Clutch Predation Research Articles

Inter-island nesting dynamics and clutch survival of green turtles Chelonia mydas within a marine protected area in the Bijagós Archipelago, West Africa

Understanding spatial heterogeneity in reproductive success among at-risk populations facing localised threats is key for conservation. Sea turtle populations often concentrate at one nesting site, diverting conservation efforts from adjacent smaller rookeries. Poilão Island, Bijagós Archipelago, Guinea-Bissau, is a notable rookery for green turtles Chelonia mydas within the João Vieira-Poilão Marine National Park, surrounded by three islands (Cavalos, Meio and João Vieira), with lower nesting activity. Poilão’s nesting suitability may decrease due to turtle population growth and sea level rise, exacerbating already high nest density. As the potential usage of secondary sites may arise, we assessed green turtle clutch survival and related threats in Poilão and its neighbouring islands. High nest density on Poilão leads to high clutch destruction by later turtles, resulting in surplus eggs on the beach surface and consequently low clutch predation (4.0%, n = 69, 2000). Here, the overall mean hatching success estimated was 67.9 ± 36.7% (n = 631, 2015–2022), contrasting with a significantly lower value on Meio in 2019 (11.9 ± 23.6%, n = 21), where clutch predation was high (83.7%, n = 98). Moderate to high clutch predation was also observed on Cavalos (36.0%, n = 64) and João Vieira (76.0%, n = 175). Cavalos and Meio likely face higher clutch flooding compared to Poilão. These findings, alongside observations of turtle exchanges between islands, may suggest a source-sink dynamic, where low reproductive output sink habitats (neighbouring islands) are utilized by migrants from Poilão (source), which currently offers the best conditions for clutch survival.

Protecting nests of the Critically Endangered South Pacific loggerhead turtle Caretta caretta from goanna Varanus spp. predation

AbstractThe South Pacific subpopulation of the loggerhead turtle Caretta caretta is categorized as Critically Endangered on the IUCN Red List because of significant population declines. Five Queensland beaches support high-density nesting of this subpopulation, but egg and hatchling survival are low at some beaches because of feral and native terrestrial predators. We quantified predation of loggerhead turtle eggs by two species of goanna, Varanus panoptes and Varanus varius, at Wreck Rock beach, one of the turtle's major nesting beaches. In addition, we conducted an experiment to determine the efficacy of a nest protection device. Predation rates at Wreck Rock beach were 15.2% for treatment and 45.8% for non-treatment clutches during the 2013–2014 nesting season. A higher probability of predation (64%) was predicted for the northern beach. Although nests were only partially predated (16.4% of the total number of eggs), nest loss to predators and beach erosion (caused by a cyclone) was 91.7%. If left unmanaged, the cumulative impact of predation and other threats, including those exacerbated by climate change, can cause unsustainable loss of loggerhead turtle nests. This study provides one of the first quantitative data sets on rates of loggerhead turtle clutch predation in the South Pacific. It enhances our understanding of goanna predation impacts and identifies an efficient predator exclusion device for mitigating the effects of terrestrial predators at Wreck Rock beach, and for protecting marine turtle nests across northern Australia and globally.

Temporary Mate Removal During Incubation Leads to Variable Compensation in a Biparental Shorebird

Theoretical models predict that parents feeding offspring should partially compensate for the reduced care of their partner. However, for incubating birds, the level of compensation may depend on how reduced care changes the risk of entire brood failure, for example due to clutch predation, and on individual variation in the timing of depletion of energy stores. Although biparental incubation dominates in non-passerines, short-term manipulations of care during incubation are scarce. Here, we describe the response of 25 semipalmated sandpipers (Calidris pusilla) to an unexpected ~12-hour absence (experimental removal) of their partner in the middle of the 21-day incubation period. During the period when the removed partner would have taken over to start its regular ~12-hour incubation bout, parents compensated partially for the absence of their partner’s care (mean: 59%). However, individuals varied in their response from no to full compensation, independent of parental sex. In contrast to incubation in undisturbed nests or by uniparental species, nest attendance of compensating parents tended to be higher during the warmer part of the day. Whereas compensation was unrelated to before-experimental share of incubation, parents that left the nest from a further distance upon human approach (more aware of or more ‘responsive’ to their environment) compensated more. The quality of incubation in the after-experimental period, i.e. after return of the partner, was lower than usual, but improved quickly over time. In seven nests where the removed parent never returned, the widowed partner attended the nest for 0-10 days (median: 4), which suggests that widowed semipalmated sandpipers can adjust their incubation behavior to that observed in uniparental incubators. To conclude, our results indicate that biparental incubators are willing to tolerate a missed or irregular incubation bout of their partner and we speculate that all individuals would compensate fully, but that some fail because they deplete their energy stores, while others may be less responsive to or initially even unaware of the absence of their partner.

Effects of food abundance and early clutch predation on reproductive timing in a high Arctic shorebird exposed to advancements in arthropod abundance.

SummaryClimate change may influence the phenology of organisms unequally across trophic levels and thus lead to phenological mismatches between predators and prey. In cases where prey availability peaks before reproducing predators reach maximal prey demand, any negative fitness consequences would selectively favor resynchronization by earlier starts of the reproductive activities of the predators. At a study site in northeast Greenland, over a period of 17 years, the median emergence of the invertebrate prey of Sanderling Calidris alba advanced with 1.27 days per year. Yet, over the same period Sanderling did not advance hatching date. Thus, Sanderlings increasingly hatched after their prey was maximally abundant. Surprisingly, the phenological mismatches did not affect chick growth, but the interaction of the annual width and height of the peak in food abundance did. Chicks grew especially better in years when the food peak was broad. Sanderling clutches were most likely to be depredated early in the season, which should delay reproduction. We propose that high early clutch predation may favor a later reproductive timing. Additionally, our data suggest that in most years food was still abundant after the median date of emergence, which may explain why Sanderlings did not advance breeding along with the advances in arthropod phenology.

Camouflage and Clutch Survival in Plovers and Terns

Animals achieve camouflage through a variety of mechanisms, of which background matching and disruptive coloration are likely the most common. Although many studies have investigated camouflage mechanisms using artificial stimuli and in lab experiments, less work has addressed camouflage in the wild. Here we examine egg camouflage in clutches laid by ground-nesting Snowy Plovers Charadrius nivosus and Least Terns Sternula antillarum breeding in mixed aggregations at Bahía de Ceuta, Sinaloa, Mexico. We obtained digital images of clutches laid by both species. We then calibrated the images and used custom computer software and edge detection algorithms to quantify measures related to three potential camouflage mechanisms: pattern complexity matching, disruptive effects and background color matching. Based on our image analyses, Snowy Plover clutches, in general, appeared to be more camouflaged than Least Tern clutches. Snowy Plover clutches also survived better than Least Tern clutches. Unexpectedly, variation in clutch survival was not explained by any measure of egg camouflage in either species. We conclude that measures of egg camouflage are poor predictors of clutch survival in this population. The behavior of the incubating parents may also affect clutch predation. Determining the significance of egg camouflage requires further testing using visual models and behavioral experiments.

Increased mammalian predators and climate ch1ange predict declines in breeding success and density of Capercaillie tetrao urogallus, an old stand specialist, in fragmented Scottish forests

Low breeding success has been associated with declines in population and range of the re-introduced Capercaillie Tetrao urogallus in Scotland. Annual breeding success from 26 Scottish forests surveyed between 1991 and 2009 averaged only 0.6 chicks per female, the lowest rate recorded in 16 previous studies. Reduced breeding success was due to proportionally fewer females rearing chicks rather than a reduction in brood size. Birds bred less well in Perthshire at the southern edge of the range, where declines in indices of female and male density were highest. Only at the core of the range (Strathspey), where birds bred better, were female densities stable. Two weather variables, April temperature in the pre-breeding period (APRTEMP), and temperature at chick hatch in June (HATCHTEMP), increased over the study period. Indices of Pine Marten Martes martes increased 3.9-fold since 1995, and those of Red Fox Vulpes vulpes by 2.2-fold, whereas those of Carrion Crow Corvus corone, raptors and forest floor vegetation showed no change. Neither forest type nor forest ground vegetation appeared to influence breeding success. Instead, females reared more chicks in years when hatch time in June was drier, and in forests with lower marten and crow indices. In addition, more females reared broods in years when Aprils were cooler. Brood size was unaffected by any of the measured variables. Densities of adult birds were lower in forests with higher fox indices. Increased predation of clutches and chicks by martens and crows within these small, fragmented forests, as well as changes in climate, may explain reductions in breeding success and hence contribute to continued declines. Successful conservation of Capercaillie in remaining Strathspey strongholds may require better predator management, including a licensed removal of martens to test the hypotheses that martens contribute to reduced breeding success. This short-medium term approach will inform longer-term predator management policies and complement aspirations to increase the area and connectivity of forest habitat to benefit Capercaillie in-part through mitigating against any impact of increasing forest-edge predators.

Dependence of clutch predation rate of Eurasian reed warbler Acrocephalus scirpaceus on nesting site selection: a model study

AbstractThe article is focused on describing a possible way of how biological data can be processed, introduces the procedures used and compares the results gained from a field ornithological study. This was carried out using statistical methods of nonparametric regression with binomial classification and probit function together with the method with forward selection, and presents the most significant outcomes. Some of the interesting findings of this study are as follows: first, a lower level of reed warbler (

Aggressive neighbors and dense nesting: nest site choice and success in high-Arctic common eiders

Minimizing the risk of nest predation has led some bird species to exploit the nest defense behavior of other species. At Nasaruvaalik Island, Nunavut, Canada, some common eiders (Somateria mollissima borealis) nest within the boundaries of Arctic tern (Sterna paradisaea) colonies, while others nest elsewhere on the island, away from the terns. We tested the effects of location (within vs. outside the tern colonies), density of common eider nests, and annual variation on the nesting parameters of common eiders. Our results suggest that nesting in association with Arctic terns does not confer an obvious benefit to eiders. Such associative nesting of eiders and terns may be the result of overlapping habitat preferences between the two species, or a general scarcity of suitable nesting habitat for ground-nesting species in the high Arctic. However, eiders nesting in higher densities with other eiders had greater nest success and lower total clutch predation, indicating a positive correlation between nest density and success.

Density-dependent effects on nesting success of temperate-breeding Canada geese

Density-dependent effects on reproduction can arise through variation in habitat quality or increased competition and interference among neighbours. Negative effects have been found in avian populations and these have been mainly attributed to food limitations. In this study, we investigated whether density-dependent effects could result from either heterogeneity in habitat suitability, interference among neighbours, or predation. To test these hypotheses, we collected data over eight years in a growing population of temperate-nesting Canada geese Branta canadensis maxima. We compared different parameters of nesting success of geese between two sites characterized by different nest densities and looked at the effects of nest proximity on these parameters within each site. At the landscape level, we found density-dependent effects due to variation in habitat quality associated with predation probabilities and flooding events. At a finer scale, nesting success declined with proximity to neighbours, probably due to increased aggressive interactions among pairs. However, complete clutch predation showed both positive and negative density-dependence, due to differences in predator community at each site. We concluded that density-dependent effects reduced nesting success of Canada geese through both heterogeneity in habitat safety and agonistic interference between neighbours but that density-dependent effects could also be positive in some instances.

Spatial and temporal patterns of habitat use by Eurasian oystercatchers (Haematopus ostralegus) in the eastern Wadden Sea revealed using GPS data loggers

Detailed knowledge on species–habitat relationships is of crucial importance for the understanding of processes in marine ecosystems. Being top-predators, birds are important bio-indicators for marine systems. The aim of this study was to elucidate precise information on foraging habitat use and foraging times of oystercatchers (Haematopus ostralegus) on wide tidal flats using global positioning system (GPS) data loggers. The study was conducted to collect hints for the negative population trends in oystercatchers in the Wadden Sea. It is the first time that GPS technique has been used in a shorebird species. Although oystercatchers are known to exhibit foraging site fidelity, a number of individuals visited multiple sites. Foraging trips at night were longer, and the targeted sites were further away than those used during the day. These patterns were likely to be caused by higher risks of clutch predation by avian predators during the day that led adults to reduce their absence to defend their clutches. Our methodological approach enabled the subtle spatio-temporal patterns of habitat use to be determined on a very fine spatio-temporal scale. We suggest further potential studies using GPS data loggers that may help to reveal the reasons for the current declines in oystercatcher populations in the German Wadden Sea.

Evaluation of a glossmeter for studying the surface appearance of avian eggs

Gloss is a striking characteristic of avian eggshells that may affect both egg recognition and clutch predation. Previously, gloss has proved difficult to quantify objectively. Newly developed handheld glossmeters seem to offer a solution. We evaluated the use of a handheld glossmeter for a range of different eggshell surfaces, shapes and sizes. In all cases, their measurements were strongly affected by egg size and seem currently suitable only for very large eggs; the size of Rhea sp. eggs or larger. A glossmeter may, however, still prove useful to quantify other structures in comparative ornithology, with flat surfaces such as feathers or nest material.

Response of parasitically laying goldeneyes to experimental nest predation

Conspecific brood parasitism (CBP) has been identified as an integral life history component in birds. Recent empirical and theoretical studies suggest that nest predation is an important ecological factor affecting the occurrence and evolution of CBP; however, behavioural responses of parasites to nest predation remain unexplored. Here we report on a field experiment addressing the ability of parasitically laying common goldeneye, Bucephala clangula, females to respond to nest predation, based either on their own experience or on public information. Females that started parasitic laying in a nest and afterwards experienced partial clutch predation stopped laying in the nest. However, nests that faced partial clutch predation earlier in the season were not avoided by other parasitically laying females later in the season; these later-laying females had not themselves experienced the partial clutch predation in the nest. Hence, considering within-season responses to nest predation, a parasite uses its own experience, rather than public information, in laying decisions and nest choice. The rate of parasitic laying in the experimental nests decreased from the first to the last year of the experiment. Because the experimental nests never produced a successful clutch, this finding suggests that parasites used information on the ever-failing nature of the nests and avoided laying in them. Our results offer new experimental evidence to support the hypothesis that safety and success of nest sites play important roles in CBP.

Clutch predation in great tinamousTinamus majorand implications for the evolution of egg color

Egg camouflage has been found to reduce predation in several ground‐nesting species. Therefore, the evolution of eggs that lack camouflage in ground nesting birds is puzzling. Even though clutch predation in the tropics is high, tinamous are the only tropical ground‐nesting birds that do not build a nest and do not lay cryptic eggs. I studied predation of great tinamou clutches in a lowland tropical forest and found that risk of predation was higher during incubation when the eggs are covered by the parent, than during laying when they are exposed, suggesting that predators primarily use cues from the incubating males to locate the clutch and not cues from the eggs. Clutch size had no effect on predation rate, even though larger clutches are more conspicuous to a human observer. Predation by visual cues is likely reduced during incubation by the camouflaged plumage and high nest attendance of males. If most predators use cues from the incubating male and not the eggs to locate clutches, then conspicuous egg color may have evolved in great tinamous as an intra‐specific signal. I evaluate hypotheses that may explain the maintenance of conspicuous egg color in tinamous.

Age-specific nest-site preference and success in eiders

Variation in nest concealment is puzzling given the expected strong selection for safe nest sites. Selecting a concealed nest may decrease the risk of clutch predation but hinder parents from escaping predators, providing a possible solution to this paradox. Because the relative value of current versus future reproduction may vary with breeder age or state, nest concealment may also vary as a function of these attributes. We tested four predictions of the female and clutch safety trade-off hypothesis in eiders (Somateria mollissima): (1) nest concealment is negatively related to escape possibilities, (2) our capture rate of females is higher in covered nests, (3) egg predation is higher in open nests, and (4) overall nest success is unrelated to nest habitat. We also analysed nest microhabitat preferences and nest success relative to breeder age and body condition, controlling for nest spatial centrality. As expected, nest concealment and potential escape angle were negatively related, and capture by us, indicating female predation vulnerability, increased with nest cover. Clutch size was smaller in open nests, suggesting higher partial clutch predation, while it was larger among experienced and good-condition breeders. The probability of successful hatching was unrelated to nest habitat, positively associated with breeder experience, and negatively associated with hatching date. Experienced females selected more concealed and centrally located nests without sacrificing potential escape angles. The age-specific spatial distribution of nests on islands was unrelated to nest initiation dates, indicating no apparent competition. The age-specific preference of eiders for concealed nests may reflect declining reproductive value with age or confidence in surviving despite selecting a concealed nest. The apparently positive relationship between female age and survival and fecundity in eiders refutes the former alternative. Individual improvement in choosing safe nest sites, coupled with differential survival of individuals performing well, most likely explains age-specific nest-site preference and success.

Habitat-specific clutch size and cost of incubation in eiders reconsidered

The energetic incubation constraint hypothesis (EICH) for clutch size states that birds breeding in poor habitat may free up resources for future reproduction by laying a smaller clutch. The eider (Somateria mollissima) is considered a candidate for supporting this hypothesis. Clutch size is smaller in exposed nests, presumably because of faster heat loss and higher incubation cost, and, hence, smaller optimal clutch size. However, an alternative explanation is partial predation: the first egg(s) are left unattended and vulnerable to predation, which may disproportionately affect exposed nests, so clutch size may be underestimated. We experimentally investigated whether predation on first-laid eggs in eiders depends on nest cover. We then re-evaluated how nesting habitat affects clutch size and incubation costs based on long-term data, accounting for confounding effects between habitat and individual quality. We also experimentally assessed adult survival costs of nesting in sheltered nests. The risk of egg predation in experimental nests decreased with cover. Confounding between individual and habitat quality is unlikely, as clutch size was also smaller in open nests within individuals, and early and late breeders had similar nest cover characteristics. A trade-off between clutch and female safety may explain nest cover variation, as the risk of female capture by us, mimicking predation on adults, increased with nest cover. Nest habitat had no effect on female hatching weight or weight loss, while lower temperature during incubation had an unanticipated positive relationship with hatching weight. There were no indications of elevated costs of incubating larger clutches, while clutch size and colony size were positively correlated, a pattern not predicted by the 'energetic incubation constraint' hypothesis. Differential partial clutch predation thus offers the more parsimonious explanation for clutch size variation among habitats in eiders, highlighting the need for caution when analysing fecundity and associated life-history parameters when habitat-specific rates of clutch predation occur.

Identifying predators of eggs and chicks of Lapwing Vanellus vanellus and Black‐tailed Godwit Limosa limosa in the Netherlands and the importance of predation on wader reproductive output

Farmland bird populations in the Netherlands have shown an accelerating decline in recent years, despite extensive conservation efforts including reserves, agri‐environment schemes and protection of nests by volunteers. Although agricultural intensification is the main cause underlying these declines, there is a growing concern that the ongoing decline of grassland‐breeding shorebirds in recent years is caused or aggravated by increasing predation. Although Red Fox Vulpes vulpes and Carrion Crow Corvus corone are often accused of causing widespread breeding losses, and calls for management of these species are made, very few field data are available on the incidence of predation on grassland shorebirds and the relative importance of different predators. To obtain such data, we identified egg predators using temperature loggers and continuous video recordings of 792 clutches, and chick predators by radiotagging 662 chicks of Black‐tailed Godwit Limosa limosa and Northern Lapwing Vanellus vanellus. In total, 22 species were identified as predators of eggs or chicks, of which Red Fox, Common Buzzard Buteo buteo, Grey Heron Ardea cinerea and Stoat Mustela erminea were the most frequent. Eggs were taken primarily by mammals and chicks more often by birds. There was great variation in predation levels and species involved in predation of clutches between sites and years, but less in chick predation. Hence, there was no correlation between predation levels on clutches and those on chicks within the same sites. In sites where more then 50% of clutches were lost to predation, however, nocturnal predators took the larger share. As temporal and spatial variation on a small scale significantly influences predation levels, a site‐specific approach based on sound knowledge of the local situation will be more effective in reducing predation on farmland birds than general, country‐wide measures. Calculations based on our data indicate that eliminating only one loss factor at a time will often not reverse a local population decline, and provide a strong argument for targeting several locally limiting factors simultaneously instead of focusing on mitigation of predation alone.

Compromise between seabird enjoyment and disturbance: the role of observed and observers

SUMMARYNatural areas are increasingly visited by people, and urban human visitors expect to watch wildlife as close as possible, but this may have associated disturbance costs. Here, effects of number of visitors and bird density on flight initiation distance (FID) as a proxy of disturbance vulnerability were evaluated in the large ground-nesting yellow-legged gull,Larus michahellis. Mean FID decreased with increasing number of visitors and with increasing gull densities, suggesting that (1) ground-nesting gulls habituate to massive human presence, while retaining their antipredatory mechanisms, and (2) dense groups of gulls were more reluctant to fly away. This density effect may be due to the increased risk of clutch predation by conspecifics at high densities and, if so, FID is a reliable metric of disturbance vulnerability in ground-nesting gulls. In conclusion, set-back distances are specific to local populations and it is unnecessary to ban or restrict human visits to ground-nesting gull colonies; redistributing visits, taking into account both the number of visitors and gull density, is preferable.

Predictors of clutch predation of a globally significant avifauna in New Zealand's braided river ecosystems

AbstractPredation by introduced mammals is decimating New Zealand's indigenous fauna. Understanding factors that influence this process allows resources for predator control to be applied with maximum effect. This study examines how predation of a secondary prey species (a relatively common but declining native plover, the banded dotterel Charadrius bicinctus) varied with reductions in abundance of a major prey source (rabbits), kill‐trapping of predators, nest density and habitat complexity. Banded dotterels mostly nest in open braided riverbeds alongside a number of endemic threatened species. We measured the fate of 753 dotterel clutches exposed to predation by cats, ferrets and hedgehogs. We found key times and places of high predation risk. Immediately after widespread reduction in rabbit populations by rabbit haemorrhagic disease (RHD), clutch predation rates were almost as high (mean, 50%) as those recorded during past rabbit poisoning programmes (mean, 57%). Both rates were significantly higher than the mean predation rate of 22% without rabbit control, suggesting a shift in predator diet immediately after rabbit population declines. Unlike after rabbit poisoning, clutch predation rate remained high in the years after RHD. Other patterns observed included higher clutch predation rate where nest density was lower, suggesting that predation can potentially cause local extinction. Clutch predation was also higher along riverbed margins where vegetation was dense. There was equivocal evidence for an effect of predator kill‐trapping on clutch predation rate. Management strategies that could potentially reduce clutch predation risks include focusing predator mitigation measures during periods of rabbit decline, maintaining them for more than one breeding season if the rabbit declines are widespread (e.g. RHD epidemics), and applying greater effort at sites with relatively low nest density and along riverbed margins where predator use is more frequent.

Nest desertion by a cowbird host: an antiparasite behavior or a response to egg loss?

Natural selection can favor songbirds that desert nests containing eggs of the parasitic brown-headed cowbird (Molothrus ater). However, the high variability in desertion of parasitized nests within species is perplexing in light of the typically high costs of parasitism. Because nest desertion can also be a response to partial clutch predation, we first asked if Bell’s vireos (Vireo bellii) deserted nests in response to the presence of cowbird eggs (antiparasite response hypothesis) or to egg removal by predators and female cowbirds (egg predation hypothesis). Second, we asked whether variation in nest desertion was due to intrinsic differences among individuals or to variation in nest contents. We monitored a large number of nests (n ¼ 494) and performed a clutch manipulation experiment to test these hypotheses. The number of vireo eggs that remained in a nest was a strong predictor of desertion both within and among pairs. Neither the presence of a single cowbird egg, which leads to nest failure for this host, nor the number of cowbird eggs received in a vireo nest influenced nest desertion. Furthermore, vireos did not desert experimental nests when we immediately exchanged cowbird eggs for vireo eggs but deserted if we removed vireo eggs and replaced them with cowbird eggs the following morning. Desertion of parasitized nests by Bell’s vireos can be almost entirely explained as a response to partial or complete clutch loss and does not appear to have been altered by selection from brood parasitism. Key words: Bell’s vireo, brown-headed cowbird, egg predation, evolutionary lag, host‐parasite coevolution, Molothrus ater, Vireo bellii. [Behav Ecol]

Effects of individual life-history traits and weather on reproductive output of Black-headed Gulls Larus ridibundus breeding in the Wadden Sea, 1991–97

Capsule Annual breeding success was relatively constant and is shown to be related to clutch size and growth rate and to be positively affected by rain during egg-laying and advanced chick-rearing phase. Aims To provide the first long-term data on breeding success of Black-headed Gulls in the Wadden Sea, to analyse its intrinsic and environmental co-variates, and to re-assess the significance of the severe winter 1995/96 on reproduction. MethodsIn a relatively small colony at the Wadden Sea coast, clutches were selected randomly and enclosed to determine clutch size, egg biometrics, hatching and breeding success, and chick development in 1991 and from 1994 to 1997. Weather data (temperature, rain, wind) were related to life-history traits (clutch size, egg volume, chick development) and reproductive success. Results Mean annual breeding success was 0.7 fledglings per pair. In 1996, breeding success tended to be lower and chick growth rate was significantly lower. Hatching success was lower in small clutches, chick mortality increased with decreasing chick growth rate. Rain during the egg-laying phase increased clutch size and decreased clutch predation. Rain during the early post-natal phase impaired chick growth. Rain during the phase of linear growth affected chick growth and fledging success positively and brood predation negatively. Conclusions Breeding success of Black-headed Gulls breeding in the Wadden Sea is relatively constant between years, probably due to the use of terrestrial and marine feeding habitats. Rain may increase the availability of intertidal and terrestrial prey and thus may affect time budgets and food provisioning of parents positively. Lower breeding success in 1996 might have been caused by a relatively dry breeding season and possibly by the preceding severe winter.