Behavior of Buff-Breasted Sandpipers (Tryngites subruficollis) during Migratory Stopover in Agricultural Fields

Background

Understanding the behavior of birds in agricultural habitats can be the first step in evaluating the conservation implications of birds'' use of landscapes shaped by modern agriculture. The existence and magnitude of risk from agricultural practices and the quality of resources agricultural lands provide will be determined largely by how birds use these habitats. Buff-breasted Sandpipers (Tryngites subruficollis) are a species of conservation concern. During spring migration large numbers of Buff-breasted Sandpipers stopover in row crop fields in the Rainwater Basin region of Nebraska. We used behavioral observations as a first step in evaluating how Buff-breasted Sandpipers use crop fields during migratory stopover.

Methodology/Principal Findings

We measured behavior during migratory stopover using scan and focal individual sampling to determine how birds were using crop fields. Foraging was the most frequent behavior observed, but the intensity of foraging changed over the course of the day with a distinct mid-day low point. Relative to other migrating shorebirds, Buff-breasted Sandpipers spent a significant proportion of their time in social interactions including courtship displays.

Conclusions/Significance

Our results show that the primary use of upland agricultural fields by migrating Buff-breasted Sandpipers is foraging while wetlands are used for maintenance and resting. The importance of foraging in row crop fields suggests that both the quality of food resources available in fields and the possible risks from dietary exposure to agricultural chemicals will be important to consider when developing conservation plans for Buff-breasted Sandpipers migrating through the Great Plains.     

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.     

Wintering site fidelity and movement patterns of Western Sandpipers Calidris mauri in the San Francisco Bay estuary

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.     

Density,habitat, and the mating system of the Western Sandpiper (Calidris mauri)

Summary The hypothesis that intense predation, variable food availability, and increased social interactions in high density populations have been important factors promoting the evolution of polygynous and promiscuous mating systems in certain Calidridine sandpipers (Holmes and Pitelka, 1966) is examined in breeding populations of the Western Sandpiper (Calidris mauri) in subarctic Alaska.Western Sandpipers breed in a habitat consisting of a complex mosaic of wet low-lying marshes and relatively well-drained, heath-covered tundra. They defend small territories and nest on the latter, while some feeding also occurs there. Most foods however are obtained off territory in the wet marshes and along the shores of lakes, rivers and sloughs.Densities in the nesting areas ranged from 132–196 pairs/40 ha at the base of a low range of hills to 200–300/40 ha on hummocks surrounded completely by marsh. These densities, the highest reported for a Calidridine sandpiper, are relatively constant from year to year.The mating system of the Western Sandpiper is monogamous, both sexes incubate and care for young. Correlated with a strong single pair-bond and stable populations is a strong tendency to return to the same site. Of sandpipers marked, an average of 57.6% of males and 48.8% of females returned in succeeding years, frequently to the same territory or its immediate vicinity. Of the pairs returning to the study area in subsequent years, 61.5% reunited.The restriction of nesting activities to heath tundra is considered to be a result of the protection it provides for nests. With the separation of nesting and feeding areas and with relatively abundant food sources that are not significantly affected by weather, a large nesting area is not required, allowing high densities of Western Sandpipers to occupy the patches of heath-covered tundra. In this ecological context, the high intensity of social interactions has not resulted in the evolution of a nonmonogamous mating system. Indeed, the increased protection afforded offspring by the presence of both parents is probably the most important selective force promoting the evolution of the monogamous mating pattern in Western Sandpipers.     

Ecological factors regulating brood attendance patterns of the Western Sandpiper Calidris mauri

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.     

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.     

Digestive organ sizes and enzyme activities of refueling western sandpipers (Calidris mauri): contrasting effects of season and age

We examined seasonal and age-related variation in digestive organ sizes and enzyme activities in female western sandpipers (Calidris mauri) refueling at a coastal stopover site in southern British Columbia. Adult sandpipers exhibited seasonal variation in pancreatic and intestinal enzyme activities but not in digestive system or organ sizes. Spring migrants had 22% higher total and 67% higher standardized pancreatic lipase activities but 37% lower total pancreatic amylase activity than fall migrants, which suggests that the spring diet was enriched with lipids but low in glycogen. Spring migrants also had 47% higher total intestinal maltase activity as well as 56% higher standardized maltase and 13% higher standardized aminopeptidase-N activities. Spring migrants had higher total enzymic capacity than fall migrants, due primarily to higher total lipase and maltase activities. During fall migration, the juvenile's digestive system was 10% larger than the adult's, and it was composed differently: juveniles had a 16% larger small intestine but a 27% smaller proventriculus. The juvenile's larger digestive system was associated with lower total enzymic capacity than the adult's due to 20% lower total chitinase and 23% lower total lipase activities. These results suggest that juvenile western sandpipers may process food differently from adults and/or have a lower-quality diet.     

Phenotypic flexibility of body composition in relation to migratory state,age, and sex in the western sandpiper (Calidris mauri)

We investigated the flexibility of body composition in relation to seasonally variable demands for endurance flight capacity and hyperphagia in a migratory shorebird. Migrating western sandpipers were sampled in spring and fall while refueling at a north temperate stopover and were compared with nonmigrating birds captured at a tropical wintering area in Panama. Sandpipers weighed 25% more at stopover, and nearly 40% of migratory mass increase consisted of lean body components. Most organs and flight muscles were 10%-100% larger during migration, and the greatest relative size increases occurred in the digestive system (including liver). Birds preparing to initiate spring migration from Panama deposited only fat, suggesting that changes in lean body components take place after migration has begun, possibly through training effects. Sex did not influence body composition. Juveniles making their first southward migration were similar to adults in structural size and body mass but had substantially enlarged alimentary tracts. Sandpipers appeared to deposit lean mass during stopover in fall but not in spring. The dramatic enlargement of the digestive system in this small species that makes short flights and fuels frequently contrasts with the reduction of digestive components in larger species that fuel only once or twice by making one or two very long flights to their destination.     

Helminth parasites of the western sandpiper, Calidris mauri (Aves), from El Paso and Hudspeth counties, Texas

Fifty western sandpipers, Calidris mauri, from El Paso and Hudspeth counties, Texas, were collected and examined for helminth parasites. Fifty-three helminths (means abundance = 1.06, SD = 2.31) consisting of 4 cestode and 1 nematode species were collected. The helminth community showed low species richness (5), low diversity and evenness (0.05, 0.14), low concentration for dominance (0.19), and all species were contagiously distributed. There were no clearly identifiable core species.     

Sex differences in the stopover ecology of Curlew Sandpipers Calidris ferruginea at a refuelling area during autumn migration

We investigated the stopover patterns of male and female Curlew Sandpipers at a stopover area in northeast Spain. Curlew Sandpipers were trapped and colour-ringed during autumn migration in 1992 and 1993. Stopover length was similar to those reported previously for this and other waders that migrate using a small number of widely separated staging areas, but were greater than stopovers reported for other waders that migrate using a large number of staging areas separated by short distances. The differences in stopover length between the birds using these two strategies could be related to the fuel reserves that have to be accumulated to reach the next staging area. Males stayed longer in the area than females. Seasonal changes in prey availability or sex differences in moulting and migratory patterns do not account for these differences in stopover ecology. Following a time-selected model of optimal migration, sex differences in stopover ecology could be related to a dominance of the larger females over the males or to a higher foraging efficiency or a shorter search and settling time in females. Whether these differences are restricted to the studied area or are widespread in other staging areas used by the species could be important for assessing the possible differences in the migration speed of Curlew Sandpipers. The finding that males leave the breeding grounds 21–35 days before females but arrive at the study area with only a 10-day difference supports the hypothesis that females migrate faster than males at least in the first half of their migration.     

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.     

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.     

宁夏中卫黄河沿岸迁徙灰鹤停歇模式

中途停歇地是候鸟重要的能量补给地及休息站,对其顺利完成迁徙有着重要作用。为了解灰鹤(Grus grus)迁徙途中的停歇模式,于2013年和2015年冬季,采用卫星跟踪技术在贵州草海保护区成功跟踪了4只灰鹤。结果显示,其中3只灰鹤在宁夏中卫市境内的黄河和清水河流域湿地停歇,表明该地区为灰鹤迁徙时的重要停歇地。春季迁徙时,灰鹤在2月初至3月初到达中卫,平均停留(24.5?6.7)d(15~30 d,n=4),2月底至3月底迁离;秋季迁徙时,10月末至11月初到达,平均停留(32.0?4.2)d(29~35 d,n=2),至11月底迁离。停留期间主要的活动范围为中宁县境内的长山头水库、黄河河心岛及同心县境内的清水河周边区域。2017年3月18日,采用夜栖地直接计数法进行实地调查,共发现灰鹤636只,其日间活动范围和卫星跟踪结果类似。此次发现的灰鹤停歇地均未纳入自然保护体系,建议当地林业部门在灰鹤迁徙期间加强管理,可适当投食,以减少灰鹤误食耕地中经农药包衣处理的玉米(Zea mays)种子造成的死亡。     

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.     

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.     

Tradeoffs between food abundance and predation danger in spatial usage of a stopover site by western sandpipers, Calidris mauri

Foragers use a variety of anti-predator behaviours to increase their safety from predators. While foraging, animals should alter usage within or between sites to balance the benefits of feeding with the costs of predation. I tested how the distribution of food abundance and predation danger interacts to explain spatial usage (i.e. distance from shore) by migratory western sandpipers ( Calidris mauri ) at Boundary Bay, British Columbia, Canada, during northward and southward migrations. At Boundary Bay there are opposing spatial gradients in the distribution of food abundance and safety from predators. Predation danger for sandpipers is high near the shoreline where there is approach cover for falcons and decreases with distance from shore. Food abundance for sandpipers declines as distance from the shoreline increases. Food and danger attributes at Boundary Bay also differ temporally, such that food abundance is higher during southward migration, and predation danger is higher during northward migration. The spatial usage by western sandpipers balances the tradeoff between the opposing spatial gradients in food and safety. For both migratory periods spatial usage of the mudflat by sandpipers is highest at distances from the shoreline where food abundance and predation danger are intermediate. During the northward migration sandpiper usage is highest between 150 and 500 m from the shoreline, and during the southward migration sandpiper usage is highest between 100 and 600 m from the shoreline. Despite temporal differences in food and danger attributes, spatial usage of the site by sandpipers does not differ between migratory periods. Understanding how the distribution of food abundance and predation danger interact to affect the within site usage by shorebirds has important implications for assessments of site quality.     

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.