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 of Purple Sandpipers (Calidris maritima) as revealed by mitochondrial DNA and microsatellites

The Purple Sandpiper (Calidris maritima) is a medium‐sized shorebird that breeds in the Arctic and winters along northern Atlantic coastlines. Migration routes and affiliations between breeding grounds and wintering grounds are incompletely understood. Some populations appear to be declining, and future management policies for this species will benefit from understanding their migration patterns. This study used two mitochondrial DNA markers and 10 microsatellite loci to analyze current population structure and historical demographic trends. Samples were obtained from breeding locations in Nunavut (Canada), Iceland, and Svalbard (Norway) and from wintering locations along the coast of Maine (USA), Nova Scotia, New Brunswick, and Newfoundland (Canada), and Scotland (UK). Mitochondrial haplotypes displayed low genetic diversity, and a shallow phylogeny indicating recent divergence. With the exception of the two Canadian breeding populations from Nunavut, there was significant genetic differentiation among samples from all breeding locations; however, none of the breeding populations was a monophyletic group. We also found differentiation between both Iceland and Svalbard breeding populations and North American wintering populations. This pattern of divergence is consistent with a previously proposed migratory pathway between Canadian breeding locations and wintering grounds in the United Kingdom, but argues against migration between breeding grounds in Iceland and Svalbard and wintering grounds in North America. Breeding birds from Svalbard also showed a genetic signature intermediate between Canadian breeders and Icelandic breeders. Our results extend current knowledge of Purple Sandpiper population genetic structure and present new information regarding migration routes to wintering grounds in North America.     

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.     

Numbers,migration phenology and survival of Purple Sandpipers Calidris maritima at Gourdon,eastern Scotland

Purple Sandpipers wintering on the Kincardine coast had a protracted autumn arrival (one-quarter and three-quarters of the birds arrived on 30 July and 21 October respectively, 83 days) but a faster spring departure (one-quarter and three-quarters departed on 9 April and 27 May respectively, 48 days). The long arrival period was partly due to differences in the migration phenology of the two main wintering populations: short-billed birds from Norway arrived before the long-billed birds, probably from Canada. There was a smaller difference in departure times of the two populations: short-billed birds left before the long-billed birds. Minimum annual survival was estimated from resightings of 92 marked birds. There was no evidence that survival differed between adults and first-years or between birds of different bill-size classes, which were of different sex and geographical origin. Minimum annual survival was estimated to be 79.5%(se = 2.8%). The similarity between the mortality rate (20.5%) and the percentage of first-year birds in populations of Purple Sandpipers probably reflects balanced population dynamics.     

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.     

Sex roles in the monogamous Purple Sandpiper Calidris maritima in Svalbard

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.     

Geographical variation in the breeding biology of the Purple Sandpiper Calidris maritima

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.     

Differences in biometrics and moult of non-breeding Red Knots Calidris canutus in southern Africa and Scotland reflect contrasting climatic conditions

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.     

A review of the subspecies status of the Icelandic Purple Sandpiper Calidris maritima littoralis

The Icelandic Purple Sandpiper Calidris maritima littoralis (C.L. Brehm, 1831) represents one member of a poorly understood subspecies complex. Currently, differences in size define two other subspecies: Calidris maritima belcheri Engelmoer & Roselaar, 1998, which breeds in north‐eastern Canada along the Hudson Bay and James Bay, and Calidris maritima maritima (Brunnich, 1764), which breeds along the Arctic coasts elsewhere in northern Canada, Greenland, Svalbard, Scotland, and Fennoscandia, to northern central Siberia. There are large size differences amongst populations of C. m. maritima, however. As an Arctic/Alpine breeding bird, C. m. littoralis could provide an interesting perspective on the evolutionary changes following a northwards expansion of a species after glacial retreat. Considering the extent of the ice sheet in the northern hemisphere during the last glaciation, and the short period of time since it ended, the correct attribution of subspecies status for C. m. maritima may reflect either rapid diversification from a single population or ancestral splits of distinct evolutionary lineages that survived in isolation at southern latitudes. We applied morphometric subspecies criteria, diagnosability by Amadon's rule, and genetic analysis of five nuclear introns, and the mitochondrial DNA markers cytochrome oxidase c subunit I (COI) and NADH dehydrogenase subunit 2 (ND2), to geographically separate breeding populations in order to examine the subspecies status of the Icelandic population. The results do not provide support for the subspecies status of the Icelandic population because the nominate and Icelandic subspecies fail to uphold Amadon's rule, and genetic analyses indicate that the study populations derive from a single shared refugium. © 2015 The Linnean Society of London     

Prebreeding moult, plumage and evidence for a presupplemental moult in the Great Knot Calidris tenuirostris

We studied the prebreeding moult and resulting plumage in a long-distance migrant sandpiper (Scolopacidae), the Great Knot Calidris tenuirostris , on the non-breeding grounds (northwest Australia), on arrival at the staging grounds after the first migratory flight (eastern China) and on or near the Russian breeding grounds (Russian data from museum specimens). We show that breeding plumage scores and breast blackness were affected not only by the increase in moulted feathers but also in the wearing down of overlaying pale tips of fresh feathers. Birds migrating from Australia and arriving in China had completed or suspended moult, but more moult must occur in Asia as Russian specimens had moulted more of their mantle and scapular feathers. Russian birds had significantly more red feathering on their upperparts than had birds in Australia or those arriving in China. The increase in reddish feathers cannot by accounted for simply by continuation of the prealternate moult. Instead, a third, presupplemental moult must occur, in which red-marked feathers replace some scapular and especially mantle feathers that were acquired in a prealternate moult only 1–3 months earlier. Great Knot sexes show little size and plumage dimorphism, whereas two other sandpipers that have supplemental plumages (Ruff Philomachus pugnax and Bar-tailed Godwit Limosa lapponica ) are thought to be highly sexually selected. Bidirectional sexual selection may therefore be involved in the evolution of a supplemental plumage in Great Knots.     

The spring migration pattern of arctic birds in southwest Iceland, as recorded by radar

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.     

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.     

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.     

Unravelling the migration and moult strategies of a long-distance migrant using stable isotopes: Red Knot Calidris canutus movements in the Americas

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.     

Migration and moult of Dunlin Calidris alpina wintering in the central Mediterranean

Dunlin migration in northeast Italy is described. An attempt to identify the main routes and staging areas used by birds wintering in the central Mediterranean is presented. The results of monthly counts from 1990–1995 revealed that the bulk of the population occupied the wintering area in October and left for the breeding grounds in April and May. The analysis of 342 Italian recoveries of foreign ringed birds showed that 65% were ringed during post-breeding migration through the Baltic Sea, whereas just a few birds had been ringed in western Europe. First-year birds arrived in autumn with a single migratory wave, peaking in October. Two categories of adults were identified during post-breeding migration: birds which directly reached Italian wintering sites and birds which arrived after they had suspended their migration for moulting: the Azov/Black Sea wetlands are suggested as possible moulting areas. Out of 2444 adults and 1627 first-years ringed between 1989 and 1996 at our study area, we obtained a total of 42 recoveries abroad and evidence of direct links between Azov/Black Sea and N Adriatic wetlands, both during autumn and spring migrations. Primary moult was observed only in adults arriving early, the second migratory wave being composed of moulted birds. Locally moulting adults adopted a moult strategy characterized by high raggedness scores, typical of resident moulters. Body mass was not affected by primary moult stage or intensity, winter mass values being reached two weeks after the average date of primary moult completion.     

Local survival in Semipalmated Sandpipers Calidris pusilla breeding at La Pérouse Bay, Canada

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 Pérouse 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.     

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.     

Foraging behaviour of non-breeding Western Sandpipers Calidris mauri as a function of sex, habitat and flocking

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 Bahía Santa María, 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 Bahía Santa María.     

Population structure and dynamics of wood mouse Apodemus sylvaticus in Iceland

Five-year-studies of two wood mouse populations (Apodemus sylvaticus L.) were carried out outside the village Vík. S. Iceland, and in a birch forest close to the lake Mývatn, N. Iceland. The population at Vík was studied on a 1.10 ha grid on a slope covered with Angelica archangelica L., the seeds of which constituted a major food resource for the mice. Altogether 948 individuals were caught, marked and followed during 1973–1978. No other free-living small mammals occur in Iceland, and predator pressure is low and considered non-existent on the Angelica slope. The population density was extremely high; highest autumn density recorded was 150 ind ha^?1. Like other Apodemus populations, the one at Vík showed a typical seasonal pattern in numbers: a winter decrease, relatively stable numbers in late summer, and an autumn peak. Breeding started in late April or early May and ended in late September; no winter breeding was observed. Three or four litters were produced, and the last litter(s) of the season greatly contributed to the autumn increase. The output of young per reproductive female throughout the year was negatively correlated with spring density, and autumn peak density with the number of females in August. Also overwinter success and weight changes appeared to be associated with population density (and weather). Survival rates did not vary much between years and seasons and were consistently rather high. Autumn survival of juveniles was positively correlated with mean minimum temperatures. Juvenile survival was, at least at times, negatively affected by the presence of adult males. Trapping outside the main grid did not indicate any marked dispersal, though some immigration to the grid must have occurred, especially in spring and early summer. Among the factors governing the population density and dynamics of wood mouse at Vík we emphasize the absence of interspecific competition and predation, the rich and dependable food resources (mainly Angelica seeds), the mild (though moist and windy) climate, the shelter of dense Angelica stands, the abundance of crevices (e.g. nest-sites), and the social interactions. The birch forest in northern Iceland was low productive and the wood mouse population differed from the Vík population by exhibiting low density, short breeding season, and high vagility; characteristics we consider to be representative for most Apodemus populations in Iceland.