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 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.     

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.     

Feeding territoriality and the reproductive consequences in brown skuas<Emphasis Type="Italic"> Catharacta antarctica lonnbergi</Emphasis>

In the maritime Antarctic, brown skuas (Catharacta antarctica lonnbergi) show two foraging strategies: some pairs occupy feeding territories in penguin colonies, while others can only feed in unoccupied areas of a penguin colony without defending a feeding territory. One-third of the studied breeding skua population in the South Shetlands occupied territories of varying size (48 to >3,000 penguin nests) and monopolised 93% of all penguin nests in sub-colonies. Skuas without feeding territories foraged in only 7% of penguin sub-colonies and in part of the main colony. Females owning feeding territories were larger in body size than females without feeding territories; no differences in size were found in males. Territory holders permanently controlled their resources but defence power diminished towards the end of the reproductive season. Territory ownership guaranteed sufficient food supply and led to a 5.5 days earlier egg-laying and chick-hatching. Short distances between nest and foraging site allowed territorial pairs a higher nest-attendance rate such that their chicks survived better (71%) than chicks from skua pairs without feeding territories (45%). Due to lower hatching success in territorial pairs, no difference in breeding success of pairs with and without feeding territories was found in 3 years. We conclude that skuas owning feeding territories in penguin colonies benefit from the predictable and stable food resource by an earlier termination of the annual breeding cycle and higher offspring survivorship.Research licence: Umweltbundesamt Bonn 13.4-94003-1/5-7.     

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.     

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.     

Long-term changes in population size,distribution and productivity of skuas (<Emphasis Type="Italic">Stercorarius</Emphasis> spp.) at Signy Island,South Orkney Islands

In this study, we investigate the numbers, productivity and territory distribution of the two species of skuas (brown Stercorarius lonnbergi and south polar Stercorarius maccormicki) breeding at Signy Island, South Orkneys, and compare the results with trends elsewhere. Comparison with previous counts indicates a biphasic increase in brown skuas at Signy Island; much faster from 1958/1959 to 1982/1983 (3.3 % per annum), than in subsequent years (0.4 % per annum from 1983/1984 to 2013/2014). Relative distribution of territories has changed little over time. The reduced rate of population growth in recent years was broadly coincident with a decrease in numbers of penguins (and therefore potential prey), which may also explain recent reductions in skua numbers at other Antarctic sites. As prey have become limiting, breeding success of brown skuas at Signy Island is now slightly lower than in the 1950s/early 1960s, but timing of breeding does not appear to have changed. Brown skuas at Signy Island may still have enough resources to start breeding, but as the season progresses and availability of resources declines, chick survival is reduced. South polar skuas have declined from ten pairs in 1982/1983 to one pair in 2013/2014, and mixed pairs have increased from one to three pairs. A review of the literature indicated that although population trend data are available for relatively few sites elsewhere in the subantarctic and Antarctic, numbers of brown skuas appear to be generally decreasing or stable, and of south polar skuas to be stable or increasing.     

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.     

Survival, site and mate fidelity in South Polar Skuas Catharacta maccormicki at Anvers Island, Antarctica

In 1974–1975, 34 adult South Polar Skuas Catharacta maccormicki were colour-ringed on 18 nest territories at Bonaparte Point, Anvers Island, near Palmer Station along the Antarctic Peninsula. Subsequently, the area was searched for these birds during the austral summers of 1975–1976 to 1984–1985 and in 1987–1988 and 1989–1990. Fifty-three percent were seen in 1984–1985, 32% in 1987–1988 and 21% in 1989–1990. Annual survival rate averaged 95% from 1974–1975 to 1984–1985; no sexual differences were detected (n = 28 of known sex). Strong territory and mate fidelity were apparent: 34 skuas averaged 1.1 nest territories and 1.7 mates each in 16 years. Only 4 of 34 individuals (all females) were known to change territories, and each territory change involved a change of mates. Although males showed higher territory fidelity than females (P < 0.01), most females (four of five) retained their territories when previous mates failed to return. Seventeen of 34 birds changed mates a total of 24 times; at least 20 mate changes followed the death or disappearance of the former mate. Males showed slightly higher mate fidelity than females (P < 0.04). Female South Polar and Brown Skuas Catharacta lonnbergi did not differ in territory or mate fidelity. From 1974–1975 to 1984–1985, 120 South Polar Skua chicks were ringed on 18 nest territories on Bonaparte Point: 17 were resighted in the Palmer area when they were 3–10 years old. All 15 returnecs were found within 3 km of their natal nest sites, and four of them occupied nest territories on Bonaparte Point.     

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.     

Behavioural ecology of lonnbergi skuas in relation to environment on the Chatham Islands,New Zealand

Abstract

A population of lonnbergi skuas (family Stercorariidae) was studied over December and January 1974–75 on Rangatira Island, Chatham group (44°22′S,176°11′W). The main aim was to compare the behavioural ecology of these birds with that of maccormicki skuas of Antarctica, and to assess the moderating effect of environment on habits. The skuas held extensive territories along the coast and in inland areas clear of forest and bracken. Five of the 11 study territories were occupied by trios of adult birds. A major effect of the vegetation was to reduce contact between neighbours on the ground at territorial boundaries, and most territorial defence was by aerial display and attack. Eggs were laid from late September, and chicks began flying in late December and January. This breeding schedule is about 2 months earlier than for lonnbergi at Signy Island or for skuas in Antarctica, but falls on a trend line of latitude against egg-laying when all populations are considered. Seventeen chicks were reared on the 11 territories, a breeding success of 1.55 chicks per nest. This high success rate was attributable largely to success in rearing both chicks when two were hatched. This is seldom achieved in more extreme climates, and is apparently determined by the intensity of sibling competitiveness. It is hypothesised that this intensity is graded, and aggression by the older chick of the pair is provoked more easily by food shortage as environmental conditions become increasingly severe. The most important prey items recorded were adult petrels, especially broad-billed prion (Pachyptila vittata vittata) and white-faced storm petrel (Pelagodroma marina maoriana) captured on the ground at night. The skuas were mostly inactive during the day, and were not seen to fish or to chase or harass any other bird on or about the island. Southern blue penguins (Eudyptula minor minor), though very common on the island, were not attacked by the skuas. Skuas were not seen to attempt to dig out petrel or penguin nesting burrows. Of the differences in ecology recorded, only the presence of three adults on territories appears to be characteristic of lonnbergi; it has not been recorded for maccormicki or hamiltoni, and is either unknown or very rare in skua.     

Experimental infection of American winter flounder (Pseudopleuronectes americanus) with Glugea stephani (Microsporida)

Young-of-the-year Pseudopleuronectes americanus were captured in Sandy Hook Bay, New Jersey and maintained in aquaria at 16–19° C for 115 days in 1982 and 127 days in 1983. Half of them were experimentally exposed to Glugea stephani and the others were sham exposed. During the course of these experiments fish from both groups died as a result of G. stephani infection, demonstrating Glwgea -induced mortality in this species. Infections were found in both control and experimentally exposed fish. Mortality of Glwgea -infected fish in the exposed group was 63% while that in the controls was 28.5%.     

SYSTEMATIC NOTES ON BIRDS OF THE CAPE PROVINCE: PART III.—PARISOMA SUBCAERULEUM (Vieillot): PART IV.—ANTHUS RICHARDI (Vieillot): PART V.—GALERIDA MAGNIROSTRIS Stephens

Fraser, M. W. 1984. Foods of Subantarctic Skuas on Inaccessible Island. Ostrich 55: 192–195.

Analysis of over 140 prey remains and 500 pellets of Subantarctic Skuas Catharacta antarctica collected from a site on Inaccessible Island. Tristan da Cunha, between October 1982 and January 1983 showed that 13 species of bird and one invertebrate were taken. Storm Petrels (Oceanitidae) made up 52% of the items recorded, Broadbilled Prions Pachyptila vittata 21%, and Common Diving Petrel Pelecanoides urinatrix 15%. Landbirds were rarely taken.     

Contribution of young fish to total production of fish in the River Thames near Reading

The production of the young stages of four species of cyprinid fish in the River Thames was estimated. From the time when underyearling fish became fully vulnerable to the special fine mesh net used, to the end of their first year, production is 39 g/ma/year compared with 83-3 g/m2/year for fish over 1-year old. The contribution of fecundity to population production varied widely, from 6.1 to 0.4 g/m2/year, but was only a small part of total production. The most productive part of the population was that found between spawning and prior to full vulnerability to the net. Production of this part of the population could be estimated only indirectly and was found to be 58-6 g/m2/year. In the four populations studied production during the first year of life was 66 to 73 % of total cohort production in 1967 and only 39 to 64% in 1968. Total fish production in the Thames was estimated at 197 k cal/m2/year; this high result is a consequence of the very high densities found: up to 96-9 fish/m2 were present in August 1967. Ten km of the Thames would contain about 54 000 000 fish in August, falling to around 8 000 000 in winter. The production: biomass ratios for the four species varied from 1–1 to 1'9 in 1967 and from 0–7 to 2-0 in 1968.
Very heavy, possibly density dependent, mortality occurs in the first year of life; the annual instantaneous mortality rates are very high, ranging from 6–4 to 8–7. Mortality is heaviest during the first 2 to 3 months after spawning; the instantaneous mortality rates during this period vary from 4-0 to 7-3.     

Modelling black fly production dynamics in blackwater streams

SUMMARY. 1. Two predictive models were employed along with intensive field sampling to estimate production of black flies ( Simulium spp.) on snags (submerged wood) in three blackwater streams on the Georgia Coastal Plain of the southeastern U.S.A. One model predicts daily growth rate from temperature and hydrograph pattern; the other predicts habitat abundance (of snags) from river height.
2. In the sixth order Ogeechee River, annual production was twice as high in 1982 (7.1 g dry mass [=DM] m⁻² of snag surface) as in 1983 (3.6 g DM m⁻²). When converted to production per m² of river bottom, values were 35–40% of the snag surface estimates. Annual production was much lower in fourth order Black Creek (1982, 1.3 g DM m⁻² of snag surface) and much higher in the sixth order Satilla River (1975, 15.6–40.0 g DM m⁻²).
3. There was a distinct bimodal pattern of black fly production in the Ogeechee River in both years, with peaks occurring in winter and summer. Similar bimodal patterns of production were found in Black Creek and in the Satilla River. Although there appears to be an intrinsic component to the bimodal pattern, production peaks (growth rate and biomass) appear to be associated with initial stages of flooding.
4. Annual production/biomass ratios (37–85) are the highest reported for black fly populations. The variation of annual P/B ratios among sites was more strongly dependent on the temporal distribution of standing stock biomass than on differences in growth rates. Variation in production among sites appears to be due to differences in current velocity, hydro-graph variability, and abundance of coexisting consumers.     

Exploitation of inland-breeding Antarctic petrels by south polar skuas

During the austral summer of 1996/1997 we studied south polar skuas at Svarthamaren, Dronning Maud Land, Antarctica, where the world's largest known colony of Antarctic petrels is found. Our censuses suggested approximately 250 full-grown skuas and 140,000 breeding pairs of petrels were present. During their breeding season, skuas did not visit the open sea at least 200 km from the site; they relied entirely on prey caught and scavenged from the petrel colony. Because the site is so isolated, we asked whether the prey (petrels) had swamped the predators (skuas), or whether there was evidence that predator numbers were limited by the size of the prey population. Particularly at the end of the petrel incubation period, we found a close correspondence between the energy required by adult skuas and their chicks, ascertained from time budget studies, and the rate at which petrel eggs disappeared from the colony. This suggests that, in this closed system, the predator population was limited by the prey population, and that predator swamping was not an advantage that petrels gained by nesting in this remote location. Received: 12 April 1999 / Accepted: 30 June 1999     

Food and feeding ecology of Wilson's storm petrel Oceanites oceanicus at South Georgia

At South Georgia, the diet of Wilson's storm petrel was studied using the regurgitates of adults arriving to feed chicks. Feeding frequency and meal size were estimated by weighing chicks twice daily, or in some cases every 3 h during daytime. Crustaceans contributed 98% of the total number of individual items and 68% of the total weight consumed; fish, 1% of the number of items and 28% of the weight. The most abundant crustacean was the amphipod Themisto gaudichaudii, which accounted for 90% of the total number but only 44% of the total weight of crustaceans eaten. Most (79%) of the Themisto were juveniles. Euphausiids were much less numerous in the diet (5% by number) but were the main group by weight (55%); most (52%) were Antarctic krill, with juveniles and sub-adults (25–50 mm long) predominating. Mysids, copepods and barnacle larvae were also present. Fish were all myctophids (lanternfish), Protomyctophum normani and P. bolini being identified; specimens were 50–85 mm long and weighed 1–4 g. Meal sizes averaged 6-5-7-5 g (14–22% adult body mass); about 75% of chicks were fed each day (mainly at night), about 10% probably receiving meals from both parents. A review of storm petrel diets emphasizes the importance of fish to Oceanodroma species and of crustaceans to Oceanites, Garrodia and Pelagodroma. Euphausiids and amphipods (chiefly Themisto and Hyperia) are the main crustacean prey and range from 5–50 mm and 0-005-0-7 g. Myctophids are the main fish prey and range from 20–100 mm and 1–5 g. Meal size ranges from 15–25% adult body mass and chicks are fed on 50–85% of days. This low delivery rate is mainly responsible for the disproportionately slow growth and long fledging period of storm petrel chicks.     

The loss of submerged plants with eutrophication II. Relationships between fish and zooplankton in a set of experimental ponds, and conclusions

SUMMARY. 1. In 1982 and 1983 sets of experimental ponds were left with their submerged plant communities intact (plant ponds) or were cleared manually of them (cleared ponds). The ponds were all fertilized with ammonium nitrate and with variable amounts of phosphate. In 1982 fish were removed from the ponds. Zooplankton communities were dominated by large Cladocera with Daphnia prominent in the cleared ponds and Simocephalus in the plant ponds. There was no detectable effect of differential phosphorus additions on zooplankton communities or populations.
2. In 1983 zooplanktivorous fish (mainly roach) were stocked in the ponds. In the plant ponds the fish did not survive, probably through severe deoxygenation and the zooplankton community again included large-bodied Simocephalus. Fish survival was variable in the cleared ponds. Where fish stocks were absent or low (0.5–1 g m⁻²) a Daphnia- dominated community persisted; at intermediate fish stocks (18.1 g m⁻²) Eudiaptomus gracilis was predominant and where fish stock was high (22.8–29.1 g m⁻²) Bosmina longirostris , and cyclopoid copepods dominated the communities. Mean biomass of the zooplankton community declined with increase in fish stock to between 5.1 and 18.1 g m⁻² then increased.
3. On the basis of results from the experimental ponds and elsewhere, a new hypothesis is put forward to account for the switch from aquatic plant to phytoplankton dominance in eutrophicated shallow lakes. It envisages dominance by either group to be possible as alternative states over a wide range of high nutrient loadings. It suggests that each state is buffered against increased loading by mechanisms involving plant and algal physiology and zooplankton grazer populations. The nature of the buffers and the reasons by which one state may be switched to the other are, discussed.     

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.     

The food and feeding relationships of black oreo, Allocyttus niger, smooth oreo, Pseudocyttus maculatus, and eight other fish species from the continental slope of the south-west Chatham Rise, New Zealand

The food and feeding habits of black oreo and smooth oreo sampled at depths of 600–1200 m from the south-west Chatham Rise in 1983 are described, with notes on prey of eight other fish species. All these fish species fed benthopelagically. Black oreo preyed on hyperiid amphipods, salps and natant decapod crustaceans; smooth oreo on salps and amphipods. Dietary composition of both species changed with size of fish. Aspects of feeding relationships between the fish species were examined including dietary similarity and prey-size selection. Findings are compared with results of research on other New Zealand deepwater fish species.