Differential survival in adult Eurasian oystercatchers Haematopus ostralegus

I explored the fitness implications of individual and sex differences in foraging strategy in the Eurasian oystercatcher Haematopus ostralegus by monitoring the survival of individually colour-ringed birds of known sex and known feeding specialisation. Over the period of this study, adult female annual and overwinter survival was significantly lower than adult male survival. However, contrary to previous findings, no differences in survival were found between birds of different feeding specialisations. Lower female survival was not due to sex differences in feeding specialisation. Possible mechanisms for sex differences in survival and the survival implications of different feeding specialisations are discussed. I conclude that sex differences in survival may be due to differences in social status. I also suggest that worm/clam feeders and mussel-stabbers, feeding specialisations previously associated with lower survival rates, may have benefited more than mussel-hammerers from milder winter temperatures in recent years.     

The influence of sediment type on the aggregative response of oystercatchers, Haematopus ostralegus, searching for cockles, Cerastoderma edule

Models that describe the dispersion patterns of predators between a series of patches that vary in prey density frequently assume that predators, in the absence of interference, will aggregate in patches with the highest prey density, at any point in time. This assumption has important implications for patterns of prey mortality, and the extent to which prey mortality is density dependent. In natural predator-prey systems, it is likely that environmental factors interact with spatial variation in prey density to influence the aggregative response of predators. We show data consistent with this idea on a population of overwintering oystercatchers foraging on cockles. There was no evidence that birds aggregated in patches with the highest biomass density of cockles. The biomass density of cockles was highest in muddy patches at the start of winter, and birds aggregated in patches that switched from being muddy at the start of winter to being sandy at some point during the winter. We argue that sediment type influences foraging costs experienced by the birds, so birds avoid feeding in muddy patches unless the fine sediment is removed from a patch, as happens during winter storms. When this happens a high biomass density of cockles suddenly becomes available and the birds aggregate in such patches. The rate of biomass loss was greatest in patches used intensively by birds for feeding, suggesting that the birds' aggregative response influences cockle mortality. We discuss the implications of our results for ideal free models.     

Genetic analysis of the population structure of socially organized oystercatchers (Haematopus ostralegus) using microsatellites

On the island of Schiermonnikoog (The Netherlands), the breeding population of oystercatchers can be divided into two groups: 'residents' and 'leapfrogs', based on their distinct social characteristics and limited probabilities of status change between breeding seasons. In order to investigate whether this social organization has caused local genetic differentiation, leapfrogs and residents were compared at eight polymorphic microsatellite loci. No significant genetic subdivision between residents and leapfrogs was observed (theta = 0.0000; 95% confidence interval (CI), -0.0027-0.0033), indicating that the oystercatcher population on the island of Schiermonnikoog has to be considered as one panmictic unit. Investigation of three additional locations in the northern part of The Netherlands did not reveal significant genetic population subdivision either (theta = -0.0005; 95% CI, -0.0045-0.0037), despite the fact that adult osytercatchers show extreme fidelity to their breeding localities. These results indicate panmixis and considerable levels of gene flow within the northern part of The Netherlands. Thus, the results from genetical analyses do not seem to be in agreement with observational data on the dispersal behaviour of breeding individuals. It is argued that the lack of population structure, locally on Schiermonnikoog as well as across larger geographical distances, is to be attributed to high levels of gene flow through dispersal of juvenile birds.     

Experimental evidence for a causal effect of pair-bond duration on reproductive performance in oystercatchers (Haematopus ostralegus)

Many studies have suggested that reproductive performance improvesduring the pair-bond, which might explain why individuals rematewith the same partner in many species. However, discussion existsabout whether the association between reproductive performanceand pair-bond duration that is reported in these studies reflectsa causal relationship. Usually it is unclear whether a positiveassociation is caused by pairs improving during their pair-bondor by high-quality pairs staying together for longer. Furthermore,reproductive performance often also depends on the age or breedingexperience of parents, which all covary with pair-bond duration.A much needed experimental approach is lacking so far. We investigatedthe effect of pair-bond duration on reproductive performancein a long-lived monogamous bird species based on natural aswell as experimental variation. The duration of oystercatcher(Haematopus ostralegus) pair-bonds, which were followed for21 years, strongly affected reproductive output, even aftercontrolling for effects of age and breeding experience. Pairsimproved during their pair-bond, and there were no indicationsof selective disappearance of low-quality pairs; however, pairsthat stayed together for very long performed badly. Experimentalremoval of one partner showed that the reproductive cost ofdivorce depended on the pair-bond duration with the old partner.In addition, after remating, the newly formed pairs stronglyimproved again, independent of the age and breeding experienceof the remated pair members. As such, this study provides thefirst experimental evidence of a causal effect of pair-bondduration on reproductive performance.     

A fundamental study revisited: Quantitative evidence for territory quality in oystercatchers (Haematopus ostralegus) using GPS data loggers

• A fundamental study by Ens et al. (1992, Journal of Animal Ecology, 61, 703) developed the concept of two different nest‐territory qualities in Eurasian oystercatchers (Haematopus ostralegus, L.), resulting in different reproductive successes. “Resident” oystercatchers use breeding territories close to the high‐tide line and occupy adjacent foraging territories on mudflats. “Leapfrog” oystercatchers breed further away from their foraging territories. In accordance with this concept, we hypothesized that both foraging trip duration and trip distance from the high‐tide line to the foraging territory would be linearly related to distance between the nest site and the high tide line. We also expected tidal stage and time of day to affect this relationship.
• The former study used visual observations of marked oystercatchers, which could not be permanently tracked. This concept model can now be tested using miniaturized GPS devices able to record data at high temporal and spatial resolutions.
• Twenty‐nine oystercatchers from two study sites were equipped with GPS devices during the incubation periods (however, not during chick rearing) over 3 years, providing data for 548 foraging trips. Trip distances from the high‐tide line were related to distance between the nest and high‐tide line. Tidal stage and time of day were included in a mixing model.
• Foraging trip distance, but not duration (which was likely more impacted by intake rate), increased with increasing distance between the nest and high‐tide line. There was a site‐specific effect of tidal stage on both trip parameters. Foraging trip duration, but not distance, was significantly longer during the hours of darkness.
• Our findings support and additionally quantify the previously developed concept. Furthermore, rather than separating breeding territory quality into two discrete classes, this classification should be extended by the linear relationship between nest‐site and foraging location. Finally, oystercatcher′s foraging territories overlapped strongly in areas of high food abundance.

     

Field tests of the accuracy of estimating prey size from bill length in oystercatchers,Haematopus ostralegus,eating mussels,Mytilus edulis

Field tests were made of the accuracy with which different observers estimated the size of mussels, Mytilus edulis, being eaten by oystercatchers, Haematopus ostralegus. Mussel size was estimated by eye by comparing their lengths with that of the bird's bill. Tests of bias were made in two ways. A ‘natural’ test involved comparing the estimated sizes of prey taken by free-living birds with the lengths of the emptied shells they left on the mussel bed. A ‘model’ test involved estimating mussel size against the bill of a model oystercatcher. Though one observer showed no bias, the other three overestimated the lengths of large mussels and underestimated those of small ones. The magnitude of the bias differed significantly between observers and, to a lesser degree, between the natural and model tests. Because of the curvilinear relationship between prey length and energy content, a bias of the magnitude found in this study could, in extreme cases, lead to serious errors in estimating a prey's energy content. It is concluded that, in studies where prey size is measured against bill length, an observer's individual bias should always be measured, preferably by the natural method. If this is not possible, then tests of the robustness of the conclusions should be made by recalculating the data assuming worst case errors.     

Taxonomic status of the Oystercatcher Haematopus ostralegus breeding in Italy

The eastern limit of H. o. longipes has moved 700 km westwards.     

Population size of Oystercatchers Haematopus ostralegus wintering in Iceland

Capsule: The first ever survey of Oystercatchers wintering in Iceland found around 11 000 individuals. This is an estimated 30% of the Icelandic population, including juveniles, suggesting that approximately 26 000 Icelandic Oystercatchers migrate to western Europe in the autumn. More Oystercatchers winter in Iceland than at similar latitudes elsewhere in Europe, which may reflect the remoteness and milder winter temperatures on this oceanic island.     

Prey depletion and foraging strategy in the Oystercatcher Haematopus ostralegus

Summary The responses of a population of Oystercatchers to their own depletion of their prey, the edible cockle Cerastoderma edule, have been examined in Strangford Lough, Northern Ireland., Cockle stocks were severely depleted each winter as a result of predation by Oystercatchers and about half the birds present in October in the main study area had left by March: the reduction was greater when the initial population of Oystercatchers was high than when it was low. Oystercatchers were initially widespread around the Lough but tended to aggregate by January into a few good sites. Within each site annual differences in the location of feeding effort were correlated with year to year variation in the location of second winter cockles.Within the feeding areas in a bay the birds apparently hunted in the short-term on the basis of expectation, ceasing to crop, the densest cockle beds once an average yield for the area has been removed; this yield was apparently estimated from the local density of the anvils on which captured cockles were broken open. This effect led to marginal areas being depleted of cockles more rapidly than high density areas, so that the birds gradually concentrated their hunting onto an increasingly restricted area of the bay. The birds initially took only second winter and larger cockles but by late winter cockle densities had fallen so much that smaller cockles were accepted: areas previously abandoned or unused for hunting were then economically viable because of the reduced expectations of the birds and were incorporated into the feeding areas.These results are seen as consistent with Royama's profitability theory.     

A silver spoon for a golden future: long-term effects of natal origin on fitness prospects of oystercatchers (Haematopus ostralegus)

1. Long-term effects of conditions during early development on fitness are important for life history evolution and population ecology. Using multistrata mark-recapture models on 20 years of data, we quantified the relation between rearing conditions and lifetime fitness in a long-lived shorebird, the oystercatcher (Haematopus ostralegus). We addressed specifically the relative contribution of short- and long-term effects of rearing conditions to overall fitness consequences. 2. Rearing conditions were defined by differences in natal habitat quality, in which there is a clear dichotomy in our study population. In the first year of life, fledglings from high-quality natal origin had a 1.3 times higher juvenile survival. Later in life (age 3-11), individuals of high-quality natal origin had a 1.6 times higher adult prebreeder survival. The most striking effect of natal habitat quality was that birds that were reared on high-quality territories had a higher probability of settling in high-quality habitat (44% vs. 6%). Lifetime reproductive success of individuals born in high-quality habitat was 2.2 times higher than that of individuals born in low-quality habitat. This difference increased further when fitness was calculated over several generations, due to a correlation between the quality of rearing conditions of parents and their offspring. 3. Long-term effects of early conditions contributed more to overall fitness differences as short-term consequences, contrary to common conceptions on this issue. 4. This study illustrates that investigating only short-term effects of early conditions can lead to the large underestimation of fitness consequences. We discuss how long-term consequences of early conditions may affect settlement decisions and source-sink population interactions.     

Field estimates of the strength of interference between oystercatchers haematopus ostralegus

Summary The rate at which oystercatchers ate mussels was higher when few oystercatchers were present. The calculated strength of this interference (m) was immensely higher than is compatible with those models which assume interference is coused solely by the time predators waste when they encounter each other.     

Anthelminthic treatment negatively affects chick survival in the Eurasian Oystercatcher Haematopus ostralegus

Eurasian Oystercatchers Haematopus ostralegus are infested with a wide range of gut parasites, but experimental evidence of their effects on host fitness is scant. We investigated prevalence of parasites, and experimentally tested the effects of gut parasites on chick survival and growth. One hundred and fifty-nine hatchlings from 66 nests were treated with a single dose anthelminthic medicine (0.5 mL Spectril + 0.0025 mL Ivomec) and compared with a sham-treated control group of 163 hatchlings from 66 nests. Unexpectedly, chicks treated with the anthelminthic drug survived less well than control chicks. Fledglings from the treated group were significantly less infected with gut parasites than untreated fledglings, although they were of similar body mass. One possible explanation for these findings is that the treatment interferes with the development of the immune system in the hatchlings. This might have caused mass mortality of treated hatchlings after the drug ceased to work and the treated chicks became susceptible to infections for the first time. Furthermore, all chicks and adults from both saltmarsh and adjacent freshwater habitat appeared free from blood parasites. Thus, in the Eurasian Oystercatcher, we found no support for the hypothesis, based on between-species comparisons, that the presence of blood parasites is related to the saltiness of the environment.     

Lipid levels in the South Island pied oystercatcher (Haematopus ostralegus finschi)

Wing lengths and some major body components were compared among the sexes and age classes of a sample of 48 South Island pied oystercatchers (Haematopus ostralegus finschi). Immature males were significantly lighter in wet weight and fat‐free weight than either immature females or adults, and were significantly lighter than adults in lean dry weight. No statistical differences were found in wing length, total lipids, or fat measure for birds in this sample. In a larger sample of 182 birds, fat measure varied similarly in the age classes, from a low in autumn to a peak in spring just before vernal migration. Massive pre‐migratory fattening typical of many small passerines does not occur in this species, but even so the average fat reserves in spring adults are sufficient for an approximate flight range of 2140 km, well in excess of need. It is hypothesised that the amount of fat deposited before vernal migration is a compromise between the energy cost of acquiring and transporting extra fat and the advantages of arrival at the breeding grounds with sufficient fat reserves for early achievement of breeding condition.     

Piping as a display of dominance by wintering Oystercatchers Haematopus ostralegus

Although Oystercatchers Haematopus ostralegus wintering in the Exe Estuary performed the piping display in any type of aggressive encounter, it was predominantly observed in two contexts. ‘Solitary piping’ occurred when one individual piped without an obvious opponent being in the immediate vicinity. In ‘piping ceremonies’, two or more individuals piped at each other for prolonged periods at close distance. Prolonged contests over mussels were usually won by the individual that had initially started piping. The most aggressive and successful birds (the dominants), engaged most often in solitary piping and piping ceremonies, using up to 10% of their time this way. This suggests that the behaviour was associated with dominance rank. The hypothesis is that the two birds are likely to start piping if their relative status has not already been established, so that a quick end to a dispute over a mussel or feeding site is not achieved. A dominant individual performs solitary piping from a distance to suppress the aggressive activities of other birds within its feeding range. If this signal fails, the dominant may join the ceremony, explaining why three was the most frequent number of birds involved in piping ceremonies. The ‘butterfly’ flight, used mainly by apparently dominant birds as they arrived on the feeding grounds, may signal their high status to birds already there. The outcome of several piping ceremonies involving birds of known dominance was consistent with this explanation. The location of piping ceremonies involving only two birds (interpreted as piping ceremonies not joined by dominants) at the extremities of the feeding ranges of the most dominant individuals is also consistent with this hypothesis, because this is where their dominance would be expected to decline and would therefore be more likely to be contested. This explanation of piping and butterfly displays assumes that individuals can recognize other individuals or quickly assess their dominance. Since dominant individuals were attacked less often than subdominants and attacking birds were rebuffed less than would be expected if recognition did not occur, we suggest that some form of dominance or individual recognition did exist. The width of the white collar did not correlate with dominance, but the individual variations in this feature may enable birds to distinguish each other and thus learn their relative status. It is intriguing that the diplomatist posture, which is typically used to rebuff misguided attackers, seems almost designed to give the opponent a particularly clear view of the white collar.     

Egg size, egg composition and reproductive success in the Oystercatcher Haematopus ostralegus

We investigated the relationship between egg size and composition and their subsequent effects on hatching and fledging success in Eurasian Oystercatchers Haematopus ostralegus on the island of Schiermonnikoog (53̀30'N, 06̀10'E) in the Dutch Wadden Sea between 1986 and 1990. Egg size variation was considerable. The ratio in egg volume of the largest over the smallest egg was more than 1.5 in each of five years. Differences between females accounted for, on average, 61% of the total variance of egg volume. Individual females produced similarly sized eggs from one year to another. Nevertheless, average egg volume per clutch declined by 0.2 cm³ per year as females aged, but this explains little of the observed variation. Female size and food supplementation had no significant effect on egg volume. In absolute terms, large eggs contain more lean dry matter and lipid than small eggs, but the proportion of both constituents decreases with egg size. Consequently, the combustible energy content of eggs increases less than proportionally with egg size. Hatchability averaged 87% and was not correlated with egg volume. Hatchling weight increased with egg size, averaging 65% of fresh egg weight. Chick survival until fledging did not increase with egg volume. Consequently, within the size range observed, large and small eggs are of comparable quality. Since both the costs and benefits of large eggs compared with small eggs seem small at best, we propose that, within the size range encountered in this population, egg size can be considered an evolutionarily neutral trait.     

The cranial and cervical osteology of the european oystercatcher Haematopus ostralegus L.

The cranial and cervical osteology of the European oystercatcher, Haematopus ostralegus L. is completely described from both whole skeletons and spirit specimens. Contrary to previous reports, the postorbital ligament shows variability in bifurcation and attachment, with the posterior branch, where present, attaching either to the suprameatic process or the zygomatic process. In addition, the quadrate ligament, which had been described as originating from the suprameatic process, appears instead to attach to the zygomatic process in some specimens. The caudal mandibular fenestra, earlier considered absent in oystercatchers, is present in all specimens examined. The cranium is additionally distinguished by a number of features which may be unique to oystercatchers. A fourth, previously unrecorded, division of the maxillopalatine strut is present, while the palatine process of the premaxilla is reduced to a thickening along the edge of the premaxillary process of the palatine. The distinctiveness of the oystercatcher cranium is particularly evident in the area of the quadratomandibular articulation, which possesses features potentially important for stabilizing the joint. A deep retroarticular notch is present, guiding a large medial jugomandibular ligament along the posterior margin of the articulation. A lateral mandibular tuberosity, which is received dorsally by an emargination of the jugal arch and quadrate, may act as an osteological brace, preventing posterior shifting of the closed mandible. The 15 cervical vertebrae are divided on the basis of structural criteria into three sections.