Incubation routine, body mass regulation and egg neglect in the Blue Petrel Halobaena caerulea

The incubation routine and mass changes of male and female Blue Petrel Halobaena caerulea were studied at the Kerguelen Islands to investigate factors influencing the durations of incubation stints and foraging trips at sea and the factors determining nest desertion and return to the nest.
The body mass at the start of an incubation shift and also when the bird was relieved varied throughout the incubation period, whereas the mass when birds deserted the nest was stable. Birds deserted the nest when their mass decreased to threshold, independent of the duration of the fast. Temporary egg neglect was observed in successful as well as in unsuccessful breeding attempts, but it increased the risk of breeding failure. The net and daily massgained at sea during the second part of the incubation period were higher than during the first part, suggesting an increase in food availability. During the first part, the mass gained at sea and time spent foraging were inversely related to the mass of the bird before it left the burrow, whereas a similar relationship did not occur thereafter.
The results suggest the occurrence of a fixed mass threshold when birds decide to leave the nest if not relieved by their partner. The mass when a bird left its nest inffuenced the time spent foraging or mass gained when food was scarce. Although decision rules to leave the nest or return from the sea are related to body condition. the possibility of neglecting the eggs temporarily enables Blue Petrels to regulate the trade-off between risks of breeding failure and risks of an increase in adult mortality. A model for behavioural decision to stop incubating or stop feeding, based on a variable set point, is proposed.     

Regulation of foraging trips and costs of incubation shifts in the Antarctic petrel (Thalassoica antarctica)

In species where incubation is shared by both parents, the mate'sability to fast on the nest may constrain the time availablefor foraging. The decision to return to the nest should thereforebe a compromise between an animal's own foraging success andits mate's ability to fast on the nest. To examine how the bodyconditions of incubating Antarctic petrels, Thalassoica antarctica,influence both the length of foraging trips and incubation shifts,we experimentally handicapped females by increasing their flightcosts during a foraging trip by adding lead weights to theirlegs. Handicapped females spent more time at sea and had lowerbody conditions at arrival to the colony than controls, and,moreover, females in poor body condition at arrival to the colonyspent generally more time at sea than those with higher bodycondition. The prolonged time period spent at sea by handicappedfemales was associated with higher desertion rates than amongcontrols. The time the incubating mates fasted increased withtheir body condition at arrival to the colony, suggesting thata high body condition of the incubating bird may reduce theprobability of nest desertion. Accordingly, our results suggestthat the time spent foraging is adjusted to the body conditionsof both the foraging and incubating mate.     

Energy expenditure, body-weight and foraging performance of Storm Petrels Hydrobates pelagicus breeding in artificial nesting chambers

Many avian species, such as Storm Petrels Hydrobates pelagicus, are intolerant of disturbance at the nest, which complicates the collection of data relating to metabolic rate and the use of body reserves during incubation. I describe the design of an artificial nest chamber, which is simple and inexpensive to construct and facilitates the collection of such data. Eighty-one nest chambers situated in a large colony of breeding Storm Petrels had high occupancy rates (29/81 in each of 2 years), and the breeding success of birds nesting in boxes was similar to that of pairs nesting in natural crevices. Direct measurement of carbon dioxide production using standard respirometry techniques and estimations of metabolic rate based on the rates of mass loss during incubation indicated close agreement between the two methods of estimating energy consumption. Assuming the metabolic requirements during incubation are furnished entirely from stomach oil, 76% of the daily mass lost represented stomach oil catabolism. The duration of incubation shifts was unrelated to the body mass, and presumably to body reserves, of Storm Petrels on arrival at the nest. Shifts were usually terminated by the return of the foraging partner. The body mass of birds returning from foraging was relatively constant and was unrelated to the amount of time spent foraging at sea, indicating that the decision rule to return from foraging was the acquisition of a threshold level of body mass (about 31 g). There was a negative relationship between the duration of foraging trips and the body mass of Storm Petrels at departure from the nest and a positive relationship between trip duration and the net mass gain at sea. The use of nestboxes based on the design described here would have a wide variety of applications in facilitating data collection for many cavity-or burrow-nesting species which are sensitive to disturbance.     

Effects of clutch sizes and incubation stage on nest desertion in the female Common Eider Somateria mollissima nesting in the high Arctic

In bird species, one of the trade-offs between reproduction and survival appears in the parental decision to desert the nest. Nest desertion is modulated by several factors including clutch size. However, the incubation stage at which predation occurs is also an important factor. In this study, we examined whether nest desertion was linked to initial clutch size, partial clutch predation (final clutch size) and the incubation stage at which it happened in a capital breeder: the female common eider (Somateria mollissima) nesting in the high Arctic. The study was performed in Kongsfjorden in 2002 on the western coast of the Svalbard Archipelago (78°55′N, 20°07′E). We observed that nest desertion was higher when the initial clutch size was small. Also, females deserted their nests more during the first third of incubation than later. Thus, as incubation proceeded, nest desertion was less likely to occur even after egg reduction. Our results pointed out that this parental decision in female eiders seemed to depend on initial clutch size and on the date into incubation of clutch reduction. Sophie Bourgeon and Francois Criscuolo contributed equally to the work     

Sex roles, parental effort and offspring desertion in the monogamous Eurasian Curlew Numenius arquata

The reasons for female desertion of offspring and the evolution of predominantly male care among monogamous bird species are not clearly understood. We studied parental effort during the incubation and chick rearing periods in the Eurasian Curlew Numenius arquata in western Finland, and compared timing of brood desertion with other populations in Europe. Males and females contributed equally to incubation and showed no differences in the intensity of mobbing behaviour towards a potential nest predator (stuffed crow) shortly after hatching. However, females deserted their offspring approximately halfway through the brooding period ( c. 16 d after hatching), while males remained with chicks until independence ( c. 35 d). Females with late-laid clutches deserted their offspring sooner after hatching than those with clutches produced earlier in the season. Curlew females deserted younger chicks in northeast Europe, where laying dates were later, breeding seasons shorter and migration distances were longer, than in western and central Europe. We suggest that the most likely reasons for offspring desertion by females may be associated with increased female survivorship and maintenance of pairbond between years.     

Incubation and brooding rhythm of the Cape Petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica

Incubation and brooding performance of the Cape Petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica, in the 1991 -1992 austral summer is reported in detail and compared with data from two other areas. There was an inverse relationship between the mean foraging trip length and mean peak weight of chicks. Variation in shift length throughout incubation resulted from a combination of seasonal effects (food supply) and behavioural adjustment to hatching. The length of the last incubation shift was independent of its number but decreased with date of initiation. The average weight of nestattending birds increased steadily during the incubation period and at the same rate in both sexes. Females attained seasonal peak body-weight before the egg hatched, whereas males just recovered their pre-breeding weight. Males took on a slightly larger share of incubation (52%, range 40–63%) and invested more time in mutual nest attendance. Length of foraging trips varied consistently in pairs and individuals, its repeatability being twice as high in females as in males. The incubation performance of individual females (length of the first and an average foraging trip) was correlated with the size of the egg they had laid and also to subsequent chick growth and fledging success. There was a tendency in successful pairs to spend more time together at the nest and to have shorter foraging trips. I suggest that variation in breeding performance among pairs was mainly a result of individual variation in female quality, with pairs where the female contributed more to incubation being more successful. There is little evidence that egg losses were caused by parental errors.     

Human-induced behaviour in Adélie penguins Pygoscelis adeliae

Summary Adélie penguins Pygoscelis adeliae appear to be little perturbed by man. We examined the incidence of nest desertion and duration of foraging trip in Adélie penguins when manipulated and fitted with devices of differing sizes. Birds with ca. 1 cm clipped from their tail feathers stayed at sea 50% longer than unmarked controls. The length of foraging trip and incidence of nest desertion increased with increasing device volume. Penguins fitted with devices did not reduce foraging trip length to that of unpackaged birds for at least 19 days. The susceptibility of Adélie penguins to disturbance should be carefully considered when activity patterns are being studied.     

Offspring desertion and parental care in the Whiskered Tern Chlidonias hybrida

In species with biparental care, a conflict of interest can arise if one mate tries to maximize its own reproductive success at the expense of the other's. One of the mates can desert the brood to accrue a number of benefits to enhance its own fitness, leaving parental care to the remaining parent. This study is the first to describe the desertion pattern in a tern species (Sternidae). We investigated offspring desertion in the Whiskered Tern Chlidonias hybrida, a species with semi‐precocial chicks. Offspring desertion was recorded in 52% of nests prior to fledging (n = 131 nests). Females also deserted during the post‐fledging period. Of the deserters, 97% were females. Desertions started when chicks were 5 days old and no longer required intense brooding. Desertions before fledging did not affect fledging success. Provisioning rates between pair members differed, and females supplied much less food than males. Female provisioning rate affected the chances of nest desertion significantly: daily desertion rates were lower when females supplied more food. After females had deserted, males increased their provisioning rates but compensated for the loss of female care only partly in two‐ and three‐chick broods. Only in small (one‐chick) broods was compensation full. We conclude that male and female Whiskered Terns adopt different reproductive strategies in the population studied here. Females invest much less in parental care than males, providing less food and deserting more frequently. Given the ready availability of food and low predation pressure, benefits appear to accrue to females that desert; selection forces may therefore not be acting against female desertion.     

Reversed sexual size dimorphism and parental care in the Red-footed Booby Sula sula

The 'division-of-labour' hypothesis predicts that males and females perform different roles in parental care and that natural selection acts differently on each sex so as to produce different body size optima suited to their particular roles. Reversed sexual size dimorphism in avian species (females larger than males) may therefore be an adaptive consequence of different roles of males and females in parental care. We investigated patterns of nest attendance, brooding, foraging and provisioning rate in a tropical seabird, the Red-footed Booby Sula sula , a species showing a reversed sexual size dimorphism. During incubation, females attended the nest more often than males, and spent more time brooding the small chick than did males during daytime. Males and females did not differ in the average duration of their foraging trips. During incubation, there was a positive relationship between nest attendance and the duration of foraging trips in males, but not in females. During the small-chick stage, for the same time spent at the nest, males spent significantly more time than females at sea. On average, females fed the chick more often than did males. In males, there was a significant and positive relationship between the probability of feeding the chick and the duration of the foraging trip, whereas in females, this probability was much less dependent on the duration of the foraging trip. Overall, female Red-footed Boobies achieved slightly, but significantly, more parental commitment than did males. However, these sexual differences in parental participation were small, suggesting a minimal division of labour in the Red-footed Booby. Our results suggest that the division of labour hypothesis is unlikely to explain fully the adult size dimorphism in Red-footed Boobies.     

Corticosterone alone does not trigger a short term behavioural shift in incubating female common eiders Somateria mollissima, but does modify long term reproductive success

The trade-off between reproductive effort and adult survival in birds is modulated by several factors. Corticosterone and prolactin have additive effects on reproductive behaviour by stimulating foraging and parental behaviours, respectively. When incubation is associated with fasting, nest desertion is supposed to be activated by an unknown refeeding signal when body condition becomes critically deteriorated. The concomitant rise in corticosterone levels has been suggested to be the triggering factor. We tested the role of corticosterone on reproductive success by observing the effect of corticosterone implants on reproductive success and on plasma prolactin concentration in female common eiders Somateria mollissima . Implanted females showed a significant increase in corticosterone and a decrease in prolactin levels. Despite their enhanced daily body mass loss, females did not abandon incubation nor did they start to refeed in the four days following implantation. These data show that the experimentally induced rise in plasma corticosterone concentration alone does not trigger nest desertion. However, after 25 days of incubation, implanted females displayed a higher rate of egg loss, suggesting lower nest attentiveness towards the end of incubation. We suggest that the short-term effects of corticosterone may be dependent on the energy state of the bird. However, the late-induced change in reproductive success is indirectly linked to corticosterone, and we suggest that either a prolactin decrease, or a depletion in protein body reserves, may participate in the long-term adjustment of incubation behaviour in female eiders.     

Body condition and retaliation in the parental effort decisions of incubating great frigatebirds (Fregata minor)

Decisions about parental effort have the potential to be affectedby an individual's body condition and, among species with biparentalcare, by the level of effort made by one's mate. Previous studies,primarily of short-lived species, have found that a reductionin the parental effort of one pair member typically leads toa compensatory increase by the mate. However, long-lived specieswith short-term pair bonds might be expected to retaliate, ratherthan compensate, for a reduction in a mate's effort. I studiedthe factors affecting parental effort decisions during incubationby the great frigatebird, a long-lived seabird that forms newpair bonds for each breeding attempt. During incubation, malesand females took turns incubating and foraging. Individualslost mass during an incubation shift and regained this massduring the subsequent foraging bout. If an individual was lefton the nest for a long period of time while its mate was foraging,it subsequently went on a long foraging trip after being relievedby its mate, despite the fact that longer shifts were likelyto lead to nest failure. This relationship between incubationshift length and duration of subsequent foraging excursion could be due to a need to regain body condition after a longfast, or it could reflect a retaliatory response to the mate'sprolonged absence. To test these alternatives, I conducteda food supplementation experiment. Individuals engaged in along incubation shift were assigned to a control group or toa treatment group that was fed until the end of that particularincubation shift. Overall, fed birds returned from the subsequentforaging trip sooner than control birds, demonstrating thatthe relationship between incubation shift duration and foragingtrip duration is due primarily to a need to increase body mass,rather than being a retaliatory response to a mate's low levelof parental effort. However, males and females differed in theextent of their responses to the experimental treatment, indicatingthat males may also exhibit some degree of retaliation.     

Using multistate recapture modelling to assess age‐specific bottlenecks in breeding success: a case study in the greater flamingo Phoenicopterus roseus

Bird reproductive performance often increases with age or experience as a result of improved foraging skills, increased reproductive effort, improved coordination between partners, or a selection process. However, it remains unclear whether age and/or experience affect equally the successive steps of the breeding process, from egg laying to incubation and chick rearing. Using data from a long‐term study of the Camargue (southern France) population of the greater flamingo Phoenicopterus roseus, we studied the influence of age on step‐specific breeding performances during a single breeding season. We used, for the first time, multistate recapture models to evaluate the effect of age on breeding attendance (as a surrogate for breeding success) during incubation, early chick rearing and late chick rearing. Our results show a significant positive influence of age on breeding attendance, but only during the incubation period. Older parents had a higher probability than younger ones of completing incubation, whereas after the chick had hatched, the influence of parental age on breeding attendance was no longer significant. Although a high rate of nest desertion by younger flamingos during the middle of the incubation period coincided with a period of heavy rainfall, including rainfall level as a covariate did not improve the fit of the models. We discuss our results in relation to the evolution of life‐history strategies in long‐lived bird species and the influence of environmental instability.     

Why do Both Parents Incubate in the Kentish Plover?

Incubation by both parents is a common parental behaviour in many avian species. Biparental incubation is expected if the survival prospects of offspring are greatly raised by shared care, relative to the costs incurred by each parent. We investigated this proposition in the Kentish plover Charadrius alexandrinus, in which both parents incubate the clutch, but one parent (either the male or the female) usually deserts after hatching of the eggs. We carried out a mate‐removal and food supplementation experiment to reveal both the role of the sexes and food abundance in maintaining biparental incubation by removing either the male or the female from the nest for a short period of time. In some nests we provided supplementary food for the parent that remained at the nest to reduce the costs of incubation, whereas other nests were left unsupplemented. Although males spent more time on incubation after their mate had been removed, females’ incubation did not change. Notwithstanding the increased male incubation, total nest attentiveness was lower at uniparental nests than at biparental controls. However, incubation behaviour was not influenced by food supplementation. We conclude that offspring desertion during incubation is apparently costly in the Kentish plover, and this cost cannot be ameliorated with supplementary food.     

Female and male Leach's Storm Petrels (Hydrobates leucorhous) pursue different foraging strategies during the incubation period

Reproduction in procellariiform birds is characterized by a single egg clutch, slow development, a long breeding season and obligate biparental care. Female Leach's Storm Petrels Hydrobates leucorhous, nearly monomorphic members of this order, produce eggs that are between 20 and 25% of adult bodyweight. We tested whether female foraging behaviour differs from male foraging behaviour during the ~ 44-day incubation period across seven breeding colonies in the Northwest Atlantic. Over six breeding seasons, we used a combination of Global Positioning System and Global Location Sensor devices to measure characteristics of individual foraging trips during the incubation period. Females travelled significantly greater distances and went farther from the breeding colony than did males on individual foraging trips. For both sexes, the longer the foraging trip, the greater the distance. Independent of trip duration, females travelled farther, and spent a greater proportion of their foraging trips prospecting widely, as defined by behavioural categories derived from a hidden Markov Model. For both sexes, trip duration decreased with date. Sex differences in these foraging metrics were apparently not a consequence of morphological differences or spatial segregation. Our data are consistent with the idea that female foraging strategies differed from male foraging strategies during incubation in ways that would be expected if females were still compensating for egg formation.     

Buildings promote higher incubation temperatures and reduce nest attentiveness in a Neotropical thrush

Incubation is an energetically costly parental task of breeding birds. Incubating parents respond to environmental variation and nest‐site features to adjust the balance between the time spent incubating (i.e. nest attentiveness) and foraging to supply their own needs. Non‐natural nesting substrates such as human buildings impose new environmental contexts that may affect time allocation of incubating birds but this topic remains little studied. Here, we tested whether nesting substrate type (buildings vs. trees) affects the temperature inside the incubation chamber (hereafter ‘nest temperature’) in the Pale‐breasted Thrush Turdus leucomelas, either during ‘day’ (with incubation recesses) or ‘night’ periods (representing uninterrupted female presence at the nest). We also tested whether nesting substrate type affects the incubation time budget using air temperature and the day of the incubation cycle as covariates. Nest temperature, when controlled for microhabitat temperature, was higher at night and in nests in buildings but did not differ between daytime and night for nests in buildings, indicating that buildings partially compensate for incubation recesses by females with regard to nest temperature stability. Females from nests placed in buildings exhibited lower nest attentiveness (the overall percentage of time spent incubating) and had longer bouts off the nest. Higher air temperatures were significantly correlated with shorter bouts on the nest and longer bouts off the nest, but without affecting nest attentiveness. We suggest that the longer bouts off the nest taken by females of nests in buildings is a consequence of higher nest temperatures promoted by man‐made structures around these nests. Use of buildings as nesting substrate may therefore increase parental fitness due to a relaxed incubation budget, and potentially drive the evolution of incubation behaviour in certain urban bird populations.     

Regulation of foraging trips and incubation routine in male and female wandering albatrosses

The resolution of the conflict between eggcare and foraging was studied in male and female wandering albatrosses. The foraging zone and range, duration of incubation shifts and foraging trips, and associated changes in body mass were studied. Costs during incubation, expressed as the time spent incubating and the proportional loss of body mass, were similar for both sexes. The mass gained at sea was related to the duration of foraging trips, but the relationship was much less significant in males, where foraging ranges, though similar on average to those of females, were very variable. Males foraged in more southerly waters than females, and gained mass more rapidly. Only females appeared to regulate the duration of foraging trips, and this compensated for the mass lost during the incubation fast. Previous breeding experience had no influence on foraging efficiency. Egg desertion because of depletion of body reserves was very rare because birds have a wide safety margin, i.e. the difference between the average body mass when relieved and that at nest desertion. This safety margin enables the birds to compensate for the high variability in the duration of foraging trips, and is probably a reason for the high breeding success of wandering albatrosses. Decisions to return from the sea to the nest or to desert the nest are probably related to the status of body reserves, and have been selected in the large wandering albatross so that both present and future reproductive success are maximised.     

Brood desertion by female shorebirds: a test of the differential parental capacity hypothesis on Kentish plovers.

The aim of this study was to examine whether the energetic costs of reproduction explain offspring desertion by female shorebirds, as is suggested by the differential parental capacity hypothesis. A prediction of the hypothesis is that, in species with biparental incubation in which females desert from brood care after hatching, the body condition of females should decline after laying to a point at which their body reserves are too low for continuing parental care. We tested this prediction on Kentish plovers (Charadrius alexandrinus) in which both sexes incubate but the females desert from brood care before the chicks fledge. We found no changes in either the body masses or body compositions of both individual male and female plovers from early incubation and throughout early chick rearing. Furthermore, the timing of brood desertion by females was not affected by their body condition. Neither did we find gender differences in the energetic costs of incubation. There were no differences in the timing of brood desertion between experimental and control females in an experiment in which we lengthened or shortened the duration of incubation by one week. These results indicate that energetic costs do not explain offspring desertion by female Kentish plovers and that the needs of chicks for parental care rather than cumulative investment by females is what determines the timing of brood desertion.     

Body mass and clutch size may modulate prolactin and corticosterone levels in eiders

Altered body condition, increased incubation costs, and egg loss are important proximate factors modulating bird parental behavior, since they inform the adult about its remaining chances of survival or about the expected current reproductive success. Hormonal changes should reflect internal or external stimuli, since corticosterone levels (inducing nest abandonment) are known to increase while body condition deteriorates, and prolactin levels (stimulating incubation) decrease following egg predation. However, in a capital incubator that based its investment on available body reserves and naturally lost about half of its body mass during incubation, corticosterone should be maintained at a low threshold to avoid protein mobilization for energy supply. This study focused on the regulation of corticosterone and prolactin release in such birds during incubation, when facing egg manipulation (control, reduced, or increased) or a stressful event. Blood samples were taken before and after clutch manipulation and at hatching. Corticosterone levels were determined before and after 30 min of captivity. Female eiders exhibited a high hypothalamic-pituitary-adrenal sensitivity, plasma concentration of corticosterone being increased by four- to fivefold following 30 min of captivity. The adrenocortical response was not modified by body mass loss but was higher in birds for which clutch size was increased. In the same way, females did not show different prolactin levels among the experimental groups. However, when incubation started, prolactin levels were correlated to body mass, suggesting that nest attendance is programmed in relation to the female initial body condition. Moreover, due to an artifactual impact of bird manipulation, increased baseline corticosterone was associated with a prolactin decrease in the control group. These data suggest that, in eiders, body mass and clutch size modification can modulate prolactin and corticosterone levels, which cross-regulate each other in order to finely control incubation behavior.     

The influence of food supply on the breeding ecology of Kittiwakes Rissa tridactyla in Shetland

We measured the breeding performance, body condition, time budgets and foraging ranges of Kittiwakes Rissa tridactyla at Sumburgh Head, Shetland, in two years of contrasting food availability. Kittiwakes in Shetland generally feed their young almost entirely on sandeels, and fisheries data indicated that stocks of sandeels in Shetland waters were at least ten times higher in 1991 than in 1990. Fledging success of Kittiwakes was nil in 1990 and 68% of eggs laid in 1991, although clutch-size and hatching success were no different between years. Post-hatching foraging trips in 1991 were of comparable duration to those recorded at other colonies in conditions of good food supply (2–3 h), while trips recorded during incubation or post-hatching in 1990 were approximately three times longer on average than at corresponding stages of the breeding season in 1991. Radio-tracking data indicated that adults generally stayed within 5 km of the colony in 1991 but flew more than 40 km from the colony on each trip in 1990. Eggs were apparently not left unattended in either year, despite the fact that this required adults to incubate for periods in excess of 44 h in 1990. The extent to which adults were able to increase trip durations, foraging ranges and incubation shift lengths between years, while maintaining hatching success, indicates the degree to which Kittiwakes are normally buffered against adverse feeding conditions during incubation. Reduced nest attendance and lower body-condition of adults post-hatching in 1990, in conjunction with complete post-hatching breeding failure, indicate that adults were beyond the limits of their buffering capacity during chick-rearing in 1990.     

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.