PREDATION AND KLEPTOPARASITISM BY SKUAS IN A SHETLAND SEABIRD COLONY

Feeding methods and relations of Great Skuas and Arctic Skuas to prey were studied in a seabird colony at Hermaness, Shetland. Great Skuas obtained food by kleptoparasitism, predation and scavenging. They induced Gannets to regurgitate by interfering with their flight; grasping the Gannet by the wing or tail or pushing it down with the feet on its back. Gannets tried to escape by descending to the surface, and regurgitated during 12% of the chases, most frequently when pursued by several birds. Great Skuas caught Puffins by swooping at flocks in the colony. Puffins flying with fish to their young were also chased, releasing food on one fifth of the attacks, or escaping down to the sea and diving. Great Skuas also took Kittiwake nestlings by hovering and grasping the chick with the bill, killing and eating it on the surface. Adult Kittiwakes from nearby nests took to the air, mobbing the predator. More Kittiwakes were engaged in mobbing at unsuccessful than at successful predation attempts, indicating that colonial breeding may be of selective value under such predation. Two different estimates pointed to a Kittiwake nestling predation of 0–12 and 014 young per pair. Fledging success of Kittiwakes was estimated at 0–87-1-06 young per pair, considerably lower than at English colonies where predators are absent. In spite of the predation, the Kittiwake colony showed no signs of decrease. Agonistic behaviour and other evidence indicate that Great Skuas defend feeding territories at the seabird colony. Skuas, gulls and Fulmars competed for food at carcasses. Fulmars dominated and chased away skuas. Arctic Skuas deprived Puffins of food. They patrolled the cliff, intercepting Puffins arriving with fish, snatching it from their victim's bill, or inducing them to release fish. Puffins continuing their inward flight lost food more often (30%) than birds descending to the sea (15%)—sometimes diving below. This opportunity to escape may explain the lower success of skuas at Hermaness than at a Puffin colony farther inland from the shore (Grant 1971). Other factors being equal, proximity to the sea may thus reduce the risk of kleptoparasitism.     

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 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.     

Diet of Black-legged Kittiwakes (Rissa tridactyla) feeding chicks at two Irish colonies highlights the importance of clupeids

Capsule We examined regurgitates from Black-legged Kittiwakes during the early chick-rearing period over two breeding seasons at two colonies in Ireland where diet has not been studied previously. Clupeids were the dominant food source at both colonies, which contrasts with other studies throughout the British Isles that suggest Kittiwakes feed mainly on sandeels during this period. Our study is limited by sample size and restricted to the early breeding season, but suggests a link between reproductive success and dietary composition which should be investigated further.     

The effect of egg size on post-hatching development in the Razorbill: an experimental study

A recent experimental study reported that Thick-billed Murre Uria lomvia chicks that hatched from large eggs grew their wing feathers more quickly than did small-egg chicks. There is little evidence of this (or any other) egg-size effect on post-hatching development in other birds. Thick-billed Murres are marine birds of the family Alcidae that employ the unique "intermediate" developmental strategy: chicks go to sea after 15-30 days at the nest site, at <30% of adult mass, accompanied by their male parent. Rapid feather growth during the brief nestling period is critical to enable chicks to make the transition from life at the nest site to life at sea quickly and safely. At the Gannet Islands, Labrador, Canada, in 1996 and 1997, I tested whether egg size has the same effect on wing-feather growth in the Razorbill Alca torda , another of the intermediate auks. To control for underlying correlations between egg size and other parental attributes, eggs were switched randomly among pairs. As in other birds, egg size strongly predicted hatchling mass, and to less extent hatchling size (tarsus length), but had no effect on the rate at which nestlings gained mass. However, egg size had the same effect on wing growth in Razorbills as in Thick-billed Murres: the wings of large-egg chicks began rapid, linear growth sooner, indicating that early development of wing feathers was enhanced in large-egg chicks. Differences in wing length established in this manner persisted through the nestling period. Egg-size effects on feather growth have not been detected in experimental studies on other birds, suggesting that effects of the magnitude seen in Razorbills and Thick-billed Murres might reflect evolutionary priorities in the post-hatching development of intermediate auks.     

Winter diet of the little auk (Alle alle) in the Northwest Atlantic

The little auk (Alle alle) is one of the most numerous seabirds in the world, but their winter prey selection has never been thoroughly studied. In the present study, we analyzed the proventricular contents of 205 little auks caught in coastal areas off southwest Greenland during December–February and off Newfoundland in March. Large Calanoid copepods are known to be the main prey during summer. We found krill to become the crucial winter prey in both areas, followed by Themisto spp. and young capelin (Mallotus villosus). No difference was found between the diet of juvenile and adult birds. Copepods constituted around 1 % of the diet and close to all copepods were in birds caught near Cape Farwell in December. These findings provide new and important insight into the forage ecology of the little auk, and they support other studies showing that large Calanoid copepods in the Arctic decent for winter hibernation at depths that are below the diving range of the little auks. More studies to determine offshore diet and annual variation are, however, needed in order to get a more complete picture.     

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

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

Diets and feeding of Fulmars Fulmarus glacialis during the breeding season:a comparison between St Kilda and Shetland colonies

Diets and nest attendance of Fulmars at Foula, Shetland and St Kilda, Outer Hebrides were examined during the breeding season. At Foula, sandeels formed the bulk of the diet, with fish offal and pelagic zooplankton of minor importance. At St Kilda, 71% of regurgitates consisted of pelagic zooplankton, probably captured at night. Dietary overlap between the two colonies was 14% by species composition. Foraging trips from Foula shortly after chick hatching generally lasted for less than 10 h, while trips from St Kilda late in chick rearing often lasted more than 24 h. The diurnal pattern of feeding suggested mainly nocturnal foraging from St Kilda but both diurnal and nocturnal foraging from Foula.     

Variation in the correction factor used for converting counts of individual Guillemots Uria aalge into breeding pairs

The correction factor k (breeding pairs/total count of birds) was measured for two nesting groups of Guillemots Uria aalge over 6–7 years. The value varied between different subgroups but there were no significant differences between groups or between years. These results, and those from other studies, suggest that a value of 0–67 is applicable to a range of colonies. Values of k for Razorbills Aha tarda were more variable. A plea is made for the presentation of original counts rather than 'corrected' figures in published work.     

The attendance and absence of adult Kittiwakes Rissa tridactyla from the nest site during the chick stage

In recent years there have been indications that the attendance of Kittiwakes Rissa tridactyla at the nest whilst containing young has changed, possibly influenced by food shortage. This paper examines the attendance of parents at the nest at North Shields, England, at a time when the breeding success was high and there was no indication of food shortage. It was evident that there was considerable variation in the attendance of individual parents, and chicks were left alone for the first time between 9 and over 40 days after hatching. Chicks were left alone overnight more frequently than during the day; this is related to the much longer overnight feeding trips by the parents.
The recording method used in this study gave a continuous trace of the presence of each adult, and it has been used to calculate the errors made from spot observations.     

Pelagic distribution and numbers of the Antarctic petrelThalassoica antarctica in the Weddell Sea during spring

Estimates of the population size of the Antarctic petrelThalassoica antarctica are hampered by its breeding locations in remote nunatak areas of the Antarctic continent. Studies at sea can provide additional information on numbers. Seabird censuses during spring 1992 showed few Antarctic petrels in the western part of the Weddell Sea, but high numbers east of the South Sandwich Islands. The vast majority of the birds occurred in a band of 300 km north to 150 km south of the outer ice edge. Numbers at sea fluctuated in agreement with synchronized colony attendance patterns in the pre-breeding phase and peaked shortly before egglaying when colonies are completely deserted. In this period, densities of Antarctic petrels in the marginal ice zone suggest that at least 2.7±0.5 million individuals occur in the Weddell Sea. It is likely that similar numbers occur immediately east of the study area. The distribution of the petrels matches the location, but not the size, of known breeding colonies in Dronning Maud Land and Coats Land, which suggests that important colonies in this area remain to be discovered. The observations imply that Antarctic petrels in the Weddell Sea commute over ice a surprisingly large distance of at least 2,000 km for a single pre-breeding visit to the colonies.     

A comparative study of the diet of auks killed during an oil incident in the Skagerrak in January 1981

Stomach contents were examined from guillemots Uria aalge (Pont.), Razorbills Alca torda Linnaeus and Little auks Alle alle (Linnaeus) killed by oil in the Skagerrak in January 1981. The range of prey species was small; the commonest fish found in the stomachs were Gobiusculus flavescens (Fabricius), Crystallogobius linearis (Düben), Sprattus sprattus (L.), Trisopterus minutus (L.) and Ammodytes sp. The relative importance of these fish depended on bird species and locality. Guillemots contained gadoids more frequently and sandeels less frequently than razorbills, and the composition of guillemot samples from Oslo Fjord differed from that of samples from Sweden. Invertebrate remains were uncommon in all bird species; the commonest items recorded were polychaete jaws. The frequency of occurrence of items in the diet of guillemots from Oslo Fjord was apparently influenced by the extent of oiling suffered by the birds.     

Seasonal differences in at-sea activity of seabirds underline high energetic demands during the breeding period

We assessed seasonal differences in at-sea activity of Lesser Black-backed Gulls Larus fuscus, Black-legged Kittiwakes Rissa tridactyla and Common Guillemots Uria aalge in the south-eastern North Sea. The three species correspond to different ecological groups, with Lesser Black-backed Gulls representing omnivorous generalists, Kittiwakes representing surface-feeding pelagic seabirds, and Guillemots representing pursuit-diving pelagic seabirds. Using data from aerial surveys, we differentiated between active (flying or scavenging at fishing vessels) and inactive behaviour (swimming). We estimated the activity budgets of all three species for the different seasons and tested for differences in activity between different seasons. All species exhibited significant seasonal differences in activity, with the highest levels of activity observed during the breeding season. Numbers of flying auks were, however, exceptionally low in autumn due to moult and guarding of not-yet fledged chicks at sea. Our results underline the high energetic demands of the breeding season that lead to increased foraging and travelling activity.     

Effects of temperature on spring bloom dynamics of epilithic diatom communities in the Gulf of Bothnia

Abstract. In order to forecast consequences of climatic changes for littoral algae in coastal ecosystems, the effects of temperature increase on spring bloom dynamics of epilithic diatom communities were studied in two ways: (1) Communities were compared from sites receiving different amounts of cooling water discharge from a nuclear power plant. (2) Communities from the exceptionally warm spring of 1989 were compared with those from the same sites in the preceding normal years. The studies were carried out in and around the Forsmark Biotest Basin, an artificial waterbody that receives brackish cooling water from the Forsmark nuclear power plant on the Swedish east coast. Species composition and biomass data from 200 epilithic diatom samples taken in the period January-May from 1983 to 1989 were analysed. Multiple regression analysis was used to show the responses of diatom cover, community diversity and abundances of individual taxa to water temperature and other environmental factors. Constrained ordination (CCA) was used to display the relationship of overall community composition to environment. Increased water temperature resulted in higher biomass through a chain of ecological effects. Strong reduction of ice cover was crucial in this chain, resulting in reduced loss of diatom cells by reduced abrading and higher primary production by higher availability of light, nutrients and substrate. The conclusion is that large blooms of fast-growing epilithic diatoms in large colonies can occupy the niche that emerges when the ice-free season is prolonged in the northern Baltic Sea and areas similar in salinity and climatic conditions. Dramatic species shifts may only be expected if the winter ice cover is totally absent.     

Diet and fish stock availability as possible factors in the mass death of auks in the North Sea

In February 1983 more than 30 000 birds died on the North Sea coasts of England and Scotland. The birds were unoiled, and there was no immediately obvious cause of death. Stomach contents of auks killed in this incident were examined to ascertain whether they provided any indication of the causal factors. Analysis of food remains showed differences between sample sites and between guillemots, Una aalge (Pont.), and razorbills, Alca tarda L. In all areas gadoid remains comprised a higher, and sandeels a lesser, proportion of the stomach contents in guillemots than in razorbills. The relative importance of clupeid remains varied with area. These differences, combined with current trends in the fisheries biology of major prey species, provided an ecological basis for a hypothesis explaining the incident. It was concluded that starvation was probably a major factor in the death of the auks, and that this may have been related to a combination of adverse weather conditions and changing patterns of abundance in North Sea sprat, Sprattus sprattus (L.), populations.     

THE EFFECTS OF EXPERIENCE AND BREEDING STATUS ON THE USE OF A WINTERING SITE BY BEWICK'S SWANS CYGNUS COLUMBIANUS BEWICKII

Bewick's Swans were identified individually at Slimbridge from 1968-69 to 1975-76, both within one winter and from one winter to the next, by bill markings or large numbered leg rings. Units, rather than individuals, were used for analysis, each unit being placed into one of two categories (New, Experienced), and into one of four classes (family, pair, single, yearling).
The Experienced category arrived proportionally earlier, had fewer one-day attendances, stayed longer, were absent less, and departed later than the New category. Experienced families were absent more than the other Experienced classes and, in the early part of the winter, left proportionally ahead of them. Arrivals of the New classes were at different rates: yearlings built up quickest, followed by singles, pairs and families. Families had more one-day attendance than yearlings and singles, and pairs were absent more than the other classes. The departures of the New classes, except of families, were closely synchronised.
The different behaviours of the classes which determine length of stay are discussed. The birds most likely to return were those with a long attendance in a previous season.     

The effect of water flow on growth and reproduction of celleporella hyalina (l.) (bryozoa : cheilostomata)

Colonies of Celleporella hyalina (L.), settled on glass Petri dishes, were grown in the Menai Straits. Water flow over the colonies was either unrestricted or increasingly restricted by short and long funnels fitted over the Petri dishes. Under all conditions, colonies grew exponentially for the first few weeks, but then growth decelerated. Growth rate at standard size was minimal in winter (December–February). Restricted water flow enhanced growth during winter but retarded growth at other times. Colonies began sexual reproduction at a size of ≈57 autozooids, but at different ages depending on growth conditions. When colonies had accumulated 1000–2000 autozooids, the number of reproductive zooids (males plus females) per autozooid became asymptotic in the ratio of 1.0 : 2.0 : 3.6 in conditions of restricted, semi-restricted, and unrestricted water flow, respectively. The proportion of sexual zooids that were reproductively active was greater among colonies experiencing greater water flow and, for all treatments, was greatest in June and least in winter. The experimentally induced trends in growth and reproduction were reversed by translocating colonies from one regime of water flow to another. Colonies had lower budding rates when grown close to neighbouring colonies than when isolated, but the ratio of reproductive zooids to autozooids was similar in both treatments. Ratios of female to male zooids varied among colonies but with no clear relation to the experimental conditions, suggesting that relative investments in male and female functions are largely under genetic control. The life history of C. hyalina is well-suited to the ephemeral frondai tissue of Laminaria saccharina L. (Lamour), commonly used as a substratum in British waters. Early sexual maturity and sexual activity, concurrent with somatic growth, maximize average sexual output in the circumstances of unpredictable longevity.     

Seasonal changes in serum glucose and condition of the plaice, Pleuronectes platessa L.

Monthly sampling of plaice caught off the north-east coast of Scotland between February 1983 and May 1984 revealed a seasonal increase in condition factor, hepatosomatic index and serum glucose, which appeared to be related to the period of active feeding. The spleen indices followed a similar seasonal increase, although this was not statistically significant, while the kidney indices showed little change over the 16 months. The peak for the gonadosomatic index was in February/March 1984, but among these plaice (aged 4–5 years) there were still large numbers of immature males and females and there was no increase in 1983. Apart from the hepatosomatic index for January 1984, there was no significant difference between male and female plaice.     

Breeding strategies of Antarctic Petrels Thalassoica antarctica and Southern Fulmars Fulmarus glacialoides in the high Antarctic and implications for reproductive success

Breeding strategies of two closely related fulmarine petrels were studied on Ardery Island, on the continental coast of East Antarctica, where short summers are expected to narrow the time-window for reproduction. Both species had a similar breeding period (97 days from laying to fledging) but Antarctic Petrels Thalassoica antarctica bred up to 16 days earlier than Southern Fulmars. During the pre-laying exodus, all Antarctic Petrels deserted the colony, whereas some Southern Fulmars Fulmarus glacialoides remained. Antarctic Petrels exhibited stronger synchronization in breeding, made longer foraging trips and spent less time guarding their chicks than Southern Fulmars. Overall breeding success of both species was similar but failures of Antarctic Petrels were concentrated in the early egg-phase and after hatching, when parents ceased guarding. Southern Fulmars lost eggs and chicks later in the breeding cycle and so wasted more parental investment in failed breeding attempts. Different breeding strategies may be imposed by flight characteristics; Southern Fulmars are less capable of crossing large expanses of pack ice and need to delay breeding until the sea ice retreats and breaks up. However, due to the short summer they risk chick failure when weather conditions deteriorate late in the season.     

Foraging behaviour of Razorbills Alca torda during chick-rearing at the largest colony in the Baltic Sea

Capsule: Foraging behaviour in the Razorbill Alca torda during breeding was similar to that found elsewhere, aside from dive shape.

Aims: To investigate the foraging behaviour of Razorbills during the breeding season at the largest colony in the central Baltic Sea.

Methods: A combination of global positioning system (GPS) and time-depth recorder (TDR) devices were used on Razorbills breeding on the island of Stora Karlsö, Baltic Sea, during the chick-rearing period.

Results: Five GPS tracks and nine TDR logs were retrieved from 12 Razorbills, and 7399 dives were analysed. Razorbills foraged south and southwest of the colony. Maximum and mean (±sd) foraging range from the colony was 72.7?km and 13.1?±?13.5?km, respectively. Mean dive depth (15.3?±?2.4?m) and duration (53.1?±?8.5?s) were similar to those of a more southern Baltic Sea Razorbill colony. Dive depth had a bimodal distribution, with 70% of dives deeper than 10?m and 30% shallower than 10?m. There was a clear diel foraging pattern with 89% of dives occurring during daytime and a higher proportion of shallow dives at night. Unexpectedly, dives were primarily U-shaped. The Razorbills spent 31% of their overall time activity budget flying or diving.

Conclusion: Aside from dive shape, foraging behaviour was consistent with that reported at other colonies of Razorbills. Inconsistency in dive shape may be due to a bimodal foraging strategy, local prey behaviour or competition with the Common Guillemot Uria aalge.