessed attacks and 585 collected prey remains, we show that shorebirds were safer in larger flocks, which tended to be attacked less often. Furthermore, species that forage relatively close to shore and in small flocks were depredated more often than expected from their relative abundance. In three species, Red Knot Calidris canutus canutus, Bar-tailed Godwit Limosa lapponica taymyrensis and Dunlin Calidris alpina, the juveniles were more vulnerable than adults. We estimated that on average 1% of the juvenile and 0.1% of the adult Red Knots present were killed by large falcons each winter. For Red Knots we simultaneously quantified annual survival on the basis of an individual colour-marking programme: mortality due to predation by falcons accounted for an estimated 6.2% (juveniles) and 0.8% (adults) of annual mortality. We suggest that juvenile Red Knots are 10 times as likely to be killed by falcons because they use riskier habitats, i.e. early and late tide foraging areas closer to shores where surprise attacks are both more common and more successful. These results indicate that the strength of indirect effects of predation operating in a shorebird population largely outweigh the effects of mortality per se.

The Purple Sandpiper Calidris maritima, an Arctic shorebird, displays unusual sex roles during breeding. In a 5-year study in Svalbard, in the Norwegian high Arctic, data were collected on the roles of the sexes in this species. Purple Sandpipers were similar to most shorebirds in that males actively courted females, established territories and vigorously defended their territories from intruders. Both sexes shared incubation duties approximately equally over the entire 21-day incubation period. Males incubated very little initially, but increased their effort significantly during the first 11 days of incubation to over 50% of the time from days 11–21. However, Purple Sandpipers contrasted with most other shorebird species in that females discontinued their breeding efforts at hatching. In nearly all cases, broods were attended solely by the male until the chicks reached fledging age (or even longer). Nonetheless, the pair bond must be described as monogamous because neither males nor females were found to re-mate or lay a second clutch during a season. This pattern of parental care, which is found only in very few other shorebird species, is discussed in an evolutionary context.

Western Sandpipers Calidris mauri are the most numerous shorebird species in the San Francisco Bay estuary during winter. A sample of 106 Western Sandpipers was captured in mist nets and radio-marked with 1-g transmitters to examine their wintering site fidelity and movements. Differences in distances moved, home range extent and core area size were examined by age, sex, season, site, time of day and tide. All birds remained in the south San Francisco Bay region during winter and exhibited strong site fidelity, with a mean home range of 22.0 km2 or only 8% of the study area. First-year birds had larger home ranges (26.6 ± 3.6 km2) than adults (17.2 ± 2.5 km2) in winter, but home range sizes of males and females were not significantly different in any period. Home range sizes were similar between seasons, but core areas were smaller in spring (6.3 ± 1.2 km2) than in early (9.6 ± 4.0 km2) or late (11.6 ± 1.6 km2) winter. Movements and home range size were similar for radio-marked birds located during day and night. The high degree of regional and local site fidelity demonstrated that the mixture of natural mud fiats and artificial salt ponds in southern San Francisco Bay provided sufficient resources for large wintering populations of Western Sandpipers.

SUMMARY The results of two expeditions to northeast Greece in May 1960 and 1961 are described together with records of a number of other parties of ornithologists who have recently visited the area. Observations include the first recorded breeding of Spur-winged Plover Hoplopterus spinosus and Isabelline Wheatear Oenanthe isabellina in Greece, and of the Fan-tailed Warbler Cisticola juncidis in north Greece; also the first recorded occurrence of Knot Calidris canutus and Spectacled Warbler Sylvia conspicillata in Greece. In the systematic list many discrepancies with previously published data are recorded for this relatively remote and ornithologically little known area of Greece. Observations are made on 111 species out of 219 recorded and on the migration observed.

The extensive overlap in morphological characters between populations of Dunlin Calidris alpina imposes problems of determining the origin of migrating and wintering birds. The morphology of the birds also varies between the sexes, and the sex of a Dunlin may often be difficult to determine. To clarify if mitochondrial DNA can be used to identify which breeding areas migrating Dunlin come from, we investigated the occurrence of different mtDNA haplotypes in Dunlin from eight breeding areas on the Russian and Siberian tundra. Four haplotypes were found and at most sites more than one haplotype occurred. The European haplotype predominated in the area west of the Taymyr Peninsula, the Siberian haplotype in central Siberia (from the Taymyr Peninsula to the Lopatka Peninsula) and the Beringian haplotype in eastern Siberia. One individual of an Alaskan haplotype, not detected previously among breeding birds outside North America, was found on Wrangel Island. The sex of each bird was identified genetically and the morphology of males and females was analysed separately. Birds with the European haplotype were generally smaller than birds with the Beringian or Alaskan haplotypes. Birds possessing the Siberian haplotype showed intermediate values in most cases. After compensating for differences between sites, males with the Siberian haplotype had significantly longer bills than males having the European haplotype. Multiple regressions indicate that mitochondrial DNA analysis improves models estimating the breeding origin of migrating Dunlin.

A hierarchical modelling approach was used to examine adult and age-specific survival in an 8-year study of breeding Semipalmated Sandpipers Calidris pusilla at La Perouse Bay, Canada. The survival of adult sandpipers was best described by a model with time dependence in local survival rate and probability of recapture. Annual variation in the local survival rate of adults was not correlated with nest success, timing of breeding or “return rates” and was not biased by an effect of first capture. Local survival rate of adult females (0.56, 95% c.l. = 0.51-0.61) was consistently lower than that of adult males (0.61, 95% c.l. = 0.56-0.66); these estimates were comparable with data from other shorebirds. The survival of returning young was best fitted by a model with both age and time dependence in local survival rate and probability of recapture. We evaluated our estimates of local survival rate with reference to patterns of breeding fidelity and philopatry in Semipalmated Sandpipers and other shorebirds.

Few studies have assessed how the dynamics of wetland bird movements relate to changing resource availability, particularly at more than one spatial scale. Within western Oregon’s Williamette Valley, we examined winter resident Dunlin Calidris alpina movements in relation to a decrease in availability of preferred shorebird foraging habitat from early to late winter of 1999‐2000. By tracking movements of 15 (early winter) and 12 (late winter) radiomarked individuals, we calculated home ranges and characterized presence/absence of a preference for shorebird foraging habitat during each winter period. Between periods, we compared: (1) percentage of shorebird habitat in home ranges to its availability in the landscape (regional preference), (2) percentage of radio locations in shorebird habitat to its availability within home ranges (local preference) and (3) relative use of roost sites. Concurrent with a 75% decrease in available shorebird habitat from early to late winter, average home range sizes increased by a factor of 3.8. At a regional scale, home ranges in early winter included a significantly greater percentage of shorebird foraging habitat than was available in the wider landscape. However, by late winter, the percent of shorebird habitat in home ranges did not match availability in the landscape. At the local scale, for both winter periods Dunlin were located in shorebird foraging habitat more often than expected given availability of habitat within home ranges [Correction added after online publication 23 May 2008: sentence amended]. An increase in the number of roosts used from early to late winter implies possible reliance on additional sites in late winter for foraging opportunities. Results suggest that wet, unvegetated habitat is sought by Dunlin throughout winter, but individuals could not select home ranges in late winter that fully compensated for seasonal loss of habitat.

This paper presents descriptions of the behaviour of Knot Calidris canutus breeding in high arctic Canada. Knot nested within territories which were of little importance as a food source. Males defended territories by singing from the ground, aerial chases, threat displays (horizontal point), fighting, and low flights in which the wings were held vertically over the back (V-wing flights). Songflights performed by males seemed to serve a mate-attraction function. The commonest heterosexual display was a tail-up display, used mainly by males prior to clutch completion. Males also used a nest scrape display and a ground point display in advertising potential nest sites to females. Behaviour preceding copulation was variable but usually included tail-up displays and the male pecking the back of the female. Anti-predator behaviour when on the nest involved remaining still until a predator was very close, whereupon distraction displays were used. Both sexes incubated but only males cared for chicks. Caring for chicks involved leading chicks to favourable feeding areas, brooding and vigilance, distraction displays, and harrassment or physical attack of avian predators. The displays used by breeding Knot tend to be unlike those of most congenerics and suggest that the Knot is a rather peripheral member of Calidris.

Summary The autumn moult pattern of adult Purple Sandpipers Calidris maritima in Iceland is described. The duration of the moult was estimated to be c. 5½-7 weeks (c. 40–50 days). Females generally started moult before males and moult did not appear to overlap breeding. Information from other areas is reviewed. A mechanism by which the duration of moult is shortened amongst various species is by an increase in the number of feathers growing concurrently during the moult. Likely reasons for the placing of the moult in the annual cycle of the Purple Sandpiper are discussed, and appear to be related to the exceptionally northerly wintering distribution of the species.

Raptor predation on waders was studied by direct observation of raptors hunting a known wader population and subsequent recovery of dead waders. In each of three winters, raptor predation was shown to be the most significant cause of mortality in most small wader species, Sparrowhawks Accipiter nisus, Merlins Falco columbarius and Peregrines F. pere-grinus attacked waders with a success rate of 11.6%. 8.8% and 6.8%, respectively. Most waders attacked or found dead were Redshank Tringa totanus and Dunlin Calidris alpina; most were killed by Sparrowhawks. Kleptoparasitism of raptors carrying prey by Carrion Crows Corvus corone significantly increased the winter mortality of some waders. Redshank populations were most affected by raptor predation: over 50% of the total population (which was found to be closed during most of the winter) and over 90% of the juvenile population were taken in two winters: juvenites were more likely to be killed by raptors.

A surveillance radar in southwest Iceland was recorded by time-lapse filming in order to monitor the migration pattern of birds departing from or passing Iceland on their way to high-arctic breeding grounds in late May and early June 1988–1990. An overwhelming majority of the radar echoes from migrating bird flocks departed from Iceland but a few seemed to pass over from further south. Timing of movements and supplementary field observations indicated that mainly four species were involved, i.e. Knot Calidris canutus, Turnstone Arenaria interpres. Sanderling C. alba and Brent Goose Branta bernicla. Departures in late May from stopover sites in Iceland took place mainly in the afternoons, peaking between 1700 and 1900h in all 3 years. The departure intensity was lowest between 0100 and 1300h. Flight paths were generally straight, and the average track direction was towards the northwest (315°), suggesting that the majority of birds were heading for a transglacial migration across the Greenland icecap on their way to breeding grounds in northwest Greenland and northern Canada. Track directions varied with wind, although to a rather small degree, indicating partial drift or pseudodrift. More echoes were registered in easterly winds (tailwinds) and fewer in northerly winds than expected from random. Airspeeds were significantly slower than groundspeeds (average 17.0 and 18.7 m/s, respectively), showing that the birds more often than not benefited from tail wind assistance.

Arctic waders often build up large fat loads and complete their migratory journeys by a few long-distance flights between traditional staging sites. Optimal fat loads and choices of staging sites differ depending on whether the birds are adapted to minimize energy or time spent on migration. In the latter case, we predict that the birds will depart for the next staging site when the instantaneous speed of migration expected after arrival at the next site, exceeds the corresponding speed at the departure site. The instantaneous migration speed is a function of the rate of fat deposition and the current fat load. As a consequence of this, overloading (birds deposit larger fat loads than needed merely for covering the flight distance to the next destination) and by-passing of possible, but low-quality staging sites, are expected under specific conditions in time-selected migration. Estimates of fat deposition rates and departure fat loads were obtained by captures of Knots Calidris canutus, Sanderlings C. alba and Turnstones Arenaria interpres in W. Iceland during spring migration. Further fat deposition data referring to spring migration of these species were compiled from the literature. Fat deposition rates at different sites, as measured by the daily gain in mass relative to lean body-mass, range between 1.0 and 3.6%/day, and departure fuel loads (in % of lean body-mass) between 27 and 73%. Comparison with flight range estimates suggests that overloading may be a regular phenomenon during spring migration of Knots, Sanderlings and Turnstones. Furthermore, fat deposition rates at different staging sites, and the general difference in migration patterns between spring and autumn, indicate that by-passing of possible staging sites may well occur. Hence, it cannot be excluded that the waders' migratory habits primarily serve to maximize the overall speed of migration.

Foraging densities of Curlew Sandpipers Calidris ferruginea and Grey Plovers Pluvialis squa-tarola at the Berg River estuary, South Africa, are examined in relation to prey and habitat attributes. There is a substantial overlap in the prey species eaten by Curlew Sandpipers and Grey Plovers, but the distribution patterns of the two species within the estuary could not be explained by the same prey attributes. The distribution of tactilely foraging Curlew Sandpipers was determined largely by the numerical abundance of nereid worms. They foraged preferentialiy on muddy rather than sandy substrata, but their distribution could not be explained on the basis of substratum penetrability. The density of visually foraging Grey Plovers was correlated with the biomass, rather than abundance, of the larger of the two common nereids on the estuary. The favoured foraging area of Grey Plovers had a high vegetation cover, but their foraging dispersion was not influenced by substratum type. The energy intake rates of Curlew Sandpipers were broadly inversely related to their foraging density, although the rate of prey capture was maximal at high prey density. These birds may use neighbour success rates in selecting foraging sites. Energy intake rates of Grey Plovers were very consistent across the estuary, suggesting that individuals occupy or defend areas which provide access to comparable food resources.

Samples of Purple Sandpipers were captured around the coasts of Britain. Analysis of their bill-length distributions enabled the sex ratios and percentages of ‘long-billed’ and ‘short-billed’ birds at each locality to be estimated. The sex ratio for the ‘long-billed’ population was estimated to be one female to 2–11 males, and one female to 1 -34 males for the ‘short-billed’ population. During winter, proportionately more ‘long-billed’ birds occurred in northern and western Scotland, Wales and southern England, whilst ‘short-billed’ birds predominated from Kincardine to Yorkshire. The total sizes of the ‘long-’ and ‘short-billed7’ populations were c. 15 000 and 4000, respectively. ‘Short-billed’ birds started arriving from Norway in early July. ‘Long-billed’ birds did not arrive until late October. Their origins are as yet unknown. No annual variations in the population structure were detected.

Although Semipalmated Sandpipers Calidris pusilla are monogamous, with biparental incubation, most females (86–97%) deserted their broods to the care of their mates, 0–11 days (average 6) after their eggs hatched. Males left the brood an average of 8 days later, shortly before or after the chicks fledged. In several instances, females that deserted in one year remained with the chicks the next year, and vice versa. Females deserted chicks at nests that hatched later in the season at an earlier age than those had hatched earlier (r2> o.6). Since females appeared to have an energy deficit in at least some years, and suffered higher mortality rates than males during breeding, it is possible that females deserted broods in order to take advantage of better feeding conditions at migratory stopovers in northeastern North America early in the season. There was little evidence of higher nesting success or earlier hatching date in reuniting pairs, although if both members of a pair returned to the breeding area, 80% reunited. Increased survival of their mate may be most advantageous to males in ensuring that they obtain a female the following year.

Adult and post-natal breeding-dispersal movements of Redshank Tringa totanus, Dunlin Calidris alpina and Ringed Plover Charadrius hiaticula were quantified in west Scotland. Data from 373 returning birds marked as breeding adults and 142 returning birds marked as chicks are presented. Unlike most previous studies, this study measured dispersal movements up to 40 km and attempted to overcome distance-related biases. For each species, adult males were significantly more nest site-faithful between years than were adult females. Likewise, first-time breeding males bred significantly closer to their natal site than did females. The settlement intensity per unit area of habitat showed marked differences between species, age classes and sex in the relative attractiveness of potential areas at different distances to settling birds. Adult dispersal was negatively and significantly related to breeding success in the previous year and positively related to capture on the nest in the previous year. Dispersal movements of adult Ringed Plover breeding on unstable cultivated habitats were significantly related to habitat quality (measured in terms of nest survival) and were always to habitat of better or equal quality.

Summary. The behavioural interactions among breeding Western Sandpipers (Calidris mauri) are described, and interpreted in terms of their adaptive characteristics. Upon arrival in early spring, males disperse onto territories through mutual antagonism. Territorial advertisement and maintenance is accomplished through display flights low over the ground, which are accompanied by soft buzzy vocalisations. These characteristics of display are related to the fact that this species occurs in dense, localised breeding populations, where longdistance communication is unnecessary. Because densities are high and feeding often occurs in communal areas off territory, violations of territorial boundaries are frequent. The result is an extremely high intensity of interactions among individuals. Chasing and physical combat between males is frequent and generally un-stereotyped. Birds nesting on territories away from the feeding sites often fly high above the other territories, and in doing so avoid being chased. Western Sandpipers form monogamous pair-bonds which are maintained until the young are ready to fly. The initial association of the pair is facilitated by a strong tendency to return to the previous site and by the advertisement of the males in which a specific vocalisation is given. Once the female is present on a male's territory, she is courted persistently by him and gradually begins to participate in his scraping displays. Pair-bonds are re-enforced by the close association of the pair during the pre-nesting period and through a simultaneous preening display. An hypothesis is developed that the participation of both sexes in incubation serves primarily as a means of pair-bond maintenance, acting to keep both parents present until after the young hatch. At this time, their presence is needed to provide protection for the precocial young against predators.

Individuals within a population may vary considerably in the way they exploit available food resources. If the sexes differ in the size of their feeding apparatus, there can be differences in foraging behaviour and habitat use, hence one sex may be more susceptible to competition. We examined relationships between sexual dimorphism in bill size and foraging behaviour, and habitat and microhabitat use of non-breeding Western Sandpipers Calidris mauri at Bahia Santa Maria, northwestern Mexico. Western Sandpipers are sexually dimorphic, with females about 15% longer-billed than males. Males used a higher proportion of pecks, had a lower probing‐pecking rate, walked at higher rates, foraged at sites with lower water content, and had greater variation in foraging technique than females. Moreover, males decreased their proportion of pecks and foraged at a higher rate than females when they changed from feeding alone to feeding in flocks, suggesting a greater safety advantage or susceptibility to conspecific interference when flock feeding. We compared behaviour and microhabitat usage in three habitats: brackish flats, mangroves, and cattail marshes. Sex-related differences in foraging behaviour and microhabitat use were consistent among habitats. Birds in brackish flats and mangroves used a higher proportion of pecks, foraged at lower rates and walked at higher rates, and foraged at deeper sites, with a lower proportion of water cover, than those in cattail marshes. Sex-related differences in foraging behaviour and microhabitat should reduce the level of competition between sexes, and may account for sex differences in Western Sandpiper distribution observed between habitats in Bahia Santa Maria.

Previous studies have shown that preen wax composition in some sandpipers shifts from the usual monoesters to diesters during the breeding season, possibly to reduce the ability of mammalian predators to find nests using olfactory cues. To investigate further the relationship between incubation and wax secretion, we examined seven sandpiper species with different incubation patterns (species in which both sexes incubate, in which only males incubate and in which only females incubate). During the breeding period, diester preen wax was secreted almost exclusively by the incubating sex in species with uniparental incubation, and by both sexes in species with biparental incubation. These findings suggest that diester preen waxes have a function that is directly related to incubation. Unexpectedly, in female-incubating Curlew Sandpiper Calidris ferruginea and Buff-breasted Sandpiper Tryngites subruficollis , some males also secreted diester preen waxes during the breeding period. This suggests that some males may in fact incubate, that these waxes may be a remnant from their evolutionary past when both sexes incubated, or that males need to be olfactorally cryptic because they are involved in the making of nest scrapes. The seasonal pattern of preen wax composition was also studied in captive male, female and female-mimicking male (‘faeder’) Ruff Philomachus pugnax . Captive female Ruff changed preen wax composition from monoesters to diesters in the spring despite the fact that no incubation took place. This suggests that circannual rhythms rather than actual incubation behaviour may trigger the shift to diester waxes. All captive male Ruff, including the faeders, continued to secrete monoesters, supporting the hypothesis that only the incubating sex secretes diesters.

Previous work has shown that Golden Plovers Pluvialis apricaria guarding chicks utilize heavily disturbed habitat at a lower rate than surrounding areas, but that such rates of avoidance are reduced when the movement of people is restricted to a surfaced footpath. However, there remained some unanswered questions, which are addressed in this paper. First, we examine to what extent habitat avoidance is dependent upon visitor numbers, and find no evidence that Golden Plovers avoided disturbed areas at a site where visitor pressure was half that previously studied. Secondly, we examine whether these relationships between habitat occupancy and disturbance apply to other upland waders and find that Dunlin Calidris alpina habitat utilization in disturbed areas showed a non-significant increase of approximately 50% following the provision of a surfaced footpath, in a manner similar to that observed for Golden Plover. Thirdly, we examine whether the large numbers of visitors (120 per weekend day) using the surfaced footpath impact on Golden Plover breeding success, despite the lack of habitat avoidance. There was no evidence that nest location, clutch survival or chick growth rates were reduced close to the footpath. Together, these results suggest that high levels of disturbance can impact upon habitat usage by upland waders, but only in limited circumstances where visitor pressure is very high (greater than at least 30 visitors per weekend day). However, access to such areas can be permitted for large numbers of visitors without impacting upon wader reproductive performance through the provision of a well-surfaced route.

Measures of annual breeding success are an important component of species monitoring programmes. It has been suggested that effective monitoring of breeding productivity for arctic breeding waders may be achieved from an analysis of annual variation in the proportion of juveniles in winter flocks. Here, we attempt to generate a recruitment index for Dunlin Calidris alpina caught during the winter in north Wales. This index revealed significant annual variation and we show that this is strongly correlated with summer temperature (but not rainfall) on the breeding grounds. Years with high recruitment were also correlated with increases in the national winter population estimate. In years of intermediate summer temperature, the recruitment index was highest and we discuss the possible implications this has for Dunlin under scenarios of future climate change. We were unable to generate a significant index for Common Redshank Tringa totanus and discuss possible reasons for this.

We studied phenology, staging time and refuelling in Broad-billed Sandpipers Limicola falcinellus stopping over during spring migration in the Sivash (Black Sea, Ukraine) in May 1991-94. In the study area, peak staging numbers of 2000-2500 individuals occurred in the third week of May. In May 1993, 460 birds were marked with a yellow dye and 126 of these were colour-ringed. Before 28 May no departure of birds dyed yellow could be detected; by 3 June all birds had departed. Colour-ringed adults in mid May 1993 staged for a minimum of 8.2 days. After the observed departure of large flocks (24 May and later) the staging time of colour-ringed birds decreased significantly with body mass at the time of capture. Of birds mist-netted in 1991-94, 99.3% were in full summer plumage and 89% were adults. In second-year birds, fuel deposition rate (measured between individuals) was 0.44 g/day. In adults caught from early May to 24 May, overall fuel deposition rate was 1.04 g/day (3.4% of lean body mass). Mean adult body mass in early May was 34.8 g, increasing to 45.5 g after 24 May. Estimated body mass at departure was 51 g. Departure body mass and flight range estimates suggest that although birds refuelled quickly, fuel loads are only just sufficient for an unbroken flight to Scandinavia and the Kola Peninsula. We suggest that Broad-billed Sandpipers use the Sivash as a crucial final take-off stopover site, and that they follow a 'jumping' migration strategy, performed under narrow time constraints.

We studied phenology, staging time and refuelling in Broad-billed Sandpipers Limicola falcinellus stopping over during spring migration in the Sivash (Black Sea, Ukraine) in May 1991-94. In the study area, peak staging numbers of 2000-2500 individuals occurred in the third week of May. In May 1993, 460 birds were marked with a yellow dye and 126 of these were colour-ringed. Before 28 May no departure of birds dyed yellow could be detected; by 3 June all birds had departed. Colour-ringed adults in mid May 1993 staged for a minimum of 8.2 days. After the observed departure of large flocks (24 May and later) the staging time of colour-ringed birds decreased significantly with body mass at the time of capture. Of birds mist-netted in 1991-94, 99.3% were in full summer plumage and 89% were adults. In second-year birds, fuel deposition rate (measured between individuals) was 0.44 g/day. In adults caught from early May to 24 May, overall fuel deposition rate was 1.04 g/day (3.4% of lean body mass). Mean adult body mass in early May was 34.8 g, increasing to 45.5 g after 24 May. Estimated body mass at departure was 51 g. Departure body mass and flight range estimates suggest that although birds refuelled quickly, fuel loads are only just sufficient for an unbroken flight to Scandinavia and the Kola Peninsula. We suggest that Broad-billed Sandpipers use the Sivash as a crucial final take-off stopover site, and that they follow a 'jumping' migration strategy, performed under narrow time constraints.

For long-distance migrants, such as many of the shorebirds, understanding the demographic implications of behavioural strategies adopted by individuals is key to understanding how environmental change will affect populations. Stable isotopes have been used in the terrestrial environment to infer migratory strategies of birds but rarely in marine or estuarine systems. Here, we show that the stable isotope ratios of carbon and nitrogen in flight feathers can be used to identify at least three discrete wintering areas of the Red Knot Calidris canutus on the eastern seaboard of the Americas, ranging from southeastern USA to Patagonia and Tierra del Fuego. In spring, birds migrate northwards via Delaware Bay, in the northeastern USA, the last stopping point before arrival in Arctic breeding areas, where they fatten up on eggs of spawning Horseshoe Crabs Limulus polyphemus . The isotope ratios of feather samples taken from birds caught in the Bay during May 2003 were compared with feathers obtained from known wintering areas in Florida (USA), Bahia Lomas (Chile) and Rio Grande (Argentina). In May 2003, 30% of birds passing through the Bay had Florida-type ‘signatures’, 58% were Bahia Lomas-type, 6% were Rio Grande-type and 7% were unclassified. Some of the southern wintering birds had started moulting flight feathers in northern areas, suspended this, and then finished their moult in the wintering areas, whereas others flew straight to the wintering areas before commencing moult. This study shows that stable isotopes can be used to infer migratory strategies of coastal-feeding shorebirds and provides the basis for identifying the moult strategy and wintering areas of birds passing through Delaware Bay. Coupled with banding and marking birds as individuals, stable isotopes provide a powerful tool for estimating population-specific demographic parameters and, in this case, further our understanding of the migration systems of the declining Nearctic populations of Red Knot.

With current losses of saltmarsh running at > 100 ha per year in the UK, creation of new intertidal habitats through managed realignment is likely to be increasingly used. Potentially, this has biodiversity as well as engineering benefits. However, assessing the conservation value of many of the current UK schemes is difficult as the biological monitoring has been generally poor, with a few notable exceptions. At the Tollesbury and Orplands realignment sites, Essex, bird communities were dominated by terrestrial species during the first year of inundation and waterbird communities rapidly developed during the second and third years. Five years after the initial breach in the sea wall, communities were similar to surrounding mudflats but with some notable exceptions. Dunlin Calidris alpina and Common Redshank Tringa totanus that prey on the early colonizing Nereis and Hydrobia used the sites in the first 2 years. Eurasian Oystercatcher Haematopus ostralegus did not occur on the realignment site as there were no large bivalves, whereas Red Knot Calidris canutus used the site after 4–5 years coincidentally with the appearance of Macoma balthica. The differences in the bird communities occurred because UK sites are often small, enclosed and poorly drained. If at a suitable height in the tidal frame, UK managed realignment sites are successful in that they have developed saltmarsh and biologically active mudflats but they may lack the full range of biodiversity found in surrounding natural intertidal habitats, even decades after inundation.

The relationship between weather and wader distributions on non-estuarine coasts is described and used to predict how future wader distributions may respond to climate change. The distributions of eight out of nine species of wader commonly wintering on the non-estuarine coasts of Britain altered between two similar surveys, in 1984/85 and 1997/ 98, that covered 78% and 38% of Britain’s 12 594 km of non-estuarine coastline, respectively. These eight species moved at least in part either eastwards along the winter isotherms or northwards. These changes in distribution broadly coincide with a distributional shift towards the species’ respective breeding grounds and are correlated with the local winter weather over the period: increasingly mild extreme temperatures and changes in mean rainfall, mean wind speed and wind-chill. Based on the scenarios for Britain’s climate in 2020 and 2080, it is predicted that the distributions of the waders will move away from the west. The non-estuarine coasts of Britain hold particularly high proportions of the international flyway populations of Ringed Plover Charadrius hiaticula , Sanderling Calidris alba , Purple Sandpiper Calidris maritima and Ruddy Turnstone Arenaria interpres that are all expected to show continuing decline to 2080. Overwintering waders appear to be good indicators of the effects of climate change.

We studied the relationships between the habitat use of migratory shorebirds and the spatial distributions of the Southwestern Atlantic Fiddler Crab Uca uruguayensis, polychaetes, sediment characteristics and tidal levels in the Rio de La Plata estuary, Argentina, where U. uruguayensis is one of the most important intertidal species. Crabs have a well-defined patchy distribution that is segregated spatially from that of polychaetes. Crab density on the surface varied across the tidal cycle, reaching maximum values during low tide. Polychaete density decreased with depth but showed no change through the tidal cycle; however, given that sediment penetrability did change during the same period, their availability to probing shorebirds is expected to change. Habitat use by shorebirds followed the spatial distribution of prey; shorebirds that foraged on polychaetes (White-rumped Sandpiper Calidris fuscicollis, Two-banded Plover Charadrius falklandicus and Hudsonian Godwit Limosa haemastica) focused their attention on the areas with the highest densities of polychaetes, whereas species that preyed mostly on crabs (Ruddy Turnstone Arenaria interpres, Whimbrel Numenius phaeopus and Grey Plover Pluvialis squatarola) predominantly used areas with crabs. This segregation occurred particularly during low tide, a period in which polychaetes became fully available. Results show that the spatial and temporal heterogeneity of mudflats in relation to the types and availability of prey has a strong effect on shorebird habitat use.

Timing and duration of primary moult in three populations of Purple Sandpipers Calidris maritima were described and discussed in relation to the birds' need to complete moult before the onset of winter, when resources are required for survival. We predicted that moult would be completed earlier by birds wintering at higher latitudes. The south Norwegian breeding population, which moults and winters along the coast of east Britain (54-57°N) had a mean starting date of 21 July for primary moult (16 July for females and 24 July for males), a mean duration of 61 days, and completed on 20 September. Resident Icelandic (64-65°N) birds had a mean starting date of 22 July for primary moult (17 July for females and 25 July for males), a mean duration of 51 days, and completed on 11 September. Birds moulting in north Norway (70°N) arrived in north Norway in suspended primary moult or without having started moult, and completed it there. They had a mean completion date of 2 November for primary moult (31 October for females and 3 November for males). Starting date and duration could not be estimated because some suspended moult for an undetermined period, but it was thought that they started in late August. It is likely that most originated from Russia. The onset of moult appears to be set by the end of breeding and there is little overlap in these two events. The earlier start of moult by females in all three populations may be because they abandon the males when the chicks hatch, leaving the males to attend the chicks. Although the duration of primary moult followed the expected trend, being fastest in north Norway and slowest in Britain, the onset of moult was so late in north Norway that they had an unexpectedly late completion date, despite their rapid moult. The late completion of primary moult in north Norway suggests that wintering in the far north may not pose the energetic constraints on Purple Sandpipers that had previously been supposed.

Studies in Iceland (66°N) and Svalbard (78°N), combined with the results of previous work, allowed geographical comparisons of different aspects of the breeding biology of the Purple Sandpiper Calidris maritima, testing predictions that they may be related to latitude and altitude (here taken as surrogates for climatic severity). The breeding density was lowest in the polar deserts of Franz Josef Land (82°N) and highest in coastal Arctic Tern Sterna paradisaea colonies in Iceland where the terns perhaps afforded protection from predators. There was no difference in the percentages of 1-year-old (first-year) birds in the breeding populations in Svalbard and Iceland. Time of breeding was related to both latitude and altitude, being later at higher latitudes and altitudes. In Iceland, larger males and males over 1 year old nested earlier than small males and 1 -year-old males, respectively. Relaying occurred in Iceland but it was not observed in Svalbard. There were no geographical differences in clutch size. Egg size was related to female size in Svalbard. There were also geographical differences in egg size associated with female size (wing length), with the largest eggs in Iceland and smallest in south Norway. Males usually attended broods and the rates at which females deserted their broods were similar in Iceland and Svalbard. Chick growth rates were similar for Iceland and Franz Josef Land. Thus, in breeding biology, Purple Sandpipers varied geographically only in breeding density, time of nesting and egg size, which was associated with female size.

Density-dependent mortality in wintering shorebirds is critical to shorebird population dynamics, but is difficult to study and demonstrate. Most winter mortality of small shorebirds probably results from predation by raptors. Over 13 winters, I studied Eurasian Sparrowhawk Accipiter nisus predation on wintering Dunlins Calidris alpina at a rocky shore in south-east Scotland, a site apparently of marginal suitability for wintering Dunlins. Most Dunlins on the study site were juveniles (first winter birds), and winter juvenile mortality rate due to Sparrowhawk predation was strongly and positively related to Dunlin density (maximum winter count). There was no indication of any marked effect of weather on Dunlin winter mortality rate, except that mortality was slightly greater in winters with lower temperatures. Dunlin mortality was unrelated to the mortality through Sparrowhawk predation of a preferred prey, the Redshank Tringa totanus, or to sighting rates of hawks, suggesting that increased hunting of Redshanks by hawks or a numerical response by hawks were not involved in density-dependent mortality of Dunlins.