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 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.     

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.     

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.     

The influence of food availability on the breeding effort and reproductive success of Arctic Skuas Stercorarius parasiticus

Territorial attendance, chick growth rate and breeding success of Arctic Skuas Stercorarius parasiticus in Shetland were lowest in the late 1980s when recruitment of Sandeels Am-modytes marinus in the surrounding waters was poor. The relationships between both fisheries-based and avian indices of food availability and annual variation in Arctic Skua chick growth and breeding success between 1976 and 1994 were better described by a threshold effect rather than linear functions. Arctic Skuas conform to the model proposed by Cairns, which predicts the responses of seabirds to changes in prey availability. Skua clutch size, egg volume, hatching success and hatching date were not reliable indices of Sandeel availability. However, annual fluctuations in Arctic Skua breeding numbers may be a useful indicator of changes in prey abundance.     

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.     

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.     

A histochemical comparison of fibre types in the M. pectoralis and M. supracoracoideus of the great skua Catharacta skua and the herring gull Larus argentatus with reference to kleptoparasitic capabilities

Skuas, which are closely related to gulls, frequently use a specialized feeding method (kleptoparasitism) by which they rob other seabirds of their food. This paper tests the idea that skuas have evolved as specialist kleptoparasites.
The fibre type composition of the M. pectoralis, M. supracoracoideus and M. iliofibularis of a great skua Catharacta skua (Brünn.) and a herring gull Larus argentatus (Pontopp.) was determined by three enzyme histochemical methods commonly used for mammalian fibre classification; the reactions for alphaGPDH, NADH-TR and mATPase activity.
In both species slow fibres were present only in the M. iliofibularis, and fast twitch glycolytic fibres were not present in any of the muscles. The M. pectoralis and M. supracoracoideus of both species consisted entirely of the fast twitch oxidative-glycolytic fibres.
The overall metabolic enzyme activities of the muscles were assessed in terms of the proportions of fibres with high, intermediate and low metabolic enzyme activity. The overall levels of oxidative and glycolytic enzyme activity were significantly higher in the M. pectoralis than in the M. supracoracoideus and significantly higher in both of these than in the M. iliofibularis. This was true of both species.
The oxidative and glycolytic activities of all three muscles of the great skua were significantly higher than those of the homologous muscles of the herring gull. This was particularly true of the M. pectoralis and M. supracoracoideus. It is suggested that this difference between great skuas and herring gulls enables the former to be more effective aerial kleptoparasites than the latter.     

Enigmatic phylogeny of skuas (Aves:Stercorariidae)

Multiple sources of evidence show that the skuas (Aves:Stercorariidae) are a monophyletic group, closely related to gulls (Laridae. On morphological and behavioural evidence the Stercorariidae are divided into two widely divergent genera, Catharacta and Stercorarius, consistent with observed levels of nuclear and mitochondrial gene divergence. Catharacta skuas are large-bodied and with one exception breed in the Southern Hemisphere. Stercorarius skuas otherwise known as jaegers) are smaller bodied and breed exclusively in the Northern Hemisphere. Evidence from both mitochondrial and nuclear genomes and from ectoparasitic lice (Insecta:Phthiraptera) shows that the Pomarine skua, S. pomarinus, which has been recognized as being somewhat intermediate in certain morphological and behavioural characteristics, is much more closely related to species in the genus Catharacta, especially to the Northern Hemisphere-breeding Great skua, C. skua, than it is to the other two Stercorarius skuas, the Arctic skua, S. parasiticus and the Longtailed skua, S. longicaudus. Three possible explanations that might account for this discordant aspect of skua phylogeny are explored. These involve (i) the segregation of ancestral polymorphism, (ii) convergent evolution of morphology and behaviour or (iii) inter-generic hybridization. The available evidence from both nuclear and mitochondrial genomes does not exclude any of these hypotheses. Thus, resolution of this enigma of skua phylogeny awaits further work.     

VARIATION IN THE CAPPED WHEATEAR

Williams, A. J. 1980. Aspects of the breeding biology of the Subantarctic Skua at Marion Island. Ostrich 51:160-167.

The breeding biology of the Subantarctic Skua Catharacta antarctica lonnbergi was studied in two austral summers at Marion Island. Eggs were laid between 23 October and 19 December. The clutch size was one (6%) or two (94%) eggs. The laying interval was two (20%) or three (80%) days and the incubation period was 29 days. Chicks could fly at 50–65 days after hatching. Chicks from first and second hatched eggs in the same brood had similar growth rates. Chicks in one- and two-chick broods had similar growth rates until 50 days after hatching when chicks reared singly were heavier. Egg mortality was 15,8%, chick mortality was 34,8% and the overall breeding success was 50,9%. The results are discussed in comparison with previous studies of Subantarctic and South Polar C. maccormicki Skuas.     

ORNITHOLOGICAL TRANSECTS IN THE NORTH ATLANTIC

An analysis is presented of observations made in the North Atlantic from 14 August to 10 September 1966, and comparisons are made with previously published data relating to this area. The majority of the records were obtained during transects along latitudes 55° N. and 58° N., but other data were collected in Labrador and Newfoundland coastal waters. During the period spent at sea 30 species were recorded, including five passerine species that came aboard ship. Manx Shearwaters were seen in Newfoundland waters where there have been few previous records. The record of a Balearic Shearwater appears to be the most northerly to date. Some evidence of correlation was found between the numbers of Great Shearwaters seen and the surface water temperature, but there was little evidence of correlation in the case of the Fulmar. Other points of interest are the records of moult in the Great Shearwater, and the records of Grey Phalaropes and Sabine's Gulls. An analysis of the skua movements suggests that the Arctic Skuas seen originated from Scandinavia, the Great Skuas from Iceland and the Pomarine and Long-tailed Skuas from the Canadian arctic.     

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).     

Chick loss in the Falkland Skua Catharacta skua antarctica

Data on reproductive success of 110 Falkland Skua Catharacta skua antarctica pairs were gathered during the austral summers of 1988–1989 and 1990–1991 on New Island, Falkland Islands. Adults laid two eggs 2–3 days apart and began incubation with the first egg. For the years combined, 1.39 chicks per nest hatched and 0.84 chicks per nest fledged (fledging was defined as surviving to 16 days of age). Brood reduction was common; 43% of the two-chick broods were reduced to one, and mortality was concentrated on the younger chick. Although asynchronous hatching and differential death are consistent with Lack's brood reduction hypothesis, application of O'Connor's quantitative criterion revealed that sibling competition may not be responsible for the observed chick mortality. Furthermore, because no aggressive interactions between chicks were observed or detected indirectly, siblicide may be absent in this population. Instead, predation modified by a variety of factors may have led to the greater mortality of the second-hatched chick.     

Brent Geese (<Emphasis Type="Italic">Branta bernicla</Emphasis>) Breeding Associations with Pomarine Skuas (<Emphasis Type="Italic">Stercorarius pomarinus</Emphasis>) on the Mainland Tundra

This article is based on data that were collected in the years 2000?2007, 2012, and 2014 in the vicinities of Medusa Bay (73°21′ N, 80°32′ E) and in 2002 at the mouth of the Uboynaya River (73°37′ N, 82°10′ E), in the northwestern part of the Taimyr Peninsula. In years when the abundance of lemmings is high, brent geese may breed not only near nests of snowy owls and rough-legged buzzards, but also sparsely in the mainland tundra, often without any protection. Some such nests are successfully incubated until hatching. A considerable part of these dispersed nests appears to be associated with a nest or territory of pomarine skuas that are able to scare away the main tundra predator, the arctic fox, to a distance of about 500 m from their nests. Brent geese that breed within this distance to theses nests gain additional protection against arctic foxes. However, brent geese do not display a tendency to place their nests closer to pomarine skua nests. The mean distance from geese nests to pomarine skua nests or centers of their territories comprised 2/3 of the mean distance between nests of pomarine skuas and turned out to be quite stable over the years and in two different tundra areas.     

1. A CENSUS AND STUDY OF BIRDS IN ALBANY OPEN THORNBUSHVELD

Williams, A. J. &; Cooper, J. 1983. The Crowned Cormorant: breeding biology, diet, and offspring reduction strategy. Ostrich 54:213-219.

Crowned Cormorants Phalacrocorax coronatus were studied at Dassen and Marcus Islands. The most frequent clutch was three eggs. Egg size varied within clutches with first-laid eggs being largest and heaviest and subsequent eggs progressively smaller and lighter, The mean laying interval was 2,2 days, the mean laying-to-hatching interval was 23,0 days, and the hatching interval was one day. The normal incubation period was 22.4 days. The weight of hatchings was related to the position of the originating egg in the laying sequence. Chicks were fed within 24 h of hatching. Chick development is described over the first 35 days. One chick could fly at 35 days. Hatching success was 48,2%. Hatching success was greatest in second-laid eggs, least in last-laid eggs. The mean number of chicks hatched at a nest was two. Mean diving time was 23,5 s. Most food was fish, particularly klipfish Clinidae and pipefish Syngnathus, 60–160 mm long. The number of offspring produced can be related to food availability by interaction of difference in egg size, hatching asynchrony, and the preferential feeding by adults of the strongest-begging chick. There is a trend towards producing two chicks, normally those from the first two eggs to be laid.     

Communally breeding skuas: breeding success of pairs, trios and groups of Catharacta lonnbergi on the Chatham Islands, New Zealand

One-third of territories on South East Island are occupied by trios or groups of communally breeding skuas. Comparisons were made of the reproductive performance of pairs, trios and groups over three (and ten) seasons to assess whether any 'advantage'–in terms of offspring production–could be demonstrated for communal breeding. Egg size was not significantly different between pairs and trios, or between years. There were no significant differences between the chicks of pairs, trios or groups in growth characteristics and chick condition appeared similarly good. Overall breeding success was generally lower for trios than for pairs. Although chick production per territory was not significantly different between pairs and trios, it was substantially lower on a per-adult basis. The breeding success of this population is generally high compared to others studied, however, there is no evidence to suggest that skua communal breeding has advantages in terms of offspring production.     

The breeding biology and factors affecting reproductive success in rockhopper penguins <Emphasis Type="Italic">Eudyptes chrysocome</Emphasis> at Macquarie Island

Adult mass changes, egg morphometrics, chick growth rates, fledging masses, reproductive success and reasons for reproductive failure were examined in rockhopper penguins at Macquarie Island from 1993/1994 to 1995/1996. Mean arrival masses, growth rates of chicks and fledging masses exhibited inter-annual variability, while egg morphometrics, hatching success (68.0±6.0%) and reproductive success (47.3±8.3%) were constant between years. Reproductive failures occurred primarily during incubation, with the majority of eggs lost to great skuas. Logistic regressions revealed that no variable significantly explained hatching success, and only in 1994/1995 was fledging success significantly correlated with the position of nest in the colony (those in the centre were more successful than those on the periphery). Reproductive success during this study was relatively high, and therefore an assessment during poor years would be instructive, particularly in relation to aspects of the penguins’ foraging ecology.     

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.     

Hybridisation between South polar skua (Catharacta maccormicki) and Brown skua (C. antarctica lonnbergi) in the Antarctic Peninsula region

Hybridisation between South polar skua (C. maccormicki) and Brown skua (C. antarctica lonnbergi) in the area of the Antarctic Peninsula is known at least since the beginning of the last century but no survey has been done so far. This paper reviews information on the species composition of skua colonies of more than 10 pairs in the Antarctic Peninsula region, and the incidence of mixed pairs. Morphometrics, population size and breeding success were examined in detail at King George Island. The northward distribution of South polar skuas extended to King George Island (62°11′ S 59°00′ W), with a small outlying population on Signy Island (60°45′ S 45°36′ W), whereas Brown skuas did not breed further south than Anvers Island archipelago (64°46′ S 64°03′ W). The proportion of mixed pairs was highest at the northern end of the ∼500-km-wide hybrid zone. Body size distribution of sympatric skuas from King George Island is clearly bimodal but overlaps considerably and hybrids cannot be identified. Skua population sizes at Potter Peninsula/King George Island remained stable between 1994 and 2004. Numbers of mixed breeding pairs fluctuated more strongly than those of pure species pairs. Breeding success of Brown skuas varied the least.     

SEABIRDS OF THE SENEGAL UPWELLING AND ADJACENT WATERS

Seabird distributions were observed off Senegal during the upwelling season there, in February-March 1976, and are interpreted against an oceanographic background. Gannets Sula bassana , Pomarine Skuas Stercorarius pomarinus , Great Skuas Catharacta skua , Lesser Black-backed Gulls Larus fuscus , Black-headed Gulls L. ridibundus , Sandwich Terns Sterna sandvicensis and Royal Terns S. maxima were the dominant species on the shelf and slope adjacent to the inshore upwelling zone. Grey Phalaropes Phalaropus fulicarius were the only seabirds associated with an oceanic 'front' farther offshore; it is suggested that this and similar boundary zones are important feeding areas during the pelagic phase of this species' annual cycle.
Leach's Storm-Petrels Oceanodroma leucorhoa were seen east to 20W, but the predominant storm-petrel of the waters between 23W and the continental slope was the Madeiran O. castro. It is suggested that the hatching period of Madeiran Storm-Petrels on the Cape Verde Islands is timed to take advantage of the arrival there of young fish in the surface waters, spawned off the African coast and drifted offshore. However Madeiran Storm-Petrels make no direct use of the coastal upwelling; nor do Little Shearwaters Puffinus assimilis or White-faced Storm-Petrels Pelagodroma marina , the other winter-breeding Cape Verdes seabirds, which probably feed close to the Islands.
The Senegal upwelling is underexploited by seabirds, in contrast to comparable upwelling systems elsewhere. It is suggested that this is due to the seasonal timing of the upwelling, the scarcity of secure nest-sites for breeding seabirds, and the competing attractions of the Benguela upwelling in the South Atlantic.