The behaviour of seabirds foraging at fishing boats around Shetland

Among the different types of fishing vessels around Shetland, whitefish trawlers attract the largest numbers of scavenging seabirds and provide the most food. Offal was almost all consumed by seabirds, predominantly by Fulmars Fulmarus glacialis , which excluded other species by their aggression. Fulmars generally ignored discarded whole fish, which were mainly taken by Great Black-backed Gulls Larus marinus , Gannets Sula bassana and Great Skuas Catharacta skua . Although flatfish were usually ignored because seabirds found them difficult to swallow and they sank faster, most discarded roundfish were consumed. Herring Gulls L. argentatus , Lesser Black-backed Gulls L. fuscus and Kittiwakes Rissa tridactyla were rarely able to obtain offal or discards. Herring Gulls and Lesser Black-backed Gulls spent much time on the periphery of feeding flocks while Kittiwakes rarely attempted even to join these. Most of the birds at trawlers were in adult plumage, and it is suggested that the low proportion of immature birds present was a further reflection of the highly competitive feeding conditions at trawlers. We suggest that likely changes in fishing practice and seabird population sizes in the immediate future may result in Herring Gulls, Lesser Black-backed Gulls and Great Skuas finding feeding on waste around trawlers increasingly difficult, so they may be further displaced by Fulmars, Gannets and Great Black-backed Gulls.     

THE SIGNIFICANCE OF KLEPTOPARASITISM DURING THE BREEDING SEASON IN A COLONY OF ARCTIC SKUAS STERCORARIUS PARASITICUS IN ICELAND

Breeding and kleptoparasitism of Arctic Skuas was studied at Vik, Iceland, in 1973. Hatching success was 88.9%; fledging success was 0.27 fledglings per pair. The heavy chick mortality occurred mostly in the first week after hatching. In the early part of the breeding season skuas fed by robbing kittiwakes of their food at sea and by eating arthropods at the breeding grounds. At the time of hatching of skua eggs, which coincided with the hatching of Puffin eggs, skuas switched their feeding activities to kleptoparasitism of Puffins and fish so obtained was the principal item of most skuas' diet thereafter. In the first week post-hatching, the energy balance of an average adult skua pirating Puffins was estimated to be negative, but changed to positive a week later, although later, coinciding with an influx of non-breeders it turned back to negative. Arctic Terns which normally breed at Vik, and are exploited by skuas, failed to breed in 1973, and the abnormally heavy chick loss is therefore attributed to this failure of the terns. The success of the skuas kleptoparasitizing Puffins depends on the proximity of the interactions to the cliff or to the ocean, where Puffins seck refuge. Skuas catch fish mostly in the air, especially if it is dropped from high. Fish dropped from low is mostly taken by competing gulls and Ravens, which mostly control the ground and lower airspace. Skuas chasing in groups enhance their success, but the average success per member decreased with group increase. However, the success of the group-member in the ‘best’ position was equal to that of a single skua. Puffins carry 1 (large)-6 (smaller) fish, an approximately equal load irrespective of number of fish. Skuas preferentially chase Puffins carrying ‘large’ fish, thus maximizing their yield per effort. Arctic Skuas responded to changes in the numbers of arriving Puffins with a functional response, but their monitoring of the food supply was far from perfect.     

Predation risk and moonlight avoidance in nocturnal seabirds

Unlike most seabird families, the vast majority of small petrel species are nocturnal on their breeding grounds. Further, they reduce markedly their activity when the light level increases. Moonlight avoidance might be a consequence of reduction in foraging profitability, as bioluminescent prey do not come to the sea surface on bright nights. Alternatively, petrels may avoid colonies during moonlit nights because of increased predation risk. We studied predation on petrels by Brown Skuas Catharacta antarctica lönnbergi at Kerguelen, and the influence of moonlight on behaviour of both skuas and petrels, to test the 'predation risk' hypothesis. On the study area, Brown Skuas hunt at night and prey heavily upon the Blue Petrel Halobaena caerulea and the Thin-billed Prion Pachyptila belcheri . Predation risk was higher on moonlit nights, as skuas caught more prey, and particularly more Blue Petrels when the light level increased. Nightly intakes of Blue Petrel and Thin-billed Prion by skuas was related to colony attendance of non-breeders rather than that of breeders. Biometry of prey also suggested that skuas caught a higher proportion of non-breeding birds than was present at the colonies. Predation risk was thus greater in non-breeders and on moonlit nights. Colony attendance by non-breeding Blue Petrels and Thin-billed Prions was also reduced during moonlit nights. Vocal activity, which is mainly by non-breeders, was also drastically reduced when the light level increased in the species suffering the highest predation rate. Our results supported the 'predation risk' hypothesis, although the 'foraging efficiency' and the 'predation risk' hypotheses are not mutually exclusive: the former might explain the moonlight avoidance behaviour of breeding, and the latter that of non-breeding individuals.     

Seabird predation by great skuas Stercorarius skua– intra‐specific competition for food?

Competition for food is widely cited as an important cost of coloniality among birds and much of the evidence in support of this hypothesis comes from studies of colonial piscivorous seabirds. However, for generalist seabirds able to switch between different prey types, the role of food availability in relation to colony size is unclear. Here we investigate patterns of the consumption of seabird prey in relation to colony size in a generalist seabird, the great skua Stercorarius skua, in Shetland, UK. At the population level skuas feed mainly on sandeels Ammodytes marinus and fishery discards, but respond to declines in fish availability to facultatively prey on other seabirds. By comparing the consumption of seabirds among seven different sized colonies, including one colony with artificially reduced numbers of skuas (Fair Isle), we investigate whether consumption of seabird prey is influenced by skua population size, while simultaneously measuring seabird prey availability. Data from five years also enables us to investigate the influence of annual variation in environmental conditions on seabird consumption. Using measures of body condition and reproductive performance we investigate the consequences of living in different sized colonies, which may provide insight into ultimate costs of nesting at high population density. Skua diets varied among colonies and the proportion of seabird prey in the diet was inversely related to skua colony size, despite similar per capita numbers of seabirds across colonies. At the colony where their numbers were artificially suppressed, skuas consumed a greater proportion of seabirds per capita. Highly significant year effects in seabird predation were observed but the pattern among colonies remained consistent over time. Two measures of adult body condition (pectoral muscle index and mean corpuscular volume) revealed that adult great skuas were in poorer condition at the largest colony (Foula), but reproductive performance did not alter significantly among colonies. This study provides evidence that intra‐specific competition among skuas may limit opportunities for obtaining seabird prey, which may be particularly important during periods of poor availability of sandeels and fishery discards, and has implications for assessing the impact of skuas on seabird populations.     

Evidence for fatal collisions and kleptoparasitism while plunge‐diving in Gannets

Plunge‐diving is a highly successful strategy for dealing with the challenges confronting birds feeding on pelagic prey. We tested for evidence of fatal injuries due to collision between conspecifics in plunge‐diving Australasian Gannets Morus serrator and Cape Gannets Morus capensis, respectively, by performing post‐mortem examinations of carcasses recovered from New Zealand waters and analysing video footage of Cape Gannet foraging events from South Africa. We found evidence of accidental collisions between Gannets and also observed a case of attempted kleptoparasitism, in which a diving Cape Gannet targeted a previously captured fish in the beak of a conspecific.     

Kleptoparasitic interactions of Arctic Skuas Stercorarius parasiticus and Black Guillemots Cepphus grylle in northeastern Hudson Bay, Canada

Kleptoparasitic behaviour of Arctic Skuas was studied at a Black Guillemot colony in northeastern Hudson Bay, Canada. Skuas procured fish in 7-2% and 19-9% of attacks, and Black Guillemots lost fish in 7-2% and 21-2% of chases in 1982 and 1983, respectively. In 1983 chase outcome was related to distance between birds at the end of a chase and number of skuas in pursuit. The skuas' foraging efficiency (energy intake/energy expended during hunting) was estimated at 45 for 1982 and 5–8 for 1983, and total daily energy expenditure was estimated at 2–4 and 2-1 × b.m.r. for these years. Energetic calculations suggested that skuas obtained sufficient food to meet their daily needs in 3–6 h of hunting in 1982 and 2–7 h in 1983. The absence of intensive kleptoparasitic activity by breeding skuas in North America is attributed to the paucity of colonies suitable for both feeding (due to available host species and colony topography) and nesting (due to presence of arctic foxes and/or polar bears).     

Predator‐avoidance behaviour in a nocturnal petrel exposed to a novel predator

Many species of bird recognize acoustic and visual cues given by their predators and have complex defence adaptations to reduce predation risk. Recognition of threats posed by specific predators and specialized anti‐predation behaviours are common. In this study we investigated predator recognition and anti‐predation behaviours in a pelagic seabird, Leach's Storm‐petrel Oceanodroma leucorhoa, at a site where predation risk from Great Skuas Stercorarius skua is exceptionally high. Leach's Storm‐petrels breed in burrows and come on land only at night. Counter‐predator adaptations were investigated correlatively in relation to changing natural light levels at night, and experimentally in relation to nocturnal visual and acoustic signals from Great Skuas. Colony attendance by Leach's Storm‐petrels was attuned to changes in light conditions at night and was highest when nights were darkest. This behaviour is likely to reduce predation risk on land; however, specific recognition of Great Skuas and specialized defence behaviours were not found. Leach's Storm‐petrels, in particular apparently non‐breeding individuals, were entirely naïve to the threat posed by Great Skuas and were captured easily in a variety of different ways, on the ground and in the air. Lack of specialized behavioural adaptations in Leach's Storm‐petrels against Great Skuas may be because spatial overlap of breeding distributions of these species appears to be a rare and recent phenomenon.     

PREY SELECTION AND TEMPORAL VARIATION IN THE DIET OF SUBANTARCTIC SKUAS AT INACCESSIBLE ISLAND,TRISTAN DA CUNHA

Ryan, P.G., & Moloney, C.L. 1991. Prey selection and temporal variation in the diet of Subantarctic Skuas at Inaccessible Island, Tristan da Cunha. Ostrich 62:52-58.

More than 2 500 prey items of Subantarctic Skuas Cuthuructa anturcricu were identified from regurgitated pellets collected at a roost of non-breeding skuas during summer 1989–90 at West Point, Inaccessible'Is-land. Most prey items (96,5%) were birds, primarily burrow-nesting procellariiforms. Sixteen bird species were recorfed in the diet, but five secies accounted for 94,6% of identifiable prey remains: Whitebellied Storm Petrel Fieettu grullariu (53,5%), Whitefaced Storm Petrel Pelugodrorna marina 15,1%), Broad-billed Prion Pacaptilu vittutu (14,0%), Great Shearwater Puffinus gruvis (7,3%), and Iommon Diving Petrel Pelecunoids urinutrix (4,7%). Petrel chicks were important in the diet, particularly Great Shearwaters. The main non-avian prey were fish and goose barnacles Lepas sp., but their importance may have been underestimated. The remains of a rat Rattus sp. presumably were derived from the neighbouring island of Tristan da Cunha; rats are not known to occur on Inaccessible Island. Prey size affected seabird availability to skuas, and dietary composition vaned throughout the summer in relation to seabird breeding cycles. Both these factors reduce the value of skua diet as an indicator of the relative abundances of burrow-nesting petrels. There was no correlation between skua diet and estimates of breeding bird abundance, but this may be a consequence of studying non-breeding skuas. Subantarctic Skua diet in 1989–90 was similar to that recorded in previous years, with consistent seasonal trends between years.     

Atlantische Papagei‐taucher in Not

Puffins, climate change and fisheries Long term changes in north Atlantic weather leading to climate changes have measurable influences on marine eco systems. Seabirds have to cope with changes in the food web, driven by rising SST and other abiotic factors. The Atlantic Puffin is a good example for this. We can study its breeding success and colony size as measurements for changes in the food resources, mainly sandeels or herring over a wide area from the southern boreal to the arctic waters and from the east to west coast colonies of the Atlantic ocean. The findings show that population changes and breeding success in several seabird species like the Puffin and Common Guillemots or Black‐legged Kittiwakes as well show strong correlations to declining sandeels as a consequence of intensive industrial fishing, changes in SST and plankton, especially the most abundant copepod Calanus finmarchicus, development.     

Interactions with Great Skuas Stercorarius skua as a factor in the long‐term decline of an Arctic Skua Stercorarius parasiticus population

Many of the UK’s seabird species have displayed high variation in breeding success since the 1980s, largely due to changes in the availability of Lesser Sandeels Ammodytes marinus, their main prey. During this time, Arctic Skuas Stercorarius parasiticus experienced a rapid decline in the UK and the species has subsequently been placed on the Red List of birds of conservation concern. Although shortage of Lesser Sandeels is likely to be an influential factor, the Arctic Skua’s breeding range overlaps with that of the Great Skua Stercorarius skua, a larger bird with a more varied diet, and interspecific interactions for nesting habitat may exert an additional pressure on Arctic Skua breeding populations. Results from four censuses, spanning 21 years, were used to model habitat use and analyse distributional change in nesting Arctic Skuas at a major colony located on Fetlar, Shetland, Scotland. The decline in Arctic Skuas was not uniform across the island and competition with Great Skuas for nest‐sites appears to have influenced localized breeding distribution. By 2006, Arctic Skuas had been almost entirely excluded from shrub heath, blanket bog and coastal heath habitats, which were identified as preferred habitat in 1986. In 2006, Arctic Skua breeding territories were mainly restricted to one core area of preferred habitat where over 90% nested in high density as this habitat became increasingly occupied by Great Skuas. The more generalist foraging habit of the Great Skua allowed the population to grow rapidly as numbers of the more specialist Arctic Skua decreased during times of low sandeel availability. Our model suggests that both interspecific competition for territories with Great Skuas and food limitation have played important roles in the decline of Arctic Skuas on Fetlar.     

The effect of food availability on the foraging behaviour of breeding Great Skuas Catharacta skua and Arctic Skuas Stercorarius parasiticus

Seabirds' behavioural activity budgets have been proposed as an indicator of changes in marine food supplies. Reduced recruitment and a low biomass of sandeels Ammodytes spp. around Shetland in the late 1980s were associated with a dramatic increase in the foraging effort of breeding skuas. We present data on the territorial attendance patterns of breeding Great Skuas Catharacta skua and Arctic Skuas Stercorarius parasiticus in Shetland in 1987 and examine three aspects of the recorded increase in foraging effort.
Time budget studies indicate that recording territorial attendance by infrequent, daily spot observations provides an accurate measure of the time spent off-territory by both skua species. None of the pairs of Arctic Skua studied ever left their chicks unattended in 1987. The same was true of 10 out of 27 pairs of Great Skuas. Although the remaining pairs of Great Skuas frequently left their chicks unprotected, evidence of pair co-ordination to minimise this suggests that their foraging schedules were still constrained by the need to protect their offspring. During periods of high chick-demand, variation between pairs' foraging efforts largely reflected variation in those of the females. Thus, we suggest that increased chick neglect in years of food shortage arises as females abandon their role of guardian for that of provider. By the end of the breeding season in 1987, both male and female Great Skuas foraged from 65 to 89% of daylight hours. The lack of any further increase in the foraging effort of Great Skuas in subsequent years of poorer reproductive success was almost certainly because time and energy constraints prevented either sex from doing so. We suggest that the comparatively short amount of time spent foraging by Arctic Skuas in 1987 may reflect an advantage conferred by their role as specialist kleptoparasites.     

Summer diet of seabirds feeding in sea-ice-covered waters near Svalbard

Summary We describe the summer diets of four seabird species. Kittiwake (Rissa tridactyla), Little Auk (Alle alle), Black Guillemot (Cepphus grylle) and Brünnich's Guillemot (Uria lomvia) collected in sea-ice-covered waters near Svalbard. Birds collected in an area filled with young sea-ice, within the seasonal sea-ice zone, were compared with birds collected from the perennial sea-ice zone dominated by multiyear ice. Pelagic Crustacea and fish dominated the diet of birds feeding in young ice, while sympagic Crustacea and fish were most important in the diet of birds feeding in multiyear ice. Boreogadus saida was the most important fish food item. B. Saida was present in the ice in both areas, while sympagic Crustacea were lacking in the area filled with young ice. Important food items in young ice were B. saida (Black Guillemots and Kittiwakes), Cajanus sp. (Little Auks) and Pandalus borealis (Brünnich's Guillemot). B. saida (Black Guillemots, Kittiwakes, Brünnich's Guillemots), Gammarus wilkitzkii (Brünnich's Guillemot, Black Guillemot) and Apherusa glacialis (Little Auk) were most important in multiyear ice. In general, Black Guillemots and Kittiwakes fed on fish, whereas Brünnich's Guillemots fed on the larger, and Little Auks on the smaller, Crustacea. The importance of sympagic species in the diet of seabirds is thought to be closely related to the age and history of the ice in the feeding area.     

Skuas on the Eurasian tundra; relative occurrence of species, ages and colour phases

The breeding ranges of the three closely related skua species in the genus Stercorarius are highly sympatric on the Arctic tundra. During the Swedish-Russian Tundra Ecology Expedition, 1994, the ages and colour phases of Pomarine Skuas Stercorarius pomarinus , Arctic Skuas Stercorarius parasiticus and Long-tailed Skuas Stercorarius longicaudus were recorded at 15 sites along a transect across 140̀ of longitude from the Kola Peninsula in the west to Wrangel Island in the east. An index of the lemming numbers was also measured. Pomarine Skua comprised 52% of the 1587 skuas seen, with 38% Long-tailed Skua and 10% Arctic Skua. The Arctic Skua occurred at low densities all along the transect. The Pomarine Skua was most common at the northernmost sites in this lemming year. It was absent from all sites with low rodent densities but also from two more southerly sites with high lemming numbers. The Long-tailed Skua showed a distribution sympatric with that of the Pomarine Skua but occurred at lower densities, and it also bred at the more southerly sites. No skuas in second-year plumage were observed. Older subadult skuas were observed in increasing numbers with age. The proportion of subadults was markedly higher in the Pomarine Skua (10%) compared with the two other species. The proportion of dark Pomarine Skuas was about 8% all along the transect. In the Arctic Skua, there was an extremely marked shift from 64% dark birds on the Kola Peninsula to an almost complete dominance by the light phase in the rest of Arctic Russia. I suggest that dark skuas are more efficient kleptoparasites over the sea while the light phase is at an advantage hunting over the tundra.     

Food and Feeding Ecology of Puffins

This study shows that although annual fluctuations occur in the main prey types of Skomer Puffins, food availability does not appear to limit breeding success. Most birds found food close to the colony, showing peaks of activity early in the day and again in late afternoon; and two out of four experimental pairs were able to rear 'twins', though the growth-rates of these were less than for single chicks. Some interesting observations are given on the feeding of the chick and on kleptoparasitism of adults by Jackdaws and gulls.     

Evidence that tufted puffins Fratercula cirrhata use colony overflights to reduce kleptoparasitism risk

Predation, foraging and mating costs are critical factors shaping life histories. Among colonial seabirds, colony overflights may enhance foraging or mating success, or diminish the risk of predation and kleptoparasitism. The latter possibility is difficult to test because low predation or kleptoparasitism rates could be due either to low danger or to effective counter-tactics by prey. Tufted puffins Fratercula cirrhata breeding at a large colony in British Columbia, Canada, deliver several loads of fish each day to their nestlings and are targets for kleptoparasitism by glaucous-winged gulls Larus glaucescens . In the present study, we documented the ecological conditions under which overflights occurred in order to assess when overflights were made and to statistically isolate the effect of overflights on kleptoparasitism risk at this site. Load-carrying puffins engaged in overflights under ecological conditions associated with relatively high rates of kleptoparasitism. Further, when ecological factors determining risk were statistically controlled, overflights were correlated with marginally lower chances of kleptoparasitism than when the risk factors were ignored. The results suggest that breeding puffins at this site use overflights for kleptoparasite avoidance. This tactic is used sparingly, suggesting it is costly. Costs of overflight behaviour might contribute to the impact of kleptoparasitism on the breeding success of tufted puffins.     

THE KLEPTOPARASITIC BEHAVIOUR OF THE ARCTIC SKUA STERCORARIUS PARASITICUS WITH THREE SPECIES OF TERN

The behaviour of Arctic Skuas when stealing fish from three species of tern was studied at the estuary of the River Ythan, Aberdeenshire. The skuas' attacks were made by flying low over the water at high speed and by approaching the terns from behind and below. The adaptiveness of this flight path is discussed. The skuas attacked singly and in groups of up to five. Group attacks were more successful than single bird attacks, but the return per bird was greatest when the skuas attacked in pairs. The success of attacks also depended upon the species of tern being attacked. The duration of attacks varied with the number of skuas in the attack, the species of tern being attacked and the outcome of the attack. Attacks were directed more frequently against Sandwich Terns than against Common Terns, correlating with the success of the attacks against these two species.     

Foods of the south polar skua Catharacta maccormicki in the eastern Larsemann Hills,Princess Elizabeth Land,East Antarctica

Regurgitated pellets and food remains were collected near nest sites, and from a club site, of south polar skuas Catharacta maccormicki in the eastern Larsemann Hills, Princess Elizabeth Land, East Antarctica, during the skuas' presence in the area. The samples indicated that the snow petrel Pagodroma nivea, the most abundant seabird species breeding locally, formed the major dietary component, comprising some 66% of food items identified in pellets and 80% of the food remains obtained. Adélie penguins Pygoscelis adeliae (which do not breed in the Larsemann Hills), other seabirds, fish and marine foods were rarely found as remains or in pellets. However, refuse (meat, fish and vegetable remains) taken as food by skuas from nearby stations occurred in pellets at all sites and formed about 12% of the food remains collected and identified. In this study, foods taken by skuas were related both to the local breeding distribution of snow petrels, and to the possession of a feeding territory. Birds without feeding territories took somewhat fewer snow petrels and included more refuse from local stations in their diet, as did those at the club site. Future monitoring of the influence of anthropogenic foods (and inedible waste materials) on the species' ecology is considered important.     

Phylogeography of the southern skua complex-rapid colonization of the southern hemisphere during a glacial period and reticulate evolution

Whilst we have now a good understanding how past glaciation influenced species at the northern hemisphere, our knowledge of patterns and modes of speciation is far more limited for the southern hemisphere. We provide mtDNA based data on the phylogeography of a circumpolar distributed southern hemisphere seabird group-the southern skua complex (Catharacta spp.). Diversification of southern skuas dates between 210,000 yBP and 150,000 yBP and coincides with a glacial spanning 230,000-140,000 yBP. Skuas most likely first inhabited the Antarctic continent, in the course of global cooling and increasing glaciation spread to the sub-antarctic islands and Tristan da Cunha and finally colonized Patagonia and the Falkland Islands at the glacial maximum. Despite significant differences between taxa most populations still exchange genes with neighboring populations of other taxa and speciation is incomplete.     

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.     

THE ANNUAL RE-OCCUPATION OF BREEDING SITES BY THE FULMAR

The Fulmar has a long period at the breeding colony prior to egg-laying. The pattern of annual occupation and build-up in numbers has been examined in detail at Marsden, Co. Durham, at a colony in which over 100 eggs are laid annually (Order 3 of Fisher's classification). The re-occupation of the cliff starts in early November with an occasional visit by one or two birds. The main period of activity at the cliff is during the morning and, as the numbers build up, the diurnal period of occupation increases. By mid-December the first birds to arrive in the colony do so before dawn and the last to leave remain well after dark until near midnight. Almost throughout the pre-egg stage, the colony is deserted each night and re-occupied the next day and birds only stay regularly overnight just before egg-laying. A similar pattern of occupation occurs after breeding but in the reverse order. The numbers of birds at the colony in January and February exceed the breeding population and include many non-breeders. The non-breeders progressively decline in numbers until May when only the breeding birds remain with a few non-breeding birds. The daily variation in the numbers of birds at the cliff is influenced by the wind speed. In general, the birds leave the colony under freshening conditions and the number present at the colony can be interpreted in terms of the wind conditions over the last three days. It is suggested that the synchronised departures are primarily feeding trips, the birds using the strong winds to reach feeding areas, except that the departure just before egg-laying is linked to egg development and synchronised laying in the colony. Competition between Fulmars and Kittiwakes for nesting sites usually results in the Kittiwakes gaining the site. This is achieved by the Kittiwakes taking over the Fulmar sites during one of the latter's departures.