Weather conditions and conspecific density influence survival of overwintering Dunlin Calidris alpina in North Wales

Capsule Inter-annual survival rates of juvenile Dunlin Calidris alpina increased with wet weather conditions but decreased with the density of conspecifics in winter, ranging from 0.34 to 0.62 in the best model. Adult inter-annual survival remained high and constant at 0.72.

Aims To estimate inter-annual survival rates of both adult and juvenile Dunlin in North Wales. To quantify the effects on survival rates of weather conditions and conspecific density.

Methods Cormack–Jolly–Seber models in Program MARK were used to analyse 21 years of ringing data (1990–2011) from Traeth Lafan, North Wales. Models were constructed using a two age-class structure. The influence of a range of weather conditions and WeBS census data on survival were examined.

Results Adult survival was best modelled at a constant rate of 0.72 (se 0.008) across the 21 years. Juvenile survival rates were found to vary most strongly with total rainfall (mm) during the season and the number of conspecifics present. Survival rates varied from 0.34 (se 0.06) to 0.62 (se 0.09), with higher survival in years with high rainfall and low numbers of birds.

Conclusion Survival rates of juvenile Dunlin are affected by both weather and density of conspecifics. These results have implications for the future of Dunlin in the UK when considering both climate change and habitat loss.     

Conservation genetics and phylogeography of southern dunlins Calidris alpina schinzii

Breeding populations of southern dunlin Calidris alpina schinzii in South Fennoscandia and the Baltic are severely fragmented and declining dramatically. Information on the genetic structure and diversity is therefore of importance for the conservation and management of these populations. Here we present the results of comparative genetic analyses of these populations with other populations of the schinzii , alpina and arctica subspecies in northern Europe. We sequenced the mitochondrial DNA control region and the Z-chromosome intron VLDLR-9, and analyzed microsatellites and AFLPs, for analyses of within-population genetic diversity. We also extended previous analyses of the phylogeographic structure of dunlins in northern Europe with a larger sample of individuals and populations. Our results revealed no evidence of reduced genetic diversity or increased levels of inbreeding in the small and fragmented populations around the Baltic Sea as compared to the more vital and larger populations elsewhere. Nevertheless, their small population sizes and presumably high degree of isolation may lead to local extinctions, indicating that demographic and ecological factors may pose a greater threat to the survival of these populations than purely genetic factors. Phylogeographically, the schinzii populations in Scandinavia and the Baltic do not form a separate genetic clade, but are part of larger cline of genetic variation encompassing several recognized subspecies of dunlins in the western Palearctic region. Only the Icelandic population showed some distinctiveness in genetic structure and might therefore be considered a separate management unit. Our study highlights the general problem of lack of genetic support for subspecies in avian taxonomy and conservation genetics.     

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.     

Body mass change and diet switch tracked by stable isotopes indicate time spent at a stopover site during autumn migration in dunlins Calidris alpina alpina

Birds may change their diet and foraging habitat during or after migration. Dunlins Calidris alpina alpina breed in the tundra of northern Europe and Russia where they feed exclusively on terrestrial prey. However, up to 80% of the flyway population uses the Wadden Sea as their first important staging site on the way to wintering grounds, feeding exclusively on marine prey. Adult birds migrate earlier than immatures and tend to fly non‐stop, whereas immatures may stage for at least a few days en route, mainly in the Baltic region. There they mostly feed on brackish water prey showing similar isotopic values compared to terrestrial prey. When they arrive in the Wadden Sea, dunlin body reserves are depleted and lower than those of individuals that have already staged for several days. We hypothesized that lighter individuals should retain a strong terrestrial isotopic blood signature, while heavier ones should show a stronger marine signature. We found a significant positive correlation between scaled mass index and carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotopes, reflecting the switch from terrestrial to marine prey during migration. A mixing model revealed differences in isotopic values between heavy and light adults and immatures, respectively, in relation to the isotopic prey signatures. Adults showed stronger marine signals compared with immatures, emphasizing the different modes of migration (i.e. a later departure in immatures) as well as the known spatial segregation of age classes in the Wadden Sea, i.e. adults use tidal flats distant from the shore while immatures use coastal areas influenced by terrestrial carbon sources. The results of this study demonstrate the value of scaled mass index in migratory birds as an indicator of time elapsed after diet switching following migration. Furthermore, this study extents the existing knowledge on the timing of dunlin migration by using an isotopic approach.     

Membrane lipid composition and cellular function

Membrane fatty acid composition, phospholipid composition, and cholesterol content can be modified in many different kinds of intact mammalian cells. The modifications are extensive enough to alter membrane fluidity and affect a number of cellular functions, including carrier-mediated transport, the properties of certain membrane-bound enzymes, binding to the insulin and opiate receptors, phagocytosis, endocytosis, depolarization-dependent exocytosis, immunologic and chemotherapeutic cytotoxicity, prostaglandin production, and cell growth. The effects of lipid modification on cellular function are very complex. They often vary from one type of cell to another, and they do not exert a uniform effect on all processes in a single cell line. Therefore, it is not yet possible to make any generalizations or to predict how a given system will respond to a particular type of lipid modification. Many of the functional responses probably are caused directly by the membrane lipid structural changes, which affect either bulk lipid fluidity or specific lipid domains. The conformation or quaternary structures of certain transporters, receptors, and enzymes probably are sensitive to changes in the structure of their lipid microenvironment, leading to changes in activity. Prostaglandin production is modulated by the availability of substrate fatty acids stored in the membrane phospholipids, but the underlying chemical mechanism still involves a change in membrane lipid structure. While this is the most likely mechanism, the possibility that the membrane lipid compositional change is an independent event that occurs concurrently but is not causally related to the functional perturbations also must be considered.     

Genetic and morphological variation in Dunlin Calidris alpina breeding in the Palearctic tundra

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.     

人为干扰对黑腹滨鹬觅食行为的影响

2003年11月至2004年3月,采用目标动物取样法和人为干扰实验法,于浙江省乐清湾滩涂进行人类活动对黑腹滨鹬(Calidrisalpina)觅食行为影响的研究。取样观察结果表明,在每天沿海堤可观察黑腹滨鹬持续觅食的1h里,96.6%的黑腹滨鹬在35m或更小的距离上通过奔跑或惊飞对人类的活动作出反应;黑腹滨鹬花费90%(人类较多的盐盆滩涂为85%,较少的乐成滩涂为94%)的时间觅食,10%的时间用于避免人类的干扰;人数、人类活动类型对觅食时间具有显著影响,人数、人离黑腹滨鹬的距离、人类活动类型对黑腹滨鹬移动的距离、移动的次数以及黑腹滨鹬对人类靠近的反应均具有显著效应。人为干扰最小接近距离实验结果表明,黑腹滨鹬对人类反应的距离在3～30m;干扰人数为1人时的最小接近距离显著大于2人;人类行走和奔跑的两种活动类型对黑腹滨鹬的最小接近距离没有显著影响。本研究验证了滨鸟觅食区内的人为活动严重影响黑腹滨鹬觅食行为的假设。     

Choosing the best foraging microhabitats: individual skills constrain the choices of dunlins Calidris alpina

Estuarine mudflats, among the most important foraging grounds for waders during the non-breeding season, consist of complex mosaics of shallow pools and dry areas during low tide. In this study, we carried out close-range focal observations to determine foraging parameters of dunlins Calidris alpina, foraging in the mudflat microhabitats of the Tagus estuary, Portugal. Birds foraging in wet patches mostly targeted the siphons of the bivalve Scrobicularia plana , while in dry patches they mostly fed on mudsnails Hydrobia ulvae . Surface visibility of prey, rather than their abundance in the sediment, explained the microhabitat-related differences in prey selection. Birds using dry patches obtained 40% less energy intake than those using wet patches, still many extensively used this poor microhabitat. Because siphons retract quickly when the sediment is disturbed, birds often failed to catch them. We found that birds that were less efficient in capturing siphons in wet patches tended to spend more time foraging on mudsnails in dry patches. This suggests that lack of skills in siphon cropping represents a major foraging constraint for dunlins wintering in the Tagus estuary. It may even cause them to forage during high tide in order to achieve their daily energetic requirements.     

Membrane lipid composition defines membrane protein spectrum

In addition to glycerophospholipids, bacterial membranes often include amino acid-containing acyloxyacyl lipids. The functional implications of these aminolipids are largely unknown. However, a recent study by Stirrup et al. expands our understanding and shows that they are major determinants for membrane properties and the relative abundance of distinct membrane proteins in bacterial membranes.     

Geographical variation in the timing of breeding and moult in dunlin Calidris alpina on the Palearctic tundra

Studies of how organisms are adapted to regional climatic conditions are valuable when predicting the effects of global climatic changes on biota. Here we report on the geographical variation in timing of breeding and moult of an Arctic breeding wader, the dunlin (Calidris alpina). The Palearctic study sites range latitudinally between 68 and 76°N and longitudinally between 46 and 179°E, and encompass a variety of local climates. The sites were visited in sequence from west to east within 1 year, and therefore the data are not affected by confounding interannual variations. The estimated breeding start ranged from 5 to 25 June across populations. Birds at more southern sites were found to breed earlier than those at more northern breeding sites. Within populations, the breeding start for first clutches spanned a period of 8 days and, when including replacement clutches, 3-4 weeks. No dunlin west of the Taimyr Peninsula were found moulting while incubating at the nest, whereas all dunlin on Taimyr Peninsula and eastwards were in active wing moult while incubating or rearing chicks. The onset of moult in these populations ranged from 23 to 27 June. The consequences of geographical variation of breeding conditions for variation in the annual cycle of this species are discussed.     

Movements of wintering Dunlin Calidris alpina and changing habitat availability in an agricultural wetland landscape

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 pub-lication 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.     

The bioavailability of various selenium compounds to a marine wading bird

The uptake of dietary selenium (about 3.5 mg/kg AF dry wt) as selenomethionine, selenocystine, selenite, selenate, and fish selenium in the plasma and red blood cells (RBC) of the oystercatcher has been investigated. The birds received the various selenium compounds subsequently, for at least 9 wk. After dietary supplementation of selenocystine, selenite, and selenate, plasma selenium was about 350 μg/L and RBC selenium 2.1 mg/kg dry wt. After supplementation of selenomethionine, the plasma concentration increased to 630 μg/L, and the RBC concentration to 4.1 mg/kg dry wt. When the fodder contained 3.1 mg/kg fish Se, an average plasma and RBC concentration of 415 μg/L and 14.4 mg/kg dry wt, respectively, was measured. The maximal increase of the selenium concentration in the plasma was attained at first sampling, 14 d after a change in dietary selenium (selenomethione or fish Se); the uptake seemed to be a concentration-regulated process. RBC concentrations (γ in mg/kg dry wt) increased with time (X in d) according toY=a?be^?cX . Fifty percent of the total increase was attained within 17d, suggesting that diffusion into the RBC played a role. The selenium concentration in the plasma was positively correlated with the (fish) Se concentration in the fodder; the RBC concentration (60 d after the change in diet) was positively correlated with the plasma concentration. When the diet contained fish Se, the blood selenium concentrations of the captive birds were similar to the concentrations measured in field birds. Fish Se is a yet undetermined selenium compound. The present experiment showed that fish Se differed from selenomethionine, selenocystine, selenite, or selenate in uptake from the food and uptake in the RBC.     

Membrane lipid composition and susceptibility to bile salt damage

Erythrocyte membranes with low sphingomyelin: choline-containing phospholipid ratios haemolyse at low concentrations of the bile salt, glycocholate. Erythrocytes with higher sphingomyelin: choline-containing phospholipid ratios require progessively greater concentrations of the bile salt for lysis.Sublytic concentrations of glycocholate remove phospholipid and acetylcholinesterase from the membranes. Membranes with low sphingomyelin: choline-containing phospholipid ratios lose both particulate (microvesicles of distinct composition) and ‘solubilized’ material, the particulate form predominating. The proportion of particulate material falls with increase of the membrane sphingomyelin: choline-containing phospholipid ratio and those membranes of highest sphingomyelin: choline-containing phospholipid ratio lose material predominantly in ‘solubilized’ form.Sheep erythrocytes treated to increase their content of phosphatidylcholine (and thereby reduce their membrane sphingomyelin: choline-containing phospholipid ratio) become more susceptible to lysis by glycocholate.These observations indicate a correlation between membrane lipid composition and the perturbation of membranes with bile salt; they also point to possible features of membranes capable of surviving exposure to the high bile salt concentrations of the biliary tract.     

Effects of grassland management on overwintering bird communities

Birds that depend on grassland and successional-scrub vegetation communities are experiencing a greater decline than any other avian assemblage in North America. Habitat loss and degradation on breeding and wintering grounds are among the leading causes of these declines. We used public and private lands in northern Virginia, USA, to explore benefits of grassland management and associated field structure on supporting overwintering bird species from 2013 to 2016. Specifically, we used non-metric multidimensional scaling and multispecies occupancy models to compare species richness and habitat associations of grassland-obligate and successional-scrub species during winter in fields comprised of native warm-season grasses (WSG) or non-native cool-season grasses (CSG) that were managed at different times of the year. Results demonstrated positive correlations of grassland-obligate species with decreased vegetation structure and a higher percentage of grass cover, whereas successional-scrub species positively correlated with increased vegetation structure and height and increased percentages of woody stems, forb cover, and bare ground. Fields of WSG supported higher estimated total and target species richness compared to fields of CSG. Estimated species richness was also influenced by management timing, with fields managed during the previous winter or left unmanaged exhibiting higher estimated richness than fields managed in summer or fall. Warm-season grass fields managed in the previous winter or left unmanaged had higher estimated species richness than any other treatment group. This study identifies important winter habitat associations (e.g., vegetation height and field openness) with species abundance and richness and can be used to make inferences about optimal management practices for overwintering avian species in eastern grasslands of North America. © 2019 The Authors. Journal of Wildlife Management Published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Breeding origin and migration pattern of dunlin (Calidris alpina) revealed by mitochondrial DNA analysis

The large-scale migration of birds has been studied extensively by recoveries of ringed birds. However, there is very little ringing data from the arctic breeding grounds of waders. Here, the migration pattern of the dunlin, Calidris alpina, is studied with population genetic markers, using haplotype frequencies to estimate the breeding origin of migrating and wintering populations. Polymerase chain reaction (PCR) and restriction analysis of DNA from the mitochondrial control region was used to study the breeding origins of morphologically similar winter populations in the western Palaearctic, and to describe the population structure of the dunlin during winter. Also migrating dunlin from various stopover sites in Europe, Africa and Asia, were analysed with respect to their mitochondrial DNA (mtDNA) haplotypes. The genetic markers clearly show that the dunlin has a parallel migration system, with populations breeding in the western Palaearctic wintering mainly in the western part of the wintering range, and dunlin populations breeding further east wintering further east. The results also show that the distance between breeding and wintering area increases eastwards in this region.     

Density-dependent mortality of wintering Dunlins Calidris alpina through predation by Eurasian Sparrowhawks Accipiter nisus

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.     

Membrane lipid composition and the interaction of pardaxin: the role of cholesterol

Modulation by pardaxin of the phase transitions of dimyristoyl phosphatidylcholine, 1-stearoyl-2-oleoyl phosphatidylcholine or 1-stearoyl-2-oleoyl phosphatidylglycerol in the presence or absence of cholesterol was studied by differential scanning calorimetry. The transition enthalpy of each of the phospholipids was lowered by pardaxin and there was a small decrease in the transition temperature. Addition of cholesterol and pardaxin to dimyristoyl phosphatidylcholine resulted in a very marked lowering of the transition temperature. Although the peptide broadens the transition of the pure phospholipids, it sharpens the transition of mixtures of the phospholipids with cholesterol. This and the observation that pardaxin also causes the formation of crystallites of anhydrous cholesterol, suggest that the peptide promotes redistribution of cholesterol in the membrane.     

Membrane composition determines pardaxin's mechanism of lipid bilayer disruption

Pardaxin is a membrane-lysing peptide originally isolated from the fish Pardachirus marmoratus. The effect of the carboxy-amide of pardaxin (P1a) on bilayers of varying composition was studied using (15)N and (31)P solid-state NMR of mechanically aligned samples and differential scanning calorimetry (DSC). (15)N NMR spectroscopy of [(15)N-Leu(19)]P1a found that the orientation of the peptide's C-terminal helix depends on membrane composition. It is located on the surface of lipid bilayers composed of 1-palmitoyl-2-oleoyl-phosphatidylcholine (POPC) and is inserted in lipid bilayers composed of 1,2-dimyristoyl-phosphatidylcholine (DMPC). The former suggests a carpet mechanism for bilayer disruption whereas the latter is consistent with a barrel-stave mechanism. The (31)P chemical shift NMR spectra showed that the peptide significantly disrupts lipid bilayers composed solely of zwitterionic lipids, particularly bilayers composed of POPC, in agreement with a carpet mechanism. P1a caused the formation of an isotropic phase in 1-palmitoyl-2-oleoyl-phosphatidylethanolamine (POPE) lipid bilayers. This, combined with DSC data that found P1a reduced the fluid lamellar-to-inverted hexagonal phase transition temperature at very low concentrations (1:50,000), is interpreted as the formation of a cubic phase and not micellization of the membrane. Experiments exploring the effect of P1a on lipid bilayers composed of 4:1 POPC:cholesterol, 4:1 POPE:cholesterol, 3:1 POPC:1-palmitoyl-2-oleoyl-phosphatidylglycerol (POPG), and 3:1 POPE:POPG were also conducted, and the presence of anionic lipids or cholesterol was found to reduce the peptide's ability to disrupt bilayers. Considered together, these data demonstrate that the mechanism of P1a is dependent on membrane composition.