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.     

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.     

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.     

Biofilm Consumption and Variable Diet Composition of Western Sandpipers (Calidris mauri) during Migratory Stopover

Many shorebird species undertake long-distance migrations punctuated by brief stays at food-rich, estuarine stopover locations. Understanding use of these food resources helps guide conservation and responsible development decisions. We determined the extent and degree to which Western Sandpiper (Calidris mauri) utilized biofilm as a food resource across a large and variable stopover location during northward (spring) migration. We investigated the spatial heterogeneity in diet composition, to determine whether shorebirds were consistently feeding on biofilm or whether diet varied between naturally and anthropogenically delineated sites. We used stable isotope analysis to estimate that biofilm conservatively comprised 22% to 53% of Western Sandpiper droppings across all sampling sites and that prey composition differed significantly between areas within the stopover location. Widespread biofilm consumption demonstrates the importance of biofilm as a dietary component. Variable diet composition suggests that habitat heterogeneity may be an important component of high quality stopover locations in the context of “state-dependant trade-offs” of Western Sandpiper population sub-groups. Future management decisions must consider and address potential impacts on the biofilm community throughout a stopover location, as single site studies of diet composition may not be adequate to develop effective management strategies for entire stopover sites.     

Survival of the fattest: body stores on migration and survival in red knots Calidris canutus islandica

Severe summer weather in Greenland and Arctic Canada in 1972 and 1974 caused very poor breeding success and elevated adult mortality in red knots Calidris canutus islandica. We show that those individual knots that are known to have survived these summers were in better than average nutritional condition shortly before departure from their late spring staging area in west Iceland. The condition index of previously banded or subsequently recovered birds captured in Iceland was positively related to the number of summers they were known to have survived. Body stores carried from the last spring staging area to the breeding grounds appear to offer Arctic-breeding shorebirds significant selective advantages: they are used for physical transformation from migration to breeding condition, and in years when weather is difficult may enable survival after arrival on the breeding grounds.     

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.     

Decision between small and large prey: reduced energy acquisition by pre-migratory purple sandpipers,Calidris maritima,on Svalbard

On Svalbard, purple sandpipers forage on the shores when preparing for autumn migration to the European continent after a period of food shortage on the tundra in summer. Their food choice was studied on a beach when they were faced with the choice between two abundant prey species: a small collembolan (springtail) and a much larger amphipod. In spite of several times higher gain of energy when foraging on amphipops, the birds did not show a distinct preference for either of the two prey species in this situation. Alternative explanations for this behaviour are discussed. We suggest that this could be a consequence of adaptations to their summer diet on Svalbard, when they rely on Collembola as food to a considerable extent. Higher variance in energy gain rate when foraging on amphipods might also be important. Collembolan densities high enough to trigger predation from birds occurred only on some days, with cloudy calm weather. This, together with the possible existence of a dietary bottleneck, may reduce the cost associated with the switch to Collembola. Accepted: 25 April 1999     

Plasma metabolites and migration physiology of semipalmated sandpipers: refueling performance at five latitudes

Long-distance bird migration is fueled by energy gathered at stopover sites along the migration route. The refueling rate at stopover sites is a determinant of time spent at stopovers and impacts the overall speed of migration. Refueling rate during spring migration may influence the fitness of individuals via changes in the probability of successful migration and reproduction during the subsequent breeding season. We evaluated four plasma lipid metabolites (triglycerides, phospholipids, β-OH-butyrate, and glycerol) as measures of refueling rate in free-living semipalmated sandpipers (Calidris pusilla) captured at non-breeding areas. We described the spatial and temporal variation in metabolite concentrations among one winter site in the Dominican Republic and four stopover sites in the South Atlantic and Mid-Atlantic Coastal Plain regions of North America. Triglycerides and β-OH-butyrate clearly identified spatial variation in refueling rate and stopover habitat quality. Metabolite profiles indicated that birds had higher refueling rates at one site in the Mid-Atlantic Coastal Plain than at three sites on the South Atlantic Coastal Plain and one site in the Dominican Republic. Temporal variation in lipid metabolites during the migration season suggested that male semipalmated sandpipers gained more weight at stopovers on the South Atlantic Coastal Plain than did females, evidence of differential migration strategies for the sexes. Plasma lipid metabolites provide information on migration physiology that may help determine stopover habitat quality and reveal how migratory populations use stopover sites to refuel and successfully complete long-distance migrations.     

Effects of feeding time constraints on body mass regulation and energy expenditure in wintering dunlin (Calidris alpina)

We examined the effects of time-restricted feeding on regulationof body mass and activity energy expenditure in captive winteringdunlin (Calidris alpina) held in outdoor aviaries at TomalesBay, California. In the first of two experiments, we comparedbirds under 24 h : 24 h (fasting : ad libitum feeding) foodrestriction with controls under continuous ad libitum feeding. In the second experiment, we compared birds under 24 h : 6 h: 12 h : 6 h (fasting : ad libitum : fasting : ad libitum)food restriction with birds under 24 h : 24 h food restriction.We estimated total energy expended on activities from dailymass balance using an additive model based on measures of grossenergy intake, thermoregulation, basal metabolism, and a sensitivity analysis of gross utilization efficiency and energy densityof reserve body tissue. Dunlin under 24 h : 24 h food restrictionovercompensated for body mass lost while fasting, increasingtheir body mass relative to controls fed ad libitum. Dunlinunder 24 h : 6 h : 12 h : 6 h food restriction were unable to recover body mass lost during the first fasting day. Whenallowed to feed, food-restricted birds reduced the amount ofenergy spent on being active and increased food intake andenergy storage relative to controls, but when forced to fast,they increased their activity energy expenditure. These patterns suggest winter body mass regulation consistent with the behaviorsof free-living dunlin in winter.     

Numbers of litters, litter size and survival in two species of microtines at two elevations

The number of litters, litter size and survival of Clethrionomys gapperi and Microtus pennsylvanicus were documented at two elevations (1450 m; 2240 m) in the Canadian Rocky Mountains to determine if those life history traits changed with the length of the breeding season. The high elevation was colder, had fewer frost-free days, and more precipitation than the low elevation. The length of the breeding season of C. gapperi and M. pennsylvanicus averaged 8 d and 34 d longer, respectively, at the low elevation than at the high elevation. In both species and at both elevations, most females survived only long enough to produce one or two litters. Litter size was significantly greater at the high elevation than at the low elevation in M. pennsylvanicus , but not different in C. gapperi. Overall, nestling survival was greater at the low elevation than at the high elevation in M. pennsylvanicus , but not different in C. gapperi. Summer and winter survival rates were not different between elevations in either species. Differences in litter size and nestling survival between elevations in M. pennsylvanicus may be related to differences in habitat quality.     

Sex-differentiated migration patterns, protandry and phenology in North European songbird populations

This study aims to investigate causes and mechanisms controlling protandrous migration patterns (the earlier breeding area arrival of males relative to females) and inter-sexual differences in timing of migration in relation to the recent climate-driven changes in phenology. Using standardised ringing data from a single site for eight North European migratory passerines collected throughout 22 years, we analysed sex-differentiated migration patterns, protandry and phenology of the entire populations. Our results show protandrous patterns for the first as well as later arriving individuals for all studied species. Males show more synchronous migration patterns compared to females and, hence, first arriving females followed males more closely than later arriving individuals. However, we found no inter-sexual differences in arrival trends as both sexes advance spring arrival over time with the largest change for the first arriving individuals. These findings seem in support of the “mate opportunity” hypothesis, as the arrival of males and females is strongly coupled and both sexes seem to compete for early arrival. Changes in timing of arrival in males and females as a response to climatic changes may influence subsequent mating decisions, with subsequent feedbacks on population dynamics such as reproductive success and individual fitness. However, during decades of consistent earlier spring arrival in all phases of migration we found no evidence of inter-sexual phenological differences.     

Changes in migration phenology and biometrical traits of Reed,Marsh and Sedge Warblers

Global environmental processes like climate change could severely affect population level migratory behaviour of long range migrant birds. We analyzed changes in migration phenology and biometrics of three closely-related long-distance migrant Acrocephalus species. We used the records of 12 063 Sedge, 12 913 Reed, and 5 409 Marsh Warblers caught and ringed between 1989–2009, at a Hungarian stopover site. Quantile regressions were used to analyse the changes in spring and autumn migration phenology. Median spring arrival date of Sedge and Reed Warblers shifted 6.5 and 7.5 days earlier, respectively. Autumn arrival of all species shifted one (Reed and Marsh Warblers) or two (Sedge Warbler) weeks later. Mean body mass of adult Reed and Marsh Warblers decreased in spring (by 0.3 and 0.2 grams, respectively) and in autumn (by 0.8 and 0.2 grams, respectively) while body mass of adult Sedge Warblers decreased only in autumn (by 0.4 grams). Mean wing length of all species increased significantly (range of change: 0.6–1 mm). Despite the fact that the studied species are closely related, all three have remarkably different migration strategies. However, similar patterns can be observed in the studied parameters, indicating that global processes may have general effects on these species, albeit through markedly different mechanisms.     

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.     

The ruv proteins of Thermotoga maritima: branch migration and resolution of Holliday junctions

In homologous recombination in bacteria, the RuvAB Holliday junction-specific helicase catalyzes Holliday junction branch migration, and the RuvC Holliday junction resolvase catalyzes formation of spliced or patched structures. RuvAB and RuvC from the hyperthermophile Thermotoga maritima were expressed in Escherichia coli and purified to homogeneity. An inverted repeat sequence with unique termini was produced by PCR, restriction endonuclease cleavage, and head-to-tail ligation. A second inverted repeat sequence was derived by amplification of a second template containing a three-nucleotide insertion. Reassociation products from a mixture of these two sequences were homoduplex linear molecules and heteroduplex heat-stable Holliday junctions, which acted as substrates for both T. maritima RuvAB and RuvC. The T. maritima RuvAB helicase catalyzed energy-dependent Holliday junction branch migration at 70 degrees C, leading to heteroduplex linear duplex molecules with two three-nucleotide loops. Either ATP or ATP gamma S hydrolysis served as the energy source. T. maritima RuvC resolved Holliday junctions at 70 degrees C. Remarkably, the cleavage site was identical to the preferred cleavage site for E. coli RuvC [(A/T)TT(downward arrow)(G/C)]. The conservation of function and the ease of purification of wild-type and mutant thermophilic proteins argues for the use of T. maritima proteins for additional biochemical and structural studies.     

Seasonal migration patterns of female sika deer in eastern Hokkaido,Japan

Fifty-seven female sika deer (Cervus nippon yesoensis), captured at the wintering area in the Shiranuka Hills in eastern Hokkaido, Japan, were radio-tracked during 1997–2001 to examine the factors affecting seasonal migration at the individual-landscape level. Ten of the 57 deers migrated between low-altitude summer home ranges and intermediate-altitude winter home ranges (the upward migrants). Twenty-nine migrated between high-altitude summer home ranges and intermediate-altitude winter home ranges (the downward migrants). Twelve used the intermediate-altitude home ranges all year round (the non-migrants). The remaining six were unknown. The summer home ranges of deer were widely scattered over an area of 5734km². Migration distances ranged between 7.2 and 101.7km. Deer showed high site fidelities to their seasonal home ranges. The upward migrants wintered in areas of less snow, higher quality of bamboo grass, and more coniferous cover than their summer home ranges. The downward migrants wintered in areas of less snow, higher quality of bamboo grass, higher winter temperature, and more southern slopes, but less coniferous cover than their summer home ranges. The non-migrants used year-round ranges with little snow, high quality of bamboo grass, and sufficient coniferous cover. We suggest that snow cover and bamboo grass are the factors affecting seasonal migration of the population and that coniferous cover is another factor for the upward migration.