Timing and duration of moult in three populations of Purple Sandpipers Calidris maritima with different moult/migration patterns

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.     

Population structure,biometrics and moult of migrant Purple Sandpipers Calidris maritima in southwest Iceland in spring

Capsule Iceland is a stop‐over site for a population of Purple Sandpipers that winter in Britain. Here, they accumulate fuel loads for onward migration along with birds that have wintered in Iceland.

Aims To establish whether Purple Sandpipers from Britain stop‐over in Iceland during spring migration and, if so, to describe their population structure, changes in mass and moult.

Methods Purple Sandpipers were cannon‐netted on the coast of the Reykjanes Peninsula in southwest Iceland during May 2003 and 2005. Birds were aged, sexed (some by DNA) and standard biometric measurements made. Active body moult was scored.

Results Bill and wing lengths showed that the Purple Sandpipers we caught were similar to one of the populations that winter in Britain rather than Icelandic breeding birds. There were more males than females throughout the migration period (63% males for first‐year‐birds and 67% for adult birds). Accounting for a bias due to a higher percentage of males in a less usual habitat (muddy/sandy bays), the values for rocky sites were 52% males for first‐year birds and 62% for adults. The percentage of first‐year birds was 19% in 2003 and 32% in 2005, though the latter figure was biased by catches in muddy/sandy bays where there was a higher percentage of young birds. The percentage of first‐year birds was 25% on just the rocky shores in 2005. Many birds were in latter stages of body moult, and males were slightly in advance of females. Increasing mass showed that they were preparing for onward migration. The average increase of 0.58 g per day was similar to the rate measured in Orkney at an earlier point on the migration route. However, a high turnover of birds could be the reason for these low values. By late May, and close to the assumed departure date, the Purple Sandpipers of the different age/sex classes had fuel indices of 24–29% (33–42% of the lean mass). This was lower than that for the high Arctic sandpipers (Knots and Sanderlings) leaving southwest Iceland for Greenland and Canada.

Conclusions Our study confirmed that Purple Sandpipers do stop‐over in Iceland, and the possible lower rate of fuel accumulation and smaller amount stored, compared with Knots and Sanderlings, suggests a different migration pattern.     

Seasonal, size- and age-related patterns in body-mass and composition of Purple Sandpipers Calidris maritima in Britain

The masses of 3229 Purple Sandpipers Calidris maritima from Britain were analysed for differences related to age, season and size. First-year birds were lighter by 2 g. There was only a slight increase in mass in mid-winter, in contrast to other waders wintering in Britain, suggesting that Purple Sandpipers are less at risk to cold weather and food shortages. Their winter fat reserves were low but their breast muscles were relatively larger than other small waders wintering in Britain. Their plumage was slightly heavier and their breast and belly feathers were longer. There was a large increase in mass in May by first-year and adult birds prior to migration. The 'long-billed' population showed a greater relative increase in May, compared with the 'short-billed' population, perhaps reflecting different migration distances.     

The winter distribution of Purple Sandpipers in Britain

A survey of numerical data from regional sources suggests that the British wintering population of Purple Sandpipers is between 14,500 and 23,000 birds, mainly concentrated in the north-east.     

Within‐year movements and site fidelity of Purple Sandpipers during the nonbreeding season

ABSTRACT Although within‐year site fidelity to specific wintering sites allows shorebirds to use prior knowledge of resources and microhabitats, such fidelity may also make populations more vulnerable to extirpation in the event of increased predation pressure, habitat loss, or disturbance. In the eastern Atlantic, Purple Sandpipers (Calidris maritima) have been found to be highly faithful to specific sites in wintering areas. However, little is known about the use of wintering areas by these sandpipers along the coast of Maine. We quantified movements of 60 radio‐marked Purple Sandpipers in a bay near the mainland and on an offshore cluster of islands along the mid‐coast of Maine during two winters (2005–2006 and 2006–2007). Birds marked in early‐ and mid‐December remained until spring migration, with no evidence of onward migration. Mean maximum distances moved did not differ significantly between either males (8.6 ± 1.0 [SE] km; N= 30) and females (7.4 ± 0.8 km; N= 30) or juveniles (9.9 ± 1.6 km; N= 9) and adults (7.8 ± 1.1 km; N= 26). We also detected no monthly (January–May) differences in maximum distances moved. Sixty percent of marked individuals moved ≤5 km between the two most distant relocations and no birds moved >25 km during the 2‐ to 4‐month tracking period. We attribute the high site fidelity primarily to the plentiful prey base in the study area. During a 2‐d period with severe cold, feeding areas at locations protected from wave action became encased in ice and birds at these locations moved up to 10 km offshore to sites with less ice. Species with strong site fidelity, like wintering Purple Sandpipers, may be at higher risk in the event of large‐scale changes in their food base, increased predation pressure, habitat loss, or disturbance. However, the short‐distance movements made when intertidal feeding areas became encased in ice suggest that Purple Sandpipers could potentially move greater distances in response to changing conditions in their wintering areas.     

MOULT OF THE PURPLE SANDPIPER CALIDRIS MARITIMA IN ICELAND

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.     

Phenology of migration and use of wintering sites by the increasing population of dark-bellied brent geese Branta bernicla bernicla

During the recovery of the population of dark-bellied brent geese Branta bernicla bernicla , traditional wintering sites in Britain have experienced increases in numbers and new sites have been colonized. Goose counts were used to describe the migration phenology and winter use of Scolt Head, north Norfolk (a traditional site) and the Burry Inlet, South Wales (a relatively new site) over the period of re-colonization and colonization, respectively. During the 1950s, only a few hundred birds wintered at Scolt Head, and most stayed for only a short period. By the late 1980s/early 1990s, numbers had reached several thousands, their period of stay was longer and there was little variation between years in median arrival (28 October) and departure (11 March) dates. A similar process was observed during the entire colonization phase at the Burry Inlet; initially birds arrived late and departed early, but arrived progressively earlier and departed later as wintering numbers increased. The autumn arrival rates at both sites were slower than the spring departure rates. During years of good breeding productivity, those adults with young arrived slightly later in Norfolk than failed breeders and non-breeders, and a higher proportion of young birds remained in spring after the main departure. However, the overall pattern of arrival and departure did not vary according to whether there had been a good or poor breeding season. Some adults of breeding age also remained until May. The effects of food depletion and use of novel feeding habitats in relation to migration phenology and winter use of sites are discussed.     

Regional, seasonal and annual variations in the structure of Purple Sandpiper Calidris maritime populations in Britain

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.     

Rapid population decline in red knots: fitness consequences of decreased refuelling rates and late arrival in Delaware Bay

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.     

Territory strategy of the migratory Brown Shrike Lanius cristatus

Brown Shrikes Lanius cristatus are both transients and winter residents in Taiwan. Large numbers of birds crowd into the available habitat during the transient period, and the cost of maintaining territories changes greatly, depending on the number of shrikes present. This study examined how wintering Brown Shrikes adapt to the changing level of competition in territory establishment. About 24% of autumn transients arrived before any wintering birds, while 73% of wintering birds arrived during autumn transient time. The frequency of aggression recorded was significantly related to the number of birds seen along the transect line that day. Among ringed birds that returned during transient time, 80% returned directly to their previous territories, but some waited for the departure of transient birds before establishing territories. Territory compression occurred at high population densities. Departure from and arrival at the study area were frequent during the winter, but no birds moved territories within the study area. On average, 25% of ringed wintering shrikes returned to the study site in the subsequent years, while no transients returned. This low site tenacity may be. in part, a result of a high persecution rate along the migration route.     

Does wintering north or south of the Sahara correlate with timing and breeding performance in black‐tailed godwits?

Migrating long distances requires time and energy, and may interact with an individual's performance during breeding. These seasonal interactions in migratory animals are best described in populations with disjunct nonbreeding distributions. The black‐tailed godwit (Limosa limosa limosa), which breeds in agricultural grasslands in Western Europe, has such a disjunct nonbreeding distribution: The majority spend the nonbreeding season in West Africa, while a growing number winters north of the Sahara on the Iberian Peninsula. To test whether crossing the Sahara has an effect on breeding season phenology and reproductive parameters, we examined differences in the timing of arrival, breeding habitat quality, lay date, egg volume, and daily nest survival among godwits (154 females and 157 males), individually marked in a breeding area in the Netherlands for which wintering destination was known on the basis of resightings. We also examined whether individual repeatability in arrival date differed between birds wintering north or south of the Sahara. Contrary to expectation, godwits wintering south of the Sahara arrived two days earlier and initiated their clutch six days earlier than godwits wintering north of the Sahara. Arrival date was equally repeatable for both groups, and egg volume larger in birds wintering north of the Sahara. Despite these differences, we found no association between wintering location and the quality of breeding habitat or nest survival. This suggests that the crossing of an important ecological barrier and doubling of the migration distance, twice a year, do not have clear negative reproductive consequences for some long‐distance migrants.     

Primary moult,biometrics, mass and age composition of Grey Plovers Pluvialis squatarola in southeastern India

Little is known about the biology of waders wintering in southern Asia; this paper deals with the Grey Plover Pluvialis squatarola, a species extensively studied only in western Europe. Adult Grey Plovers wintering in southeastern India underwent primary moult in autumn; the duration was estimated to be 127 days, with mean starting date 1 September and mean completion date 5 January. Some first-year Grey Plovers initiated primary moult in late winter and spring, and completed this moult the following spring. The average mass of adults on arrival in September was 200 g, fluctuated close to 220 g from October to February, and increased to 280 g near the end of May. The mass variation did not show the January peak observed in western Europe. Breeding productivity, measured as the percentage of first-year birds in winter catches, varied between 5% and 70% over six years, and showed a positive correlation with that of Dark-bellied Brent Geese Branta b. bernicla in western Europe and Curlew Sandpipers Calidris ferruginea in South Africa.     

Travel schedules to the high arctic: barnacle geese trade-off the timing of migration with accumulation of fat deposits

On their way from the wintering area to the breeding grounds in Spitsbergen, barnacle geese Branta leucopsis stage on islands off the coast of Norway. The aim of this study was to describe when the geese migrate in relation to the body stores deposited and explore questions related to the concept of optimal migration schedules and on the possible mechanisms involved. We estimated fat stores by repeated assessments of the abdominal profile index of individually marked females throughout staging. Reproductive success was derived from observations of the same individuals later in the annual cycle. Females arriving late, or with low fat stores at arrival, achieved higher fat deposition rates, probably by spending more time foraging. But they were unable to match final fat scores of birds that arrived earlier or with larger fat stores. Reproductive success was correlated with the timing of migration and individuals departing at intermediate dates achieved highest success. The exact date of peak reproductive success depended on the size of fat stores accumulated, such that low-quality birds (depositing less fat) benefited most from an early departure to the breeding grounds. Observations in the breeding colonies showed that these birds did not initiate a nest earlier but they spent a longer time in Spitsbergen before settling. The length of stay in Norway was close to the prediction derived from an optimisation model relating spring events to eventual breeding success. Poorest performing birds stayed longer than expected, perhaps depositing more fat to avoid the risk of starvation. Two possible mechanisms of the timing of migration were contrasted and it seemed that the geese departed for migration as soon as they were unable to accumulate any more fat stores.     

Temperature‐related increases in grass growth and greater competition for food drive earlier migrational departure of wintering Whooper Swans

Understanding the departure decisions of migratory birds is critical for determining how changing climatic conditions will influence subsequent arrival times on the breeding grounds. A long‐term dataset (1972–2008) of Whooper Swan Cygnus cygnus departure dates from a wintering site in Ireland was used to assess the factors determining the timing of migration. Early and late migrating swans showed different departure patterns. Earlier wintering ground departure was more pronounced for the first 50% of the population than the last 10% of departing individuals. Earlier departure was associated with an increase in February temperatures at the wintering site for all departure phases except the date when the last individual departed. The date by which the first 50% of Swans had departed was earlier with increasing numbers of wintering Swans, suggesting that competition on the wintering grounds may further influence the timing of departure. The results also suggested that departure is mediated by the influence of spring temperature on food resources, with increased February grass growth in warmer years enabling earlier departure of migrating Swans. To determine why arrival dates in the breeding ground have altered, environmental conditions in the wintering grounds must be taken into account.     

High within‐winter and annual survival rates in a declining Afro‐Palaearctic migratory bird suggest that wintering conditions do not limit populations

For migratory birds, it is necessary to estimate annual and overwinter survival rates, identify factors that influence survival, and assess whether survival varies with age and sex if we are to understand population dynamics and thus inform conservation. This study is one of the first to document overwinter and annual survival from the wintering grounds of a declining Afro‐Palaearctic migrant bird, the Whinchat Saxicola rubetra. We monitored a population of marked individuals for which dispersal was low and detectability was high, allowing accurate estimates of survival. Annual survival was at least 52% and did not differ significantly across demographic groups or with habitat characteristics or residency time in the previous winter. Overwinter survival was very high and monthly survival at least 98% at some sites. Although winter residency varied spatially and with age, lower residency did not correlate with reduced annual survival, suggesting occupancy of multiple wintering sites rather than higher winter mortality of individuals with shorter residency. Our results suggest that mortality occurs primarily outside the wintering period, probably during migration, and that wintering conditions have minimal influence on survival. The similarity between survival rates for all age and sex classes when measured on the wintering grounds implies that any difference in survival with age or sex occurs only during the first migration or during the post‐fledging stage, and that selection of wintering habitat, or territory quality, makes little difference to survival in Whinchats. Our findings suggest that the wintering grounds do not limit populations as much as the migratory and breeding stages, with implications for the conservation of declining Afro‐Palaearctic migrants more widely.     

Fast fuelling but light flight in Broad-billed Sandpipers Limicola falcinellus: stopover ecology at a final take-off site in spring (Sivash, Ukraine)

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.     

Population-specific adjustment of the annual cycle in a super-swift trans-Saharan migrant

For migratory birds optimal timing of the onset of reproduction is vital, especially when suitable conditions for reproduction occur only for a short while during the year. With increasing latitude the suitable period becomes shorter and we expect the organization of annual cycle to be more synchronized to the local conditions across individuals of same population. This should result in low variation of arrival and departure date in breeding sites at higher latitudes. We quantify the temporal and geographical variation in pre- and post-breeding migration between individuals from four different populations of alpine swifts Tachymarptis melba along a latitudinal gradient. We tracked 215 individuals in three years with geolocators. The two western and two eastern populations showed separate migratory flyways and places of residence in Africa. Length of stay at the breeding sites was negatively correlated with latitude and differed by more than a month between populations. Duration of migration was similarly short in all populations (median 6.2 days in autumn and 8.7 days in spring). However, variation in timing of migration was unrelated to latitude and individuals everywhere arrived in the same asynchrony at the breeding site.     

Inferring geographic origin, population structure and migration routes of a wintering population of Mediterranean gulls from resightings data

Winter congregations of migratory birds are made by individuals of different origins and generally assumed to be variable across space and time, but the demographic characteristics of these temporal populations are poorly known. We used 2,216 observations of 472 colour-ringed individuals to estimate the annual local survival of Mediterranean gulls Larus melanocephalus wintering in NE Spain. In addition, by gathering the ringing information on the 19,856 individuals marked as fledglings in 18 countries between 1990 and 2009, we were able to infer the composition of population in relation to the country of origin. We coupled these estimates with geographic information to contrast hypotheses on the migratory pattern most likely used by the gulls in their first migration from their natal colonies to the wintering area. The probability of reaching the study area was negatively associated with the distance from the natal colony. Data were consistent with a migratory strategy that combines fluvial and coastal routes in an optimal way, seeking minimal distance along favourable terrain. We found that, after the first year, annual local survival at the wintering site (0.81 on average) was comparable with the one estimated at the breeding colonies, indicating a high individual fidelity to the areas used in winter. Our work shows that winter groupings may behave as real populations, shaped by breeding output and survival, and that the geographic origin of wintering birds can be explained by a simple model. The study of winter congregations can help understand a species’ population structure and movement strategies.     

Bill length: a reliable method for sexing Purple Sandpipers

ABSTRACT.   Purple Sandpipers ( Calidris maritima ) are sexually dimorphic, with females larger than males. We tested the reliability of using bill length to sex individuals and estimate the sex ratio at a stopover site in Iceland in May 2003 and 2005. Feather samples from 65 of 324 captured birds were used for molecular sexing, and generalized linear models (GLMs) were used to discriminate the sexes from body measurements of the molecularly sexed birds. About 3% of the 324 individuals were misclassified by the Harding-Cassie method, and the proportion of males was 0.657 compared to 0.656 according to the best GLM. Our results showed that the reliability of determining the sex and sex ratio of Purple Sandpipers using a Harding-Cassie plot of bill length measurements was high, but that reliability was improved by including other variables in a GLM. The estimate of an uneven sex ratio in the population we studied was not due to a systematic error, and supports the conclusion of earlier studies that Purple Sandpipers exhibit an uneven sex ratio in favor of males.     

Aggressive behaviours and correlates of dominance in Purple Sandpipers Calidris maritima at a communal winter roost

Aggression and the competitive ability of individual Purple Sandpipers were studied at a communal winter roost in northeast England. Aggression was most intense in high winds and more frequent in moderate breezes than when wind speeds were high or low. Higher wind speeds resulted in a drop in aggression rate, as birds had to face into the wind to maintain their balance. The ability of an individual to secure a sheltered roost position was related to its size and sex, larger individuals being dominant over those smaller than themselves. Purple Sandpipers exhibit reversed sexual size dimorphism and females were thus dominant over males. There was also weak evidence that adults were dominant over first-winter birds. The importance of size and sex in determining dominance at the roost is discussed in relation to the evolution of reversed sexual size dimorphism in Purple Sandpipers and similar shorebirds.