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Abstract Identifying and understanding patterns in movement data are amongst the principal aims of movement ecology. By quantifying the similarity of movement trajectories, inferences can be made about diverse processes, ranging from individual specialisation to the ontogeny of foraging strategies. Movement analysis is not unique to ecology however, and methods for estimating the similarity of movement trajectories have been developed in other fields but are currently under-utilised by ecologists. Here, we introduce five commonly used measures of trajectory similarity: dynamic time warping (DTW), longest common subsequence (LCSS), edit distance for real sequences (EDR), Fréchet distance and nearest neighbour distance (NND), of which only NND is routinely used by ecologists. We investigate the performance of each of these measures by simulating movement trajectories using an Ornstein-Uhlenbeck (OU) model in which we varied the following parameters: (1) the point of attraction, (2) the strength of attraction to this point and (3) the noise or volatility added to the movement process in order to determine which measures were most responsive to such changes. In addition, we demonstrate how these measures can be applied using movement trajectories of breeding northern gannets (Morus bassanus) by performing trajectory clustering on a large ecological dataset. Simulations showed that DTW and Fréchet distance were most responsive to changes in movement parameters and were able to distinguish between all the different parameter combinations we trialled. In contrast, NND was the least sensitive measure trialled. When applied to our gannet dataset, the five similarity measures were highly correlated despite differences in their underlying calculation. Clustering of trajectories within and across individuals allowed us to easily visualise and compare patterns of space use over time across a large dataset. Trajectory clusters reflected the bearing on which birds departed the colony and highlighted the use of well-known bathymetric features. As both the volume of movement data and the need to quantify similarity amongst animal trajectories grow, the measures described here and the bridge they provide to other fields of research will become increasingly useful in ecology. Significance statement As the use of tracking technology increases, there is a need to develop analytical techniques to process such large volumes of data. One area in which this would be useful is the comparison of individual movement trajectories. In response, a variety of measures of trajectory similarity have been developed within the information sciences. However, such measures are rarely used by ecologists who may be unaware of them. To remedy this, we apply five common measures of trajectory similarity to both simulated data and real ecological dataset comprising of movement trajectories of breeding northern gannets. Dynamic time warping and Fréchet distance performed best on simulated data. Using trajectory similarity measures on our gannet dataset, we identified distinct foraging clusters centred on different bathymetric features, demonstrating one application of such similarity measures. As new technology and analysis techniques proliferate across ecology and the information sciences, closer ties between these fields promise further innovative analysis of movement data.

In many taxa, environmental changes that alter resource availability and energetics, such as climate change and land use change, are associated with changes in body size. We use wing length as a proxy for overall structural body size to examine a paradoxical trend of declining wing length within a Yellowhammer Emberiza citrinella population sampled over 21 years, in which it has been previously shown that longer wings are associated with higher survival rates. Higher temperatures during the previous winter (prior to the moult determining current wing length) explained 23% of wing length decrease within our population, but changes may also be correlated with non-climatic environmental variation such as changes in farming mechanisms linked to food availability. We found no evidence for within-individual wing length shrinkage with age, but our data suggested a progressive decline in the sizes of immature birds recruiting to the population. This trend was weaker, although not significantly so, among adults, suggesting that the decline in the sizes of recruits was offset by higher subsequent survival of larger birds post-recruitment. These data suggest that ecological processes can contribute more than selection to observed phenotypic trends and highlight the importance of long-term studies for providing longitudinal insights into population processes.

Current fishing extraction methods often generate huge quantities of dead or dying biomass that is returned to the sea in the form of discards. This practice produces a readily available clumped resource for many scavengers such as seabirds, but in the face of declining stocks and via policy change, the amount of discards produced is set to decline in the future. To understand how discards have influenced seabird foraging in the past and how birds may respond to future change requires studies examining consistent individual foraging choices. There is increasing evidence that populations may be made up of generalist or specialist foragers and this is key to the population's ability to adapt to change. Here we test for consistent individual foraging behaviour of northern gannets Morus bassanus in relation to fishing vessels and examine consequences of scavenging behaviour in terms of foraging effort and body condition. Using a combination of bird-borne bio-logging devices (GPS and Time Depth Recorders) with high resolution GPS data acquired through vessel monitoring systems on fishing boats, we examined the overlap between birds and fisheries. We found that during repeat foraging trips in the same breeding season, gannets regularly foraged at fishing boats but there were also clear among individual differences in the extent of fisheries overlap. Furthermore, we show for the first time that these differences represent consistent strategies – individual differences in scavenging were highly repeatable across multiple trips within a period of several weeks. However, despite this finding, we found no differences in foraging effort or body condition between scavengers and non-scavengers. Moreover, scavenging strategy did not appear to influence diving behaviour or vary by sex. Scavenging on discards appears to be a strategy employed consistently by a subsection of the population and future work should examine whether these specialisations persist throughout and between years and what causes these individual differences, exploring possible demographic and fitness consequences in light of global changes to fish stocks and fisheries management.

SUMMARYStudies of blood parasite infection in nestling birds rarely find a high prevalence of infection. This is likely due to a combination of short nestling periods (limiting the age at which nestlings can be sampled) and long parasite prepatent periods before gametocytes can be detected in peripheral blood. Here we examine rates of blood parasite infection in nestlings from three Columbid species in the UK. We use this system to address two key hypotheses in the epidemiology of avian haemoparasites: first, that nestlings in open nests have a higher prevalence of infection; and second, that nestlings sampled at 14 days old have a higher apparent infection rate than those sampled at 7 days old. Open-nesting individuals had a 54% infection rate compared with 25% for box-nesters, probably due to an increased exposure of open-nesting species to dipteran vectors. Nestlings sampled at 14 days had a 68% infection rate compared with 32% in nestlings sampled at 7 days, suggesting that rates of infection in the nest are high. Further work should examine nestlings post-fledging to identify rates of successful parasite infection (as opposed to abortive development within a dead-end host) as well as impacts on host post-fledging survival and behaviour.

SUMMARYStudies incorporating the ecology of clinical and sub-clinical disease in wild populations of conservation concern are rare. Here we examine sub-clinical infection by Trichomonas gallinae in a declining population of free-living European Turtle Doves and suggest caseous lesions cause mortality in adults and nestlings through subsequent starvation and/or suffocation. We found a 100% infection rate by T. gallinae in adult and nestling Turtle Doves (n = 25) and observed clinical signs in three adults and four nestlings (28%). Adults with clinical signs displayed no differences in any skeletal measures of size but had a mean 3·7% reduction in wing length, with no overlap compared to those without clinical signs. We also identified T. gallinae as the suggested cause of mortality in one Red-legged Partridge although disease presentation was different. A minimum of four strains of T. gallinae, characterized at the ITS/5·8S/ITS2 ribosomal region, were isolated from Turtle Doves. However, all birds with clinical signs (Turtle Doves and the Red-legged Partridge) carried a single strain of T. gallinae, suggesting that parasite spill over between Columbidae and Galliformes is a possibility that should be further investigated. Overall, we highlight the importance of monitoring populations for sub-clinical infection rather than just clinical disease.

Commercial selective logging and the conversion of primary and degraded forests to agriculture are the biggest threats to tropical biodiversity. Our understanding of the impacts of these disturbances and the resulting local extinctions on the functional roles performed by the remaining species is limited. We address this issue by examining functional diversity (FD), which quantifies a range of traits that affect a species' ecological role in a community as a single continuous metric. We calculated FD for birds across a gradient of disturbance from primary forest through intensively logged forest to oil palm plantations on previously forested land in Borneo, Southeast Asia, a hotspot of imperilled biodiversity. Logged rainforest retained similar levels of FD to unlogged rainforest, even after two logging rotations, but the conversion of logged forest to oil palm resulted in dramatic reductions in FD. The few remaining species in oil palm filled a disproportionately wide range of functional roles but showed very little clustering in terms of functional traits, suggesting that any further extinctions from oil palm would reduce FD even further. Determining the extent to which the changes we recorded were due to under-utilization of resources within oil palm or a reduction in the resources present is an important next step. Nonetheless our study improves our understanding of the stability and resilience of functional diversity in these ecosystems and of the implications of land-use changes for ecosystem functioning.

The transition to independence is a critical period of development and a focus of parent–offspring conflict over the optimum level of parental care, but there is continuing uncertainty over how much this transition is influenced by parents or offspring. We experimentally cross-fostered Manx shearwater chicks differing in age by 10–14 days and tested two predictions: 1) food-provisioning rate in the period leading up to fledging is related to the duration of parental care rather than to the age of chicks; 2) parents protect themselves from exploitative offspring by becoming insensitive to begging behavior of chicks over the period leading up to fledging. We also examined whether fledging age was under endogenous hormonal control or influenced mainly by parents. Switching chicks had no effect on fledging age, which was mainly controlled by an internal mechanism linked to a marked and rapid increase in corticosterone secretion, with no difference among treatments in the timing or magnitude of this increase. In contrast, parents reduced their frequency of food delivery according to the number of days elapsed since they started provisioning, regardless of age of chicks and despite younger foster chicks having lower body condition and begging more intensely than older foster chicks or controls. These data provide clear experimental evidence of parent–offspring conflict over parental feeding frequency in late chick development and hence chick body condition at fledging. As predicted, parents resolved this conflict in their favor by responding much less to begging over the period prior to fledging than at earlier stages of chick development.

The inclusion of carbon stock enhancements under the REDD+ framework is likely to drive a rapid increase in biosequestration projects that seek to remove carbon from the atmosphere through rehabilitation of degraded rainforests. Concern has recently been expressed, however, that management interventions to increase carbon stocks may conflict with biodiversity conservation. Focusing on a large-scale rainforest rehabilitation project in northern Borneo, we examine the broad impacts of selective logging and subsequent carbon enhancement across a wide range of invertebrate fauna by comparing the abundance of 28 higher-level taxa within two separate rainforest strata (leaf-litter and understorey) across unlogged, naturally-regenerating and rehabilitated forest. We additionally assess changes in functional composition by examining responses of different feeding guilds. Responses of individual taxa to forest management were idiosyncratic but logging resulted in more than a 20% increase in total invertebrate abundance, with fewer than 20% of taxa in either stratum having significantly lower abundance in logged forest. Rehabilitation resulted in a marked reduction in abundance, particularly among leaf-litter detritivores, but overall, there were much smaller differences between unlogged and rehabilitated forest than between unlogged and naturally regenerating forest in both total invertebrate abundance and the abundances of different feeding guilds. This applied to both strata with the exception of understorey herbivores, which were more abundant in rehabilitated forest than elsewhere. These results support previous data for birds suggesting that carbon stock enhancement in these forests has only limited adverse effects on biodiversity, but with some impacts on abundance within particular guilds.

Explanations for the coexistence of migratory and sedentary (resident) birds are much debated: some authors have suggested that migrants make use of surplus resources not exploited by residents, whereas others have recorded ecological shifts in residents during periods when migrants are present. These shifts have been ascribed to the presence of migrants but could equally reflect seasonal variation in the distribution of resources required by sedentary species. Here we assess how the abundance of migrants affects the territorial behaviour and habitat use of resident Sardinian Warblers Sylvia melanocephala at a stop-over site on the Rock of Gibraltar. There was a significant positive relationship between the abundance of migrants and the distances moved, time spent calling and the species and functional diversity of plants used by Sardinian Warblers on a daily basis, suggesting that migrants have a significant impact on residents.

We examined how short-term impacts of experimentally increased foraging effort by one parent reverberate around the family in a monomorphic seabird (little auk, Alle alle), and whether these effects depend on offspring sex. In many species, more effort is required to rear sons successfully than daughters. However, undernourishment may have stronger adverse consequences for male offspring, which could result in a lower fitness benefit of additional parental effort when rearing a son. We tested two alternative hypotheses concerning the responses of partners to handicapping parents via feather clipping: partners rearing a son are (1) more willing or able to compensate for the reduced contribution of their mate, or (2) less willing or able to compensate, compared to those rearing a daughter. Hypothesis 1 predicts that sons will be no more adversely affected than daughters, and the impact on parents will be greater when rearing a son. Hypothesis 2 predicts that sons will be more adversely affected than daughters, and parents raising a son less affected. Although experimental chicks of both sexes fledged in poorer condition than controls, sons attained higher mass and more rapid growth than daughters in both groups. Clipped parents lost a similar proportion of their initial mass regardless of chick sex, whereas partners of clipped birds lost more mass when rearing a son. These results support hypothesis 1: impacts of increased foraging effort by one parent were felt by offspring, regardless of their sex, and by the partners of manipulated birds, particularly when the offspring was male.

Movement patterns of predators should allow them to detect and respond to prey patches at different spatial scales, particularly through the adoption of area-restricted search (ARS) behaviour. Here we use fine-scale movement and activity data combined with first-passage time (FPT) analysis to examine the foraging strategy of northern gannets Morus bassanus in the western North Sea, and to test the following hypotheses: (i) birds adopt a hierarchical foraging strategy characterized by nested ARS behaviour; (ii) the locations and characteristics of ARS zones are strongly influenced by physical oceanography; (iii) the initiation of ARS behaviour is triggered by the detection and pursuit of prey; (iv) ARS behaviour is strongly linked to increased foraging effort, particularly within nested ARS areas.Birds on 13 of 15 foraging trips adopted ARS behaviour at a scale of 9.1 +/- 1.9 km, and birds on 10 of these 13 trips adopted a second, nested ARS scale of 1.5 +/- 0.8 km, supporting hypothesis 1 above. ARS zones were located 117 +/- 55 km from the colony and over half were within 5 km of a tidal mixing front similar to 50 km offshore, supporting hypothesis 2 above. The initiation of ARS behaviour was usually followed after only a short time interval (typically similar to 5 min) by the commencement of diving. Gannets do not dive until after they have located prey, and so this pattern strongly suggests that ARS behaviour was triggered by prey detection, supporting hypothesis 3 above. However, similar to 33% of dives in mixed coastal water and 16% of dives in stratified water were not associated with any detectable ARS behaviour. Hence, while ARS behaviour resulted from the detection and pursuit of prey, encounters with prey species did not inevitably induce ARS behaviour. Following the initiation of ARS behaviour, dive rates were almost four times higher within ARS zones than elsewhere and almost three times higher in zones with nested ARS behaviour than in those without, supporting hypothesis 4 above and suggesting that the foraging success of birds was linked to their ability to match the hierarchical distribution of prey.

Sexual differences in food provisioning rates of monomorphic seabirds are well known but poorly understood. Here, we address three hypotheses that attempt to explain female-biased food provisioning in common guillemots Uria aalge: (1) males spend more time in nest defence, (2) females have greater foraging efficiency, and (3) males allocate a greater proportion of foraging effort to self-maintenance. We found that males spent no more time with chicks than females but made longer trips and travelled further from the colony. There was extensive overlap between sexes in core foraging areas, indicating that females were not excluding males from feeding opportunities close to the colony. However, as a result of their longer trips, the total foraging areas of males were much greater than those of females. There was no difference between sexes in overall dive rate per hour at sea, in behaviour during individual dives or in a number of other measures of foraging efficiency including the frequency, depth and duration of dives and the dive: pause ratio during the final dive bout of each trip, which was presumably used by both sexes to obtain prey for the chick. These data strongly suggest that sexes did not differ in their ability to locate and capture prey. Yet males made almost twice as many dives per trip as females, suggesting that males made more dives than females for their own benefit. These results support the hypothesis that female-biased food provisioning arose from a difference between sexes in the allocation of foraging effort between parents and offspring, in anticipation of a prolonged period of male-only post-fledging care of the chick, and not from differences in foraging efficiency or time spent in nest defence.

According to life history theory, it is particularly important for long-lived iteroparous species to balance effort invested in reproduction against risks to future reproductive potential, leading to a conflict between investment in current versus potential future offspring. If providing care for offspring entails a risk of predation, parents must compromise between ensuring offspring growth and survival and ensuring their own survival. We addressed three hypotheses for the outcome of this conflict, which focus on different costs and benefits, and relate variation in parental effort to: (1) the parent's own risk of predation; (2) the reproductive value of the offspring; and (3) the harm that offspring would suffer in the absence of such care. We tested these hypotheses using Manx shearwaters, Puffinus puffinus, which, in common with many burrow-nesting seabirds, are active above ground only at night as a means of reducing attacks by diurnal avian predators. We found that parents were far less likely to return to the colony on moonlit nights, when they were vulnerable to such predators, and tended to delay their arrival until after moonset, which strongly supported hypothesis (1). Also, parents of chicks in good condition were more likely to attend the burrow on moonlit nights than those of chicks in poor condition, which supported hypothesis (2) but not hypothesis (3). Growth of chicks was suppressed on clear nights around the full moon, such that they experienced well-below average growth during one week in four. This could help explain the accumulation of fat stores by the chick that has evolved in some nocturnal seabirds.

The pattern of chick feeding in the Fulmar Fulmarus glacialisat St Kilda, Scotland, was examined by repeated weighing of chicks throughout 14 consecutive days during the first half of the chick-rearing period in 1994. After correcting for metabolic weight losses, the sizes of positive mass increments between weighings were used to assess meal sizes and feeding frequency for each chick. Individual meals fed to chicks averaged 80.8 g (s.d. ± 21.0 g), or approximately 10% of adult mass. Each chick received 0 to 4 meals per day, with an average of 1.9 meals per chick per day, giving an average interval of around 25 h between meals delivered by each parent. The distribution of time intervals between feeds for each chick (whether single or double meals) followed a negative exponential function with a maximum value of 80 h. These results are not compatible with the idea that the purpose of large fat deposits in procellariiform chicks is to guarantee survival over long intervals between feeds. Over 14 days, the chicks' mean daily food requirements for zero-growth increased from 98 g to 160 g. This corresponded with an increase in feeding frequency but not meal size. Chicks with lower scores for body condition after feeding by both parents received more meals during the subsequent 16 h and had shorter intervals to the next feed, indicating that adults regulated feeding frequency in accordance with chick condition at the previous feed. This does not agree with the hypothesis that lipid accumulation by nestling Procellariiformes is a response to stochastic variation in food delivery associated with an absence of regulation. In view of the diversity of growth and feeding patterns present among the Procellariiformes, it is possible that lipid accumulation in this group does not have a unitary explanation.

Growth and foraging strategies in procellariiforms show a great deal of variation, but the fulmarine petrels are notable in that chicks are fed frequently and develop unusually rapidly. This study examined age-related and daily variation in provisioning of the Northern Fulmar Fulmarus glacialis throughout the chick-rearing period at Fair Isle in 1997. In common with many other petrels, meal mass showed an initial rise with age, probably because of a gradual increase in chick gut capacity, but then levelled off. By comparison, feeding frequency showed little age-specific variation until chicks reached the oldest age-class, when the number of meals declined to less than a third of the previous level as chicks underwent mass recession prior to fledging. Compared with the limited day-to-day variation in mean provisioning rates for the whole sample, food delivery to particular chicks was much more variable, suggesting that differences in feeding rates were determined by stochastic factors influencing the feeding success of individual parents. The caloric density of feeds and their size in relation to adult mass were lower in Northern Fulmars than in most other Procellariiformes. This implies that adults are not heavily dependent on stomach oil formation to raise the energy content of the payload, but rely on a high feeding frequency to maintain adequate rates of energy transfer to chicks.

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.

Birds such as great skuas Catharacta skua adapted for successful breeding at high latitudes may experience problems of heat dissipation in mild climates. Great skuas spend time bathing at freshwater sites close to breeding territories and here, we examine impacts of heat stress on bathing, foraging and nest attendance of adults during three breeding seasons with marked variation in the availability of prey (1-group sandeels Ammodytes marinus). Adults exhibited diurnal variation in bathing activity that matched heat-stress conditions. Moreover more birds bathed on days of higher average heat stress, suggesting that bathing plays a role in thermoregulation. Bathing numbers were lower in years of poor food availability, when adult attendance at territories was low, probably because lower attendance reduced the opportunity for parents to bathe without leaving chicks unattended. Chicks are normally guarded by female parents and fed by males but under conditions of low food availability territorial attendance of breeding pairs was particularly low on days of high heat stress, with chicks regularly left unattended at air temperatures exceeding 14 degrees C. Unattended chicks are at risk of being killed by neighbouring conspecifics and survival of chicks to fledging was low in the two years of low sandeel stocks. Our study indicates that for great skuas, indirect effects of climate change on prey stocks and direct effects on heat stress experienced by adults may be additive

Age-specific variation in the reproductive performance of great skuas in Shetland, U.K., was investigated during a period of fluctuating sandeel availability between 1988 and 1993. Increased sandeel abundance was associated with earlier laying dates, increased clutch volumes and improved fledging success. Sandeel abundance had no effects on clutch size and hatching success. Parental age improved haying date and clutch size in a nonlinear manner. Laying date became earlier in the youngest age classes, with the effect being negligible in older birds. Clutch size increased with age up to 18 and then declined in older birds. Clutch volume increased with age and the probability of nest predation declined with age. There were no significant interactive effects of age and year on laying date nor clutch volume, indicating that birds of all ages benefited equally from increases in sandeel abundance. This suggests that lack of breeding experience rather than deficient foraging skills are responsible for young birds laying later. There was a significant interactive effect of year and age on fledging success that was associated with changes in sandeel availability. Success was uniformly low for birds of all ages in 1988-90 when sandeel abundance was low. Success increased with age in 1991 and 1992 during which time food supply improved. During 1993 food supply was abundant and fledging success was high in all age classes. Age-specific improvements in laying date, clutch size and nest predation were best explained by learning of skills associated with experience of breeding. Improvements in fledging success with age were probably best explained by the learning of skills associated with foraging.

In animals that produce few offspring during their lifetime, the ability to raise high-quality offspring through the provision of parental care is particularly important for individual fitness. In this paper, we use repeatability analysis of data from two separate time-periods, in the 1970s and the 1990s, to examine parental and environmental effects on chick growth and food provisioning in a long-lived seabird, the Manx shearwater Puffinus puffinus. We test the hypothesis that asymptotic body masses of chicks are most strongly influenced by an individual's genotype, with growth rates intermediate and food provisioning rates most strongly affected by environmental conditions during growth. Peak body masses of chicks raised by the same parents showed significant repeatability in both samples, whereas mass growth rates up to the attainment of peak mass showed significant repeatability only in the 1970s. Several different measures of food provisioning by parents showed no individual consistency in either time-period, in keeping with our predictions. Food provisioning rate was a major determinant of annual variation in chick growth, and so it may seem surprising that Manx shearwaters showed repeatable patterns of chick growth but not of food provisioning, and several possible explanations for this pattern are discussed.

Sex differences in the foraging behaviour of adults have been observed in a number of sexually size-dimorphic birds, and the usual inference has been that these sex-specific differences are driven primarily by differences in body size. An alternative explanation is that foraging differences result from sex differences unrelated to size, such as sex-specific nutritional requirements. To examine these alternative hypotheses, the foraging behaviour of parents was compared between two sympatric and congeneric species of seabird, the Brown Booby Sula leucogaster , which is highly sexually size-dimorphic (females 38% larger) and the Red-footed Booby S. sula , in which sex differences in body size are less marked (females 15% larger). Using temperature and depth loggers, we found that there were highly significant differences in the foraging trip durations and diving behaviour of male and female Brown Boobies. These sex differences were less marked in Red-footed Boobies. Thus, our interspecies comparison revealed that the magnitude of the difference between the sexes matched the sexual size dimorphism of the species, providing support for the size hypothesis.

The foraging behaviour of any animal is intricately linked to ecological constraints. Extrinsic factors, such as daylength, may limit the time available for foraging. Intrinsic factors, such as the coordination of mates at a nest site, may exacerbate this limitation. Using activity loggers, we examined the effects of extrinsic and intrinsic limitations on the foraging behaviour of the sympatric red-footed booby, Sula sula, and brown booby, Sula leucogaster, during incubation, and tested the prediction that individuals would work harder when time available for foraging was limiting. Both species share incubation duties and forage by day. Brown boobies made significantly shorter foraging trips than red-footed boobies, and thus, both parents could forage during the same day. Therefore, departure time and time available for foraging will also be influenced by when the partner returns to the colony. Overall foraging trip duration decreased linearly with respect to departure time. Our study suggests that the time available for foraging is limited by a combination of extrinsic and intrinsic processes in both species. Crucially, the rate of flying and diving, indicators of prey search and pursuit, respectively, increased with decreasing time available for foraging, whereas time spent inactive on the sea surface decreased, as did the time between departure and diving. Our data therefore suggest that boobies adjust their foraging behaviour according to extrinsic constraints.

Many species of seabirds are known to undertake foraging trips that vary in duration, lasting from a few hours up to several days. However, the important question of how individuals allocate their time during foraging trips of different durations has received relatively little attention until recently. Using activity loggers, we examined the foraging behavior of chick-rearing northern gannets, Morus bassanus (L., 1758), during trips of different durations, and tested predictions concerning how foraging activity varies across trips. There was no evidence of a relationship between dive frequency during the first 3 h of a trip and trip duration, suggesting that the decision to continue on a longer trip was not affected by an adult's initial rate of encounter with prey. Flight constituted approximately 50% of total trip time, and the dive rate of birds per daylight hour was apparently unaffected by trip duration. Birds dived at similar rates on the outward and return sections of their foraging trips, which suggests that they may have been "topping up" on food on their return. Overall our results suggest that, unlike other pelagic seabirds, northern gannets at the Bass Rock do not adjust their individual foraging strategies among trips of different durations.

Sex differences in provisioning rules have been found in a number of bird species, but the reasons remain unclear. Studies of begging in species with single-chick broods exclude the influence of nestling competition and may provide especially useful models for the study of signalling during parent–offspring conflict. We tested whether sex differences in provisioning rules occur in a species with an obligate brood size of one, the Manx shearwater. We found that chicks conveyed information about their body condition through begging, but male and female parents responded differently to that information. Females varied meal sizes according to the begging intensity of the chick and adjusted subsequent trip duration according to the chick's body condition after feeding, but males did not alter meal size or adjust trip duration. We discuss these findings in the context of recorded differences in the contributions to food provisioning by male and female parents, and we discuss why females may respond more sensitively than males to changes in chicks' nutritional requirements.

Among species where there is a risk to leaving offspring unattended, parents usually take alternating shifts guarding their young. However, they may occasionally exhibit brood neglect by leaving their offspring unattended at the nest. To investigate this phenomenon further, we examined the foraging behavior of the northern gannet (Morus bassanus) during chick-rearing. This species has a prolonged nestling period (13 weeks) and the single chick is usually guarded by one or other of its parents, because unattended chicks are frequently attacked by conspecifics. We tested the prediction that the foraging behavior of adults when they left their offspring alone at the nest (unattended trips) would differ in character to when adults left offspring with their partner (attended trips). Brood neglect typically occurred after a longer-than-average attendance period at the nest. Unattended trips were, on average, about half the duration of attended trips, and therefore much closer to the colony. There was also a difference in departure direction between attended and unattended trips, with unattended trips tending to be northeast of the colony. Chicks were fed by parents after both attended and unattended trips, but the frequency and the duration of unattended trips increased as chicks grew older whereas the duration of attended trips decreased as chicks grew. These results indicate that parents may be making a trade-off between risk of attack to their offspring and food-intake rate, and that the solution to this trade-off is dependent on chick age.

In many vertebrates, male offspring are affected more than female offspring by adverse conditions during growth, resulting in facultative adjustment of offspring sex ratio by parents in response to social and environmental conditions during breeding. The greater vulnerability of male offspring is generally attributed to their higher energy requirements associated with their larger size, although greater sensitivity to adverse conditions could be related to other factors such as negative effects of androgens on male physiology. To control for sexual differences in body size, we examined variation in offspring sex ratio in the Common Tern Sterna hirundo, a species with negligible sexual size dimorphism. In this species, the last-laid egg (termed the c-egg) is smaller than the first two and hatches last, so that the chick obtains relatively little food and hence has a low probability of survival to fledging. This species thus provides a powerful model for examining sex-linked mortality and sex ratio variation under natural conditions. We found that the sex ratio of c-eggs, but not of earlier laid eggs, was significantly biased in favour of females. Chicks hatched from c-eggs (termed c-chicks) had low survival but female c-chicks had significantly higher survival than male c-chicks. These data provide strong evidence that factors other than sexual size dimorphism are responsible for producing greater vulnerability of male offspring to adverse conditions during growth.

The costs and benefits of dominance status have been investigated in the past and it has generally been reported that subdominant birds are at a nutritional disadvantage owing to their low dominance status. The nutritional condition of birds during winter can be important in determining their likelihood of survival. This is particularly so in small passerines that are sensitive to severe weather conditions. Ptilochronology is an accurate method used to produce a long-term estimate of body condition spanning the number of days that it takes to grow a new feather. Ptilochronology was used during this study to estimate the nutritional condition of Coal Tits Pares ater during one winter and how condition was affected by dominance status. Dominant Coal Tits produced poorer quality feathers, which they grew at a slower rate, than did subdominant conspecifics. This study highlights a nutritional cost to high dominance status that could have long-term consequences because the induced tail feathers will not be replaced for at least 5 months.

Both within and among seabird species, different aspects of breeding biology may respond to changes in prey availability in distinct ways, and the identification of species-specific breeding parameters that are sensitive to food availability is useful for monitoring purposes. We present data from a 5-year study (1995–1999) of the breeding ecology of Horned Puffins (Fratercula corniculata) in Alaska. The El Niño – Southern Oscillation event of 1997–1998 provided an opportunity to examine the sensitivity of various breeding parameters to a reduction in prey availability caused by the anomalous oceanographic conditions of 1998. Horned Puffins were able to maintain high fledging success (83–97%) over the 5 years of the study, despite the poor local feeding conditions in 1998. The rate of increase in chick mass was lowest in 1998, and evidence suggests that chicks also fledged at the youngest ages in that year. The impacts of reduced food availability on growth varied among body structures, suggesting differential allocation of energy and nutrients. There was no variation among years in either chick diet or the mass of food loads delivered by adults. We suggest that rates of chick growth, specifically mass increase, may be a good parameter to measure for use in monitoring Horned Puffins.

Chick growth and food provisioning of northern fulmars Fulmarus glacialis were studied at Fair Isle, Shetland, in 2 years of contrasting environmental conditions. Low sandeel Ammodytes marinus availability resulted in very low breeding success in many seabirds in Shetland in 1998, whereas breeding success in 1997 indicated a normal season. Fulmars at Fair Isle experienced lower chick survival in 1998 than in 1997. There were also significant differences in diet and food provisioning in the 2 years; sandeels were the most abundant prey items in 1997, but were completely absent from the diet in 1998. High nestling mortality occurred shortly after hatching in 1998. Those chicks that survived this episode had previously been heavier for their age, and remained heavy throughout development in comparison to chicks in 1997, despite a continued absence of sandeels from the diet. Surviving chicks in 1998 were subsequently fed comparatively small meals at frequent intervals, but with no overall difference in the amount of food delivered. We discuss the implications of differential mortality and resulting growth patterns for the use of seabird breeding parameters to monitor marine environments.

The survival rates of breeding adult Great Skuas Catharacta skua were examined at Foula, the largest colony in the world, where numbers have been declining since the late 1970s. Resightings of colour-ringed breeding adults over a 12-year period were analysed using Cormack-Jolly Seber models to estimate survival rates. Annual survival rates averaged 0.89 but varied among years between 0.82 and 0.93, with annual variations being temporally associated with variations in sandeel abundance during the breeding season. Most birds appeared to die outside the breeding season and so it is possible that nutritional stress and reproductive costs of breeding in years of poor food supply affect survivorship on migration or in the wintering range. Survival rates of adult Great Skuas were affected by their age according to a quadratic equation, with survival increasing significantly with age from 0.73 in 5-year-old-birds to between 0.85 and 0.96 in birds from 7 to 22 years old, with a sharp decline to between 0.75 and 0.87 in birds over 22 years old. Year effects were evident when controlling for age, indicating that annual variations in survival rates are not explained by changes in age-composition of the marked population among years.

This study combines conventional dietary assessment with stable isotope techniques to describe Great Skua (Stercorarius skua) diet. Stable carbon and nitrogen isotope ratios in feathers of Great Skua chicks collected over three years were analyzed in conjunction with pellet and regurgitate collections. A significant drop in trophic status was detected in 1997, which likely resulted from an increase in herring and mackerel in the diet. These items were almost certainly obtained from a commercial trawler, as evidenced by a significant drop in territorial attendance during the ship's absence. Feathers yielded significantly different δ13C values among years, and part of this may have been related to a period of enhanced phytoplankton growth during 1996. A combined approach, such as the one described here, is likely to become increasingly useful in elucidating the diets of polyphagous birds. Variación Anual en la Dieta de Stercorarius skua: La Importancia de Pesqueros Comerciales y Predación de Aves Marinas Revelada por la Combinación de Análisis de Dieta Resumen. Este estudio combina evaluaciones de dieta convencionales con técnicas de isótopos estables para describir la dieta de Stercorarius skua. Analizamos la proporción de isótopos estables de carbono y nitrógeno en las plumas de pichones de S. skua colectadas durante más de tres años, en conjunto con colecciones de pellets y regurjitados. Un descenso significativo en la posición trófica fue detectado en 1997 probablemente como consecuencia del incremento de arenque y caballa en la dieta. Estos ítems fueron casi con seguridad obtenidos de un pesquero de arrastre comercial, como lo sugiere la caída significativa de la presencia en el territorio durante la ausencia del barco. Las plumas generaron valores significativamente diferentes de δ13C entre años, parte de lo cual puede haber estado relacionado con un período de gran crecimiento del fitoplancton durante 1996. Un enfoque combinado como el descripto aquí será probablemente cada vez más útil para elucidar la dieta de aves polífagas.

In many species of bird, nestlings undergo a period of mass recession, associated with a large reduction in food provisioning, before they fledge. This has been ascribed to parental desertion, although it could equally result from voluntary limitation of feeding (i.e. anorexia) among chicks. We examined the interactions between Manx shearwater, Puffinus puffinus, parents and chicks, to test whether the prefledging reduction in feeding was controlled by parents, chicks or both. In a cross-fostering experiment, chicks of different ages were exchanged between nests. We predicted that under a parental control model, cross-fostering would have no effect on food provisioning by the parents, but under chick control, the level of provisioning would be adjusted according to the age-specific requirements of the foster chick. Older chicks placed in the nests of younger chicks entered mass recession at an older age and had higher food-provisioning rates than unmanipulated chicks over the 10 days prior to fledging, supporting the parental control model. Younger chicks placed in the nests of older chicks had lower fledging masses than unmanipulated chicks, as a result of lower food-provisioning rates prior to fledging, which also supports this model. However, parents continued to feed younger chicks beyond the date at which their own chick would have fledged. This suggests that chicks also influenced food provisioning, and that mass recession resulted from interactions between a relatively fixed provisioning pattern in parents and age-specific food requirements of chicks.

Recent attention has focused on the foraging and food-provisioning strategies that allow procellariiform seabirds (e.g. albatrosses and shearwaters) to exploit pelagic food resources whilst providing food for dependent offspring at the nest. Several studies have found sexual differences in foraging and provisioning strategies in sexually dimorphic procellariiforms. However, there are few such data for species such as Manx shearwaters, that have negligible sexual size dimorphism. We used radiotracking coupled with periodic weighing of chicks to examine foraging trip durations and overall contributions to food provisioning by individual male and female Manx shearwaters at Skomer Island, U.K. Both sexes made trips of 1–4 days but females also made longer trips of up to 7 days. As a result, mean trip duration was significantly longer in females (2.1 days) than in males (1.5 days). Males fed their chick on a greater proportion of nights, delivered food to chicks at a higher rate, and made a greater contribution to overall food provisioning than females. Meal size increased with foraging trip duration but the yield to the chick in g/day was greater after short trips. Trips of 5–7 days would give females access to the highly productive nursery grounds for sardines, Sardina pilchardus, in the southeast Bay of Biscay, which were previously thought to be exploited by birds from Skomer but were more recently thought to be beyond the foraging range of chick-rearing adults, based on a mean trip duration of 2 days.

Arctic Terns (Sterna paradisaea) and Common Terns (S. hirundo) are similar in many aspects of their breeding ecology, but Common Terns generally lay three eggs per clutch whereas Arctic Terns lay two. In our study, Common Terns had a higher rate of food delivery and energy supply to the nest and higher nest attendance, indicating that they made trips of shorter average duration. This suggests that the number of chicks raised by these two species was primarily limited by the rate at which parents could supply food. However, estimated daily metabolizable energy intake of chicks was about 30% higher in Common Terns than in Arctic Terns. Common Tern chicks apparently spent a higher proportion of daily energy intake on maintenance of body temperature. It remains unknown whether this difference was because Common Tern parents could not brood three chicks as effectively as Arctic Terns brooded two or because the energy requirements for heat production in the third-hatched Common Tern chick were particularly high. If brooding did play a less important role in the energy budgets of Common Terns, the number of chicks that Arctic Terns could raise may have been limited not only by the rate at which parents could supply food to the nest but also by the requirements of chicks for brooding. We suggest that more detailed studies on the role of brooding constraints in limiting brood size in these species are required to clarify this matter.

The single young of shearwaters grows slowly and is fed at long, often highly variable, intervals. We monitored the growth patterns of nestling Short-tailed Shearwaters Puffinus tenuirostris exchanged between nests in order to assess whether the patterns of parental visits or the demands of their young determined the rate of food provisioning in this species. These cross-fostering trials indicate a strong parental influence upon the rate of food delivery, but did not exclude the possibility of a minor influence from chicks.

Although periodic weighing of chicks is used widely as a means to assess meal mass and feeding frequency in seabirds, there have been few critical assessments of its effectiveness. Using established methods, we were able to predict whether feeding had occurred with an accuracy of over 96% at nests where adult attendance was being monitored independently. This was despite the occasional provision of small meals which did not result in positive mass increments of chicks between weighings. However, information from repeat weighing could not be used to distinguish food delivery by both rather than a single parent within the weighing interval according to the relative sizes of mass increments. This may be because chicks are unable to cope with large meals from both adults because of gut capacity constraints. In addition, adults occasionally return from foraging trips with no food for the chick. These results have important implications for studies in which adult foraging trip durations are estimated solely using periodic weighing data, particularly in species with relatively short feeding intervals.

SUMMARYTrichomonas gallinaeis an emerging pathogen in wild birds, linked to recent declines in finch (Fringillidae) populations across Europe. Globally, the main hosts for this parasite are species of Columbidae (doves and pigeons); here we carry out the first investigation into the presence and incidence ofTrichomonasin four species of Columbidae in the UK, through live sampling of wild-caught birds and subsequent PCR. We report the first known UK cases ofTrichomonasinfection in 86% of European Turtle DovesStreptopelia turtursampled, along with 86% of Eurasian Collared DovesStreptopelia decaocto, 47% of WoodpigeonsColumba palumbusand 40% of Stock DovesColumba oenas. Birds were more likely to be infected if the farm provided supplementary food for gamebirds. We found three strains ofT. gallinaeand one strain clustering within theTrichomonas tenaxclade, not previously associated with avian hosts in the UK. OneT. gallinaestrain was identical at the ITS/5.8S/ITS2 ribosomal region to that responsible for the finch trichomonosis epizootic. We highlight the importance of increasing our knowledge of the diversity and ecological implications ofTrichomonasparasites in order further to understand the sub-clinical impacts of parasite infection.

The pattern of chick feeding in Little Shearwater Puffinus assimilis on Selvagem Grande was examined by weighing chicks at 4–h intervals throughout eight successive nights and daily for a further 11 days (19 days in all). Individual meals fed to chicks averaged 23.2 g (s.d. ±4.7) or 13.6% of adult mass. Mass increments over 24 h (NET) were linearly related to the sum of positive mass increments over 4–h intervals during the night (SUM) by the equation NET = 0.36SUM - 5.89 (r2= 0.60). Using this relationship, I estimated that over a period of 18 nights, a mean of 95% of chicks were fed each night, and the mean interval between feeds was 1.05 nights, with a maximum of three nights. There was no significant day-to-day variation in feeding rate. These results were not compatible with the prevalent idea that the purpose of large fat deposits in Procellariiformes is to tide chicks over periodic fasts resulting from poor feeding conditions. On average, chicks required 16 g of food per day to maintain constant mass and converted 33% of their intake of food above this requirement into biomass. Meal size and feeding frequency were independent of chick size and body condition (body-mass corrected for body-size), and the masses of food received by individual chicks each night varied in direct proportion to previous values. These results suggest that the rate of food supply to chicks was not regulated by adjustment according to chicks' nutritional requirements. To some extent, this supports the hypothesis that lipid accumulation among Procellariiformes is related to stochastic variation in food supply rate, resulting from an absence of regulation of feeding. However, feeding was not stochastic, in that adults tended to deliver consistent amounts of food to their chicks, and the pattern of feeding among even the worst-fed chicks was inconsistent with a need for large lipid stores based upon chance variation in food delivery.

This paper tests the predictions of parental investment theory and other hypotheses relating to variation in brood defence by examining aggression displayed by great skuas Catharacta skua towards intruders within their territories. Aggression serves the function of nest defence; hatching success of adults breeding in Shetland increased with aggression displayed during incubation, and the correlation between aggression and hatching success was apparent in three separate age classes. Adults displayed higher levels of aggression and greater parental investment in reproduction in years of poor food supply. This was not due to an increase in adults' expectations of future benefits of brood defence with increased investment. since hatching success was unaffected by food supply, and breeding success was lower in years of poor food supply. The observed increase in aggression therefore supports parental investment theory. Aggression increased with body condition index (in terms of mass corrected for body size) for females, but decreased with increasing body condition index for males. This probably reflects size-specific differences in the relative benefits of weight and manoeuvrability as means of promoting effective brood defence and reducing the risk of injury to parents. Aggression may also reflect adult quality, with body condition reflecting quality in opposite ways in males and females, as a result of their different roles during the breeding season.

SUMMARYSpecies of bird that use their wings for underwater propulsion are thought to face evolutionary trade-offs between flight and diving, leading to the prediction that species with different wing areas relative to body mass (i.e. different wing loadings) also differ in the relative importance of flight and diving activity during foraging trips. We tested this hypothesis for two similarly sized species of Alcidae (common guillemots and razorbills) by using bird-borne devices to examine three-dimensional foraging behaviour at a single colony. Guillemots have 30% higher wing loading than razorbills and, in keeping with this difference, razorbills spent twice as long in flight as a proportion of trip duration whereas guillemots spent twice as long in diving activity. Razorbills made a large number of short, relatively shallow dives and spent little time in the bottom phase of the dive whereas guillemots made fewer dives but frequently attained depths suggesting that they were near the seabed (ca. 35–70 m). The bottom phase of dives by guillemots was relatively long, indicating that they spent considerable time searching for and pursuing prey. Guillemots also spent a greater proportion of each dive bout underwater and had faster rates of descent, indicating that they were more adept at maximising time for pursuit and capture of prey. These differences in foraging behaviour may partly reflect guillemots feeding their chicks single large prey obtained near the bottom and razorbills feeding their chicks multiple prey from the water column. Nonetheless, our data support the notion that interspecific differences in wing loadings of auks reflect an evolutionary trade-off between aerial and underwater locomotion.

While the role of corticosterone in mediating the response of birds to acute stress is well established, it has recently been proposed that a decrease in prolactin levels following stress may complement corticosterone in redirecting resources away from breeding activities and towards behaviors promoting immediate survival. Here, for the first time, we detail changes in the prolactin stress response of birds throughout the breeding cycle. We then discuss the modulation of the corticosterone and prolactin stress responses over successive stages of breeding, differing in reproductive value and parental effort. In a long-lived Procellariiform seabird, the Manx shearwater Puffinus puffinus, we found that prolactin levels decreased in response to acute stress during incubation and mid chick-rearing but increased in response to stress during late chick-rearing and in non parenting birds, a pattern similar to that previously described for mammals. The high corticosterone stress response in pre-breeders was consistent with predictions based on reproductive value, but a similar response during late chick-rearing was not. This probably reflected foraging effort and a heightened importance of the parents’ own nutritional status at this stage of the season, in advance of post-breeding migration. We also found that baseline prolactin levels were maintained at high levels during chick-rearing and were only slightly lower during late chick-rearing and in failed breeders and non-breeders. These data suggest that prolactin may play a role in nestling care long beyond the brooding phase, that this is not due to birds spending long periods away from the colony and that prolactin secretion may be necessary for nest-guarding behavior.

Great skuas on Foula, Shetland have responded to a decline in the availability of sandeels since the late 1970s by increasing the proportion of other items in their diets. This change is correlated with the annual recruitment of sandeels in Shetland waters. Since 1983 there has been a 10-fold increase in predation by great skuas upon other seabirds, as Furness & Hislop (1981) suggested might occur in response to a low availability of sandeels. Changes in diet have been accompanied by a 50% reduction in adult territorial attendance as adults increased their foraging effort, such that between 1987 and 1989 breeding adults were probably working as hard as they were able to. Despite this, breeding success was less than 40% in 1987 and less than 15% in 1988 and 1989. The major cause of breeding failure was predation of unguarded chicks by adults from neighbouring territories. The willingness of adults to expose their chicks to high predation risk is probably maintained because of a positive correlation between chick pre-fledging growth and post-fledging survival, which is expressed up to the age of two years and which will place a strong pressure upon adults to feed their chicks as well as possible. The high expenditure of effort by adults in 1987 and 1988 did not affect the weights of those birds incubating eggs in 1988 and 1989, but there was a slight (3%) decrease in egg size between the late 1970s and the late 1980s. Changes in the age structure of the breeding population and the absence in 1989 of 28% of adults colour-ringed during incubation in 1988 suggest an increase in the rate of egress since the 1970s. These changes probably represent an increase in the long-term costs of reproduction to adults at this colony.

Procellariiform seabirds such as the Manx shearwater Puffinus puffinus, rear only one chick at a time but may breed many times in their lives; parents should thus limit food delivery to the chick in keeping with the balance between current and future reproductive output. Yet procellariiform chicks accumulate large quantities of lipid, which may provide a buffer against pronounced and unpredictable variation in food provisioning, resulting in part from an inability of parents to regulate food supply to the nest. We switched chicks between nests to examine the roles of parents and offspring in controlling food delivery. The serial autocorrelation in age-specific body masses for unmanipulated chicks decreased from 0.61 (P< 0.01) to 0.35 (NS) over a period of 15 days and remained nonsignificant thereafter. By contrast, the serial autocorrelation for switched chicks increased from 0.64 (P< 0.01) to 0.83 (P< 0.001) and the serial cross-correlation rose from 0.23 (NS) to 0.50 (P< 0.05). These results supported both chick determination and parental determination models of food provisioning, indicating that chicks conveyed information about their nutritional status, which parents acted upon by adjusting their rate of food delivery. We discuss these results in relation to the optimization of nestling lipid reserves and parental foraging effort. We suggest that information conveyed by the chick's begging intensity serves to reduce the provisioning rate to well-fed chicks, but parents cannot or do not increase food provisioning to poorly fed chicks. Such adjustment of food provisioning does not refute the hypothesis that nestling obesity provides a buffer against highly variable food delivery. Copyright 1999 The Association for the Study of Animal Behaviour.

In keeping with many species of waterbirds, there is little difference in the body size or external appearance of adult male and female Northern Gannets (Morus bassanus). Previous work has indicated that some morphological characteristics and behavioral patterns differ on average between sexes but there has yet to be a quantitative evaluation of the accuracy or applicability of these different criteria in determining sex. The aim of this study was to use DNA-based techniques to determine the sexes of adults and to use this information to evaluate a range of behavioral and morphological criteria for sex determination. Nape-biting behavior was frequently observed, was performed almost exclusively (>99% of observations) by males and was completely accurate in assigning sexes on the basis of multiple observations at each nest: this confirms previous results which indicated that nape-biting is highly sex-specific. Collection of nest material was also performed primarily by males, but this seldom occur...

Arctic Terns (Sterna paradisaea) and Common Terns (S. hirundo) are similar in many aspects of their breeding ecology, but Common Terns generally lay three eggs per clutch whereas Arctic Terns lay two. In our study, Common Terns had a higher rate of food delivery and energy supply to the nest and higher nest attendance, indicating that they made trips of shorter average duration. This suggests that the number of chicks raised by these two species was primarily limited by the rate at which parents could supply food. However, estimated daily metabolizable energy intake of chicks was about 30% higher in Common Terns than in Arctic Terns. Common Tern chicks apparently spent a higher proportion of daily energy intake on maintenance of body temperature. It remains unknown whether this difference was because Common Tern parents could not brood three chicks as effectively as Arctic Terns brooded two or because the energy requirements for heat production in the third-hatched Common Tern chick were particul...

Cory's Shearwaters, in common with other species of Procellariiformes, accumulate large quantities of body fat after hatching. This pattern of development has been related both to temporal variation in feeding conditions around the colony and to stochastic variation in foraging success and food provisioning by individual parents. Chicks need not necessarily always accept food provided by their parents, yet few studies have examined parental attendance independently of the masses of food fed to chicks overnight. This paper examines variability in provisioning and growth of nestling Cory's Shearwaters over three years between 1991 and 1994, and uses observations of individually marked adults coupled with radio-tracking to examine the nest attendance patterns of individual parents. Chicks were fed on 77% of nights on average, with little temporal variation in food delivery at the level of the colony. Individual chicks were nonetheless fed highly variably, and this to some extent supports the notion that nestling obesity results from chronic overfeeding by the parents to reduce the impact of variable food provisioning at an individual level. Chicks were attended by 1.5 parents overnight on average, and chicks were attended but not fed on 16% of occasions. Parents may have returned on these occasions without any food for the chick, or the chick may have refused to accept food from the parents. A minority of nests accounted for most of the occasions when a chick was attended but not fed, and there was evidence to suggest that these chicks were consistently heavy for their size. A refusal by these chicks to accept food on some occasions is consistent with the notion that lipid stores of chicks should be optimized rather than maximized, and this requires further investigation.

We used a supplementary feeding experiment to investigate the existence of a causal relationship between the nutritional status of nestling Puffins and the subsequent rates of food provisioning by their parents. Chicks given supplementary food received less frequent meals than controls, with the effect that the total amount of food received by the experimental group (parental delivery plus supplementary food) was similar to the amount delivered by parents in the control group. This indicated compensatory regulation of provisioning by parents in the experimental group. Increases in body size and body mass were virtually identical in the two groups until shortly before fledging, when supplemented chicks attained higher body mass than controls. This difference appeared to result mainly from the fact that unmanipulated chicks normally receive little food from their parents towards the end of the nestling period; supplements thus constituted a large proportion of chicks' normal daily food intake over this period, and compensatory reduction in food delivery by parents did not completely match the extra food given to chicks in the supplemented group. If growth rates of chicks were usually limited by poor food supply and adults delivered food to the chick at the maximum rate permitted by food availability irrespective of the chick's immediate nutritional requirements, then regulation of food delivery by parents would not be expected. The fact that adults were regulating food delivery to their offspring below the maximum attainable rate suggests that in this study at least, nestling growth rate was not limited by a scarce and unpredictable food supply.

We examined the pattern of chick feeding in Cory's shearwater Calonectris diomedea on Selvagem Grande by weighing 50 chicks at 4-h intervals throughout nine successive nights, then daily for a further 11 days (20 days in all). Meal sizes were estimated from the sum of positive mass increments recorded over 4-h intervals (SUM). Mass increments over 24-h (NET) were linearly related to SUMs by the equation NET = 0.83 SUM −40.1 (r 2 = 0.82). Individual meals fed to chicks averaged 74 g (SD = 26.4 g, range = 20 - 100 g). A mean of 79% of chicks were fed each night and the mean interval between feeds was 1.3 nights, with a maximum of four nights. There was little day-to-day variation in feeding rate, and no relationship with ambient light conditions over a half lunar cycle