The incidence of the tick parasite Ixodes uriae in Kittiwake Rissa tridactyla colonies in relation to the age of the colony, and a mechanism of infecting new colonies

Kittiwakes Rissa tridactyla (683 chicks and 64 adults) were examined for ectoparasites in three breeding areas: Isle of May (Firth of Forth, Scotland; eight colonies of approximately known age of occupation on natural sites), North Shields and Gateshead (Tyne and Wear, England; well established colonies of known age on buildings). All the 208 ticks collected were Ixodes uriae. On the Isle of May, a colony which had only been established for 7 years was already infested by ticks. Tick incidence increased significantly with the length of colonization of the area during the first 30 years. Age of colonization did not have an effect after 30 years. In Gateshead only a single adult among the 127 birds examined had a tick. No ticks were found on the 106 chicks born at North Shields checked in 1987 or in 1988. However, among nine visiting newly-fledged chicks from neighbouring colonies caught at the North Shields colony, six had either ticks or scars of previous infestation. Some visiting chicks remained for several days in the host colony and were even fed by the owners of visited nests. Ixodes uriae may thus spread from one colony to another during visits of newly-fledged infested chicks to neighbouring colonies. The absence of any infestation in old colonies (38 years in North Shields and 17 years in Gateshead) on buildings that are regularly visited by newly fledged infested chicks, suggests that buildings may lack the rock refuges necessary for tick survival from one year to the next. In this respect Kittiwake colonies on buildings are rather atypical.     

Costs and benefits of colony size vary during the breeding cycle in Black-headed Gulls Chroicocephalus ridibundus

We studied the effects of colony size on individual reproductive success in a multi-site population of Black-headed Gulls Chroicocephalus ridibundus where colony size ranged from 10 to 5,000 pairs. By focusing on family size, the number of chicks attended by individually marked parents, and accounting for between-individual variation, we detected a negative colony-size effect during the very first days of life of the chicks that was compensated by a subsequent increase in the proportion of surviving chicks with colony size. We suggest that this result originates in the interplay between overcrowding costs acting on hatching success, and benefits of colonial breeding, most probably more efficient food-searching (foraging enhancement), acting on chick survival. However, the frequency of complete colony failure increased with decreasing colony size. Taking this hazard risk into account yielded a corrected estimate of the effect of colony size on breeding success, and indicated that the largest colonies were the most productive. This pattern is congruent with the previous finding that larger colonies are more attractive to dispersing breeders.     

Recruitment to a seabird population depends on environmental factors and on population size

1. A novel capture-mark-recapture (CMR) method was used to build a multistate model of recruitment by young birds to a breeding population of common guillemots Uria aalge on the Isle of May, Scotland. Recruitment of a total of 2757 individually marked guillemots over 17 years was modelled as a process where individuals had to move from an unobservable state at sea, through a nonbreeding state present in the colony, to the breeding state. The probabilities of individuals returning to the colony in a given year, at age 2 and 3-4 years, were positively correlated with an environmental covariate, the winter North Atlantic Oscillation index (WNAO) in the previous years. 2. For 2 year olds, there was a negative relationship with breeding population size, suggesting that density dependence operated in this colony through limitation of food or some other resource. 3. Survival over the first 2 years of life varied with cohort, but was unrelated to the WNAO. Mean survival over this 2-year period was high at 0.576 (95% CI: 0.444; 0.708). 4. This high survival, combined with a low 'local' survival after age 5 years of 0.695 (0-654; 0.733) and observations of Isle of May chicks at other colonies, suggests that most surviving chicks return to the natal colony before deciding whether to recruit there or move elsewhere.     

The life cycle of the tick Amblyomma loculosum in a sooty tern Sterna fuscata colony in the Seychelles

An infestation of a sooty tern colony on Bird Island, Seychelles, with the ixodid tick Amblovomma loculosum was studied during visits to the colony in 1993 to 1995. During the infestation, adult ticks were numerous during the birds' incubation, larvae were abundant when young chicks were present, and there was some evidence that nymphs were found when older chicks were present. The life cycle of the tick appeared closely synchronized with that of its host. Following a breeding failure of the sooty terns, caused by adverse weather in August-September 1994, few adult ticks were found during the birds' breeding season of 1995, suggesting that the absence of chicks late in the 1994 breeding season led to starvation of the ticks and their failure to survive their diapause until the onset of the birds' next breeding season.     

Population trends in Kittiwake Rissa tridactyla colonies in relation to tick infestation

Parasite and disease transmission has long been considered one of the main costs of group living, but there is still little empirical evidence. Most seabird species breed in dense colonies and occupy the same breeding sites for many years and thus could be exposed to deleterious levels of haematophagous ectoparasite infestations. The aims of this study were to provide information on the epidemiology of the tick Ixodes uriae and to investigate the relationship between Kittiwake Rissa tridactyla colony population trends and local levels of tick infestation in chicks. It is based on a cross-sectional survey of a large sample of British Kittiwake colonies (22 colonies more than 30 years old and situated on natural cliffs). Ixodes uriae was by far the most common ectoparasite detected and was found in all colonies examined. Ticks were aggregated between nests, but not between siblings. Very young and old chicks were far less parasitized than chicks of intermediate age and were excluded from the analysis. Among colonies for which we had good data on population trends in Kittiwakes, those decreasing in number had a higher prevalence and density of ticks than those which were increasing. We suggest that this correlation reveals a cause-and-effect relationship. By influencing the local reproductive success, ectoparasites can affect the recruitment of new breeders and the breeders' site fidelity in the following year, thus influencing the local trend of numbers. This causal link highlights the potential effect of parasites on their host population and thus needs to be investigated further.     

Bone crushing carnivores and their significance to osteodystrophy in griffon vulture chicks

The natural removal of 89 ungulate carcasses by predators and scavengers was monitored in various wildlife reserves and ranching areas of South Africa. Carnivores responsible for the bone remains were determined. Spotted hyaenas and, to a lesser extent, Brown hyaenas, were the only carnivores that regularly chewed bones. When hyaenas were absent, months of weathering were required before the smaller bones became disarticulated and able to be removed by vultures. The bone-collecting behaviour and related aspects of breeding of two species of griffon vulture were studied at five different nesting colonies in southern Africa—one Cape vulture colony and one White-backed vulture colony in or near wildlife reserves, as well as two Cape vulture colonies and one White-backed vulture colony in ranching land. A total of 2825 bones was found in or below the vulture nests. These bones were categorized and measured. Hyaena-produced bone fragments were found only in the colonies in the wild areas—none of the 387 chicks examined here had osteodystrophy (metabolic bone disease). By contrast, in the ranching areas, vultures collected larger and less fragmented bones. Many Cape vulture chicks had osteodystrophy (130 of 1917 examined), as did two White-backed vulture chicks (of 196 examined). In 1977, artificial feeding stations, 'restaurants' where carcass skeletons were crushed, were established for Cape vultures. Since then, the incidence of osteodystrophy has declined from 17% to 2–5% in 1983. It is clear that bone fragments are an essential dietary requirement, providing calcium for correct skeletal growth of griffon vulture chicks.     

Birders of Africa: History of a Network

A total of 114 Marabou Stork chicks were ringed in the years 2003–2007 at the most southerly African breeding colony, Swaziland. Just one of 35 chicks fitted with colour rings was resighted by December 2007, while 17 of 79 chicks fitted with patagial tags were resighted. Juvenile Marabou Storks can disperse over 1 500 km within their first year after fledging, sufficient to cover the large distances between breeding colonies.     

Intraspecific variation in size and density of Avocet colonies: effects of nest-distances on hatching and breeding success

In many colonial bird species there is considerable intraspecific variation in colony size and inter‐nest distance (colony density). Possible causes of this variation and its effects on hatching success (survival of eggs) and breeding success (probability of a pair raising chicks) were studied in 48 Avocet Recurvirostra avosetta colonies in Schleswig‐Holstein (Germany) between 1991 and 1996. Colony density was influenced by time of year and habitat (categories: island or mainland, close to or far from feeding grounds). Colonies on islands had the highest densities. When all available space at a colony site was used, colonies became very dense (mean nearest‐neighbour nest distance less than 1 m). Colony size (number of clutches) was influenced by time of year, but not by habitat. Hatching success was low in high density colonies and in very low density ‘colonies’ (single nests) and high over a broad range of intermediate nest densities. The low success rate of single nests was caused by a very high predation rate, whereas the low success rate in very dense colonies was caused by a high rate of nest abandonment. Nest abandonment in very dense colonies was associated with a high level of aggressiveness among Avocets during the egg‐laying period. Due to territorial behaviour, Avocets seemed to be expelled from the densest breeding sites. In very dense colonies, high frequencies of clutches of unusual size occurred due to conspecific nest parasitism. The number of Avocets taking part in attacks on potential egg predators was small and (in colonies of more than one clutch) depended neither on colony size nor on colony density. Despite a low hatching success in very dense colonies, individuals breeding in the densest colonies had significantly better chances of raising chicks than Avocets breeding in less dense colonies. Coloniality seemed to be obligatory for Avocets in order to ensure hatching success. The size and density of colonies seemed to be associated with the availability of suitable nesting habitats (islands).     

Life cycle of the tick Ixodes uriae in penguin colonies: relationships with host breeding activity.

A survey of the temporal pattern of population structure and feeding activity of the seabird tick Ixodes uriae was conducted for the first time in two host species colonies: King penguin (Aptenodytes patagonicus halli) and Macaroni penguin (Eudyptes chrysolophus chrysolophus). The life cycle of the tick was investigated over 3 years in a King penguin colony and 2 years in a Macaroni penguin colony at Possession Island (Crozet Archipelago). There was a marked seasonal feeding activity pattern of ticks in both host species, connected with the presence of birds during the breeding season. Although the King penguin colonies were occupied throughout the year by birds, the favourable period for engorgement was limited to 3.5-4.5 months, and almost all the ticks overwintered in the unengorged state. Consequently, I. uriae probably completed its life cycle over 3 years in King penguin colonies. In contrast, this life cycle could be shortened to 2 years in Macaroni penguin colonies, as a result of a different timetable of the presence of birds for breeding and moulting. The relationships between such plasticity and the host behaviour and subantarctic climatic conditions are discussed.     

Activity periods and questing behavior of the seabird tick Ixodes uriae (Acari: Ixodidae) on Gull Island, Newfoundland: the role of puffin chicks

Questing behavior of Ixodes uriae and their associated seasonal, host-feeding patterns are crucial to our understanding of tick life history strategies and the ecology of diseases that they transmit. Consequently, we quantified questing behavior of nymphs and adult female I. uriae ticks at Gull Island, a seabird colony in Newfoundland, Canada, to examine seasonal variation of off-host and on-host tick activity. We sampled a total of 133 adult Atlantic puffins (Fratercula arctica), 152 puffin chicks, and 145 herring gull (Larus argentatus) chicks for ticks during the breeding seasons of 2004 and 2005. Questing ticks were sampled by dragging a white flannel cloth across the grassy breeding areas during the mo of May, June, July, and August. Nymph questing activity reached a peak during mid-July (79 and 110 individuals/hr in 2004 and 2005, respectively). The prevalence of nymphs and adult female ticks on different seabird hosts varied between years and during the seasons. Puffin chicks had the highest prevalence (above 70% in July) of nymphs in both years and this was correlated with questing activity. Female ticks rarely fed on puffin chicks, but were prevalent on adult puffins and gulls, although prevalence and questing of ticks were not correlated in these hosts. These patterns of off-host and on-host tick activity suggests that I. uriae ticks likely use a combination of questing and passive waiting, e.g., in puffin burrows, to detect hosts, depending on the tick stage and the host species.     

Hand-Rearing,Release and Survival of African Penguin Chicks Abandoned Before Independence by Moulting Parents

The African penguin Spheniscus demersus has an ‘Endangered’ conservation status and a decreasing population. Following abandonment, 841 African penguin chicks in 2006 and 481 in 2007 were admitted to SANCCOB (Southern African Foundation for the Conservation of Coastal Birds) for hand-rearing from colonies in the Western Cape, South Africa, after large numbers of breeding adults commenced moult with chicks still in the nest. Of those admitted, 91% and 73% respectively were released into the wild. There were veterinary concerns about avian malaria, airsacculitis and pneumonia, feather-loss and pododermatitis (bumblefoot). Post-release juvenile (0.32, s.e.  = 0.08) and adult (0.76, s.e.  = 0.10) survival rates were similar to African penguin chicks reared after oil spills and to recent survival rates recorded for naturally-reared birds. By December 2012, 12 birds had bred, six at their colony of origin, and the apparent recruitment rate was 0.11 (s.e.  = 0.03). Hand-rearing of abandoned penguin chicks is recommended as a conservation tool to limit mortality and to bolster the population at specific colonies. The feasibility of conservation translocations for the creation of new colonies for this species using hand-reared chicks warrants investigation. Any such programme would be predicated on adequate disease surveillance programmes established to minimise the risk of disease introduction to wild birds.     

Within- and between-year variation in the juvenile survival of Common Guillemots Uria aalge

We studied juvenile survival of 20 cohorts of Common Guillemot Uria aalge chicks colour-ringed on the Isle of May, Scotland, using both live observations at the colony and dead recoveries, allowing estimation of fidelity to the colony as well as survival. In this seabird, chicks leave the colony when only partly grown and are cared for by the male parent for several weeks afterwards. First-year survival varied strongly between cohorts, with a mean of 56% (range 30–91%). We did not identify any covariates which could explain this variation, whether relating to climate, population size or prey density. Survival was low during two regime shift episodes in the North Sea (1987–90 and 2000 onwards). Early hatched chicks were substantially more likely to survive than those hatching later in most years, whereas body condition at ringing had no detectable effect. Ringing recoveries indicated that mortality was highest in mid-winter, i.e. well after the cessation of paternal care. These results do not support the hypothesis that variation in prey quantity or energy content before fledging is a primary driver of variation in juvenile survival. Rather, it seems that chicks of high-quality parents are more likely to survive, as high-quality females tend to lay earlier in the season, and high-quality males presumably are better able to prepare their chicks to survive their first winter at sea. Very few (4%) Guillemots emigrated permanently before age 3 years, but from age 5 onwards 25–30% of birds annually left the colony or otherwise became unobservable.     

Impacts of human visitors on breeding success and long-term population trends in Adélie Penguins at Casey,Antarctica

Breeding populations of Aélie Penguins Pygoscelis adeliae have been counted at two localities near Casey Station in Wilkes Land, East Antarctica since 1959/60 and 1968/69. At Whitney Point, the breeding population increased from 1122 pairs in 14 colonies in 1959/60 to 4714 pairs in 36 colonies in 1992/93. All new colonies at Whitney Point established on relict colony sites identified in 1959/60. On Shirley Island, the total breeding population has remained at 7770 pairs ±10% between 1968/69 and 1992/93, except in 1990/91 when the population peaked at 8719 pairs. An association between the age of a colony and its rate of increase was observed at Whitney Point, with new colonies (those established since 1971/72) increasing more rapidly than colonies extant in 1959/60. At Shirley Island, where most of the colonies extant in 1968/69 have decreased in population, the establishment and growth of 13 colonies has offset this decrease; these new colonies also exhibited the association between age and rate of increase. Breeding success (chicks fledged per nest) was significantly lower for Shirley Island colonies than for those at Whitney Point. Human visitors to Shirley Island from Casey station are believed to be responsible for the observed changes in the distribution and abundance of breeding pairs and for maintaining the stable population by reducing overall breeding success through the disturbance associated with visits.     

Consequences of foraging trip duration on provisioning behaviour and fledging condition of common murres Uria aalgae

We examine the provisioning constraints of a pursuit‐diving seabird in a cold ocean regime by comparing the behaviour of common murres Uria aalge rearing chicks at two colonies in the Northwest Atlantic during 1998‐2000. Funk Island is the largest (340,000–400,000 breeding pairs) and most offshore (60 km) colony of common murres in eastern Canada. Seventy‐five percent of the Northwest Atlantic population of common murres breeds on this island. Great Island is one island within the Witless Bay Ecological Reserve, which is the second largest breeding aggregation (100 000 breeding pairs) and is located near‐shore (2 km). The primary forage fish species in Newfoundland waters is capelin Mallotus villosus, which spawns on or near coastal beaches during summer. Therefore, the two study colonies differ in their distance to food resources and colony size. It is within this natural context that we compare: (1) prey types and frequency of delivery (amount of prey), (2) parental time budgets, and (3) the mass and condition (mass/wing length) of fledglings at both colonies. Similarly sized female capelin (100–150 mm) were delivered to chicks at both colonies. Foraging time per day per parent, a proxy of foraging effort, was similar at both colonies (Great Island: 5.1 h; Funk Island: 5.5 h), as was the percentage of time spent with mates (Great Island: 12.3%; Funk Island: 10.9%). Foraging trips, however, were longer at Funk Island (4.1 h) than at Great Island (2.9 h). This resulted in lower feeding rates of chicks (0.17 feeds per h) and poorer condition of fledglings (2.9 g/mm) at Funk Island compared to those at Great Island (0.22 feeds per h; 3.9 g/mm). We hypothesize that provisioning efforts are constrained at Funk Island by (1) distant food resources and increased competitor density, resulting in longer foraging trip durations and (2) the time spent paired with mates at the colony, which may reflect a minimum time required to maintain breeding sites due to higher breeding densities at Funk Island compared to Great Island. Demographic consequences of this poor fledgling condition at Funk Island are unknown, but fledglings may sufficiently accelerate growth at sea due to their closer proximity to an important nursery area. If fledgling survival is compromised, however, the lower potential for growth at Funk Island will impact the entire Northwest Atlantic population of common murres.     

Does foraging efficiency vary with colony size in the fairy martin Petrochelidon ariel?

Colonial breeding occurs in a wide range of taxa, however the advantages promoting its evolution and maintenance remain poorly understood. In many avian species, breeding colonies vary by several orders of magnitude and one approach to investigating the evolution of coloniality has been to examine how potential costs and benefits vary with colony size. Several hypotheses predict that foraging efficiency may improve with colony size, through benefits associated with social foraging and information exchange. However, it is argued that competition for limited food resources will also increase with colony size, potentially reducing foraging success. Here we use a number of measures (brood feeding rates, chick condition and survival, and adult condition) to estimate foraging efficiency in the fairy martin Petrochelidon ariel, across a range of colony sizes in a single season (17 colonies, size range 28–139 pairs). Brood provisioning rates were collected from multiple colonies simultaneously using an electronic monitoring system, controlling for temporal variation in environmental conditions. Provisioning rate was correlated with nestling condition, though we found no clear relationship between provisioning rate and colony size for either male or female parents. However, chicks were generally in worse condition and broods more likely to fail or experience partial loss in larger colonies. Moreover, the average condition of adults declined with colony size. Overall, these findings suggest that foraging efficiency declines with colony size in fairy martins, supporting the increased competition hypothesis. However, other factors, such as an increased ectoparasitise load in large colonies or change in the composition of phenotypes with colony size may have also contributed to these patterns.     

Philopatry and colony fidelity of Shags Phalacrocorax aristotelis on the east coast of Britain

Shag Phalacrocorax aristotelis colonies from the Firth of Forth south to the Fame Islands (SE Scotland and NE England) were surveyed in 1982 and 1983 for nesting Shags ringed as chicks or adults in the previous 20 years. On average, 5% of 863 Shags ringed as chicks were breeding away from their natal colony, and only 1% of 401 marked adults had left their breeding colony. The proportion p of chicks that settled at or beyond distance D from the natal colony followed the relationship p = 0.263D-^0.771, implying that more Shags moved 10 km or beyond to breed than expected from diffusion models or a constant-rate dispersal model; adjusted for birds that settled outside the study area, the relationship was p = 0.314D^-0771. There was no effect of colony size on emigration rate, but females showed a greater tendency to settle away from the natal colony than did males. At the natal colony, more males (69%) than females (47%) nested within 300 m of their birthplace. Two mechanisms appear to determine where a Shag settles to breed: a navigational one governing the return to the natal colony, plus the competitive process of nest-site establishment.     

Studying the effects of multiple invasive mammals on Cory��s shearwater nest survival

The most common invasive mammals??mice, rats, and cats??have been introduced to islands around the world, where they continue to negatively affect native biodiversity. The eradication of those invasive mammals has had positive effects on many species of seabirds. However, the removal of one invasive mammal species may result in abundance changes of other species due to trophic and competitive interactions among species. Understanding the overall impact of several invasive species is a key challenge when evaluating the possible effects of eradication programmes. Here we assess the influence of the three most common invasive mammals on nest survival of Cory??s shearwater (Calonectris diomedea). We monitored six breeding colonies over 3?years and measured the activity of mice, rats and cats to examine the influence of invasive mammals on nest survival. We found that nest survival showed a similar temporal trend in all years, with lowest weekly survival probabilities shortly after chicks hatched. Cats were identified as major predators of chicks, but no measure of colony-specific cat activity was able to adequately explain variation in shearwater nest survival. Nest survival was on average 0.38 (95?% confidence interval 0.20?C0.53) and varied among colonies as well as over time. We found a small positive influence of rats on nest survival, which may indicate that the presence of small rodents as alternative prey may reduce cat predation of chicks. Our findings suggest that the eradication of rodents alone may exacerbate the adverse effects of cats on shearwater nest survival.     

Disturbance-dependent Yellow-legged Gull (Larus michahellis) predation on Larid chicks decreases with chick age

Predation is one of the key factors shaping the dynamics of animal populations. In birds, nest loss due to predation can be a significant cause of low reproductive success. Ground-nesting birds are among the bird groups most susceptible to predation, mainly because their nests are easily accessible to a broad suite of potential predators. For these birds, anthropogenic disturbances can generate changes in nest predation risk by altering their antipredator behaviour and also by altering the behaviour of the predator species, i.e. the predator becoming much more aware of predation opportunities due to frequent disturbances and/or motivated to repeat predation attempts when some are successful. To date, most previous studies investigating this have focused on a single effect, either predation or disturbance, on chick survival. It remains unknown how the risk of predation with and without disturbance varies with chick age. In this study, we used behavioural observations to assess how the interaction between predators and disturbance affects predation risk in chicks and how this interacts with chick age. Specifically, we investigated the effect of disturbance caused by humans and stray dogs on the predation of Slender-billed Gull Chroicocephalus genei chicks by Yellow-legged Gulls Larus michahellis, and whether this depended on the age of the chicks. Our results revealed that disturbance had a significant positive effect on predation measures of Slender-billed Gull chicks by Yellow-legged Gulls, but that this effect was mediated both by disturbance type and the age of chicks. Stray dogs entering the colony had a stronger disturbance effect on chicks than passing humans, increasing predation risk by Yellow-legged Gulls. Our results also showed that chick age interacts with disturbance type to determine the predation risk. This is probably mediated by chicks' capacity to escape predation by gathering in a single large crèche that runs into the water when disturbed. To preserve Slender-billed Gull colonies in one of its few remaining breeding sites in Tunisia, and as gulls tend to react even when the disturbance occurs relatively far from the colonies, it is crucial to (1) restrict human access to dikes and islets where large colonies breed and (2) construct artificial islets attractive to gulls and inaccessible to stray dogs.     

Foraging rhythms in Adélie penguins (Pygoscelis adeliae) at hope bay,Antarctica; determination and control

Summary We studied diel periodicity in activity of Adélie penguins (Pygoscelis adeliae) during the austral summer solstice and one month later by counting, hourly, numbers of birds leaving colonies to forage, numbers arriving back after foraging, numbers in a colony and numbers resting outside the colonies. During the solstice large numbers of birds were arriving at and departing from the colony at all times of the day although there was a tendency for more birds to be at sea when light intensity was highest at mid-day. Generally, birds not brooding chicks did not rest on land. A month later, when visibility was poor at mid-night, the percentage of birds at sea was highly positively correlated with light intensity. Birds returning from foraging in the evening fed chicks immediately and then either rested in the colonies or on snow patches between the colonies and the sea until the following morning.     

Brood reduction in temperate and sub-tropical ospreys

Summary In an effort to understand patterns and causes of nestling loss in Ospreys (Pandion haliaetus), I studied brood reduction in three eastern U.S. Osprey colonies during 1978 and 1979. The colonies, located in Florida Bay (1) and on coastal Long Island, N.Y. (2), differed in the average daily amount of food delivered to nestlings; Florida nests received 43% and 11% less fish per day than nests in the two N.Y. colonies, largely because latitude and season restricted day length and thus foraging time for the winter-breeding Florida Ospreys. Increased distance from stable food sources accounted for the lower rate of feeding at one of the N.Y. colonies. Variation in clutch size in the three colonies reflected differences in latitude more than in food availability; average clutch sizes in Long Island were larger than Florida clutches by 0.5 of an egg, but were similar to each other and to those in other northeastern U.S. Osprey populations.Increased nestling loss coincided with reduced food delivery rates and, in food stressed colonies, this loss was 2–3 times greater than any recorded for Ospreys. Starvation was the primary cause of nestling death, with mortality concentrated on third chicks, which hatched on average 3.9 d later and from eggs 5.6% smaller than chicks hatching first. Sibling aggression accounted for the preferential feeding of older nestmates,but only in colonies or nests where food was limited. Aggressive chicks nearly always stopped fighting after being fed. This behavior provided a reversible mechanism for controling brood reduction that was based on nutrition. Growth rates of young measured during the first half of the growth period were more variable between colonies than within nests. This is interpreted as reflecting both the differences in colony food delivery rates as well as the evolutionary pressures of sibling competition to equalize the growth of nestmates.