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.     

Indexing winter gull numbers in Great Britain using data from the 1953 to 2004 Winter Gull Roost Surveys

Capsule Winter Gull Roost Survey data spanning 50 years were used to generate population indices.

Aims To evaluate how wintering numbers of five gull species have changed in Great Britain over the last five decades.

Methods Generalized linear models were used to relate gull numbers to habitat, site and year factors, and so derive species‐specific indices for nine regions of Great Britain. Regional models considered data from different timescales depending on coverage.

Results Patterns of change varied by species and region. All species showed increases in number over the period 1953 to 2004. In most regions, Black‐headed Gull Chroicocephalus ridibundus numbers have declined since peaks between 1973 and 1993; Common Gulls Larus canus have also declined recently in some regions. Lesser Black‐backed Gull L. fuscus numbers have increased dramatically since 1953, whereas numbers of Herring Gull L. argentatus showed large declines between 1963 and 1983. Great Black‐backed Gull L. marinus numbers have increased in the west and the Midlands, but recently declined in eastern regions.

Conclusions Numbers of wintering gulls in Great Britain have shown rapid changes over the last five decades, reflecting changes in the sizes of breeding populations. These changes are likely to be associated with changes in human activities and resource availability.     

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.     

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.     

Riding the tide: intriguing observations of gulls resting at sea during breeding

Adult Lesser Black‐backed Gulls Larus fuscus were followed during the breeding season using a high‐resolution GPS tracking system. Little is known about the foraging movements of these birds and what they do when they are away from the colony. The study revealed intriguing yet infrequent behavioural patterns showing that birds would sit on the sea surface drifting passively with the tidal current for several hours, resting at sea rather than in the breeding colony. The potential function of the observed patterns is discussed as well as the relevance of these observations to other species.     

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.     

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.     

Effects of geolocator attachments on breeding parameters of Lesser Kestrels

ABSTRACT Light level geolocators, also known as GLS loggers, are electronic devices intended for tracking the location of wide‐ranging animals using ambient light to estimate latitude and longitude. Miniaturized geolocators that can be used on relatively small migratory birds have recently become available, but little is known about the potential harmful effects of geolocators on birds. We examined the possible effects of 1.5‐g geolocators (dimensions: 21 × 6.5 × 9 mm) on the breeding success and survival of migratory Lesser Kestrels (Falco naumanni). During the 2007 breeding season, kestrels were fitted with geolocators using two attachment methods (Teflon wing harnesses and darvic bands), and geolocators were removed in 2008 after the birds returned to the breeding grounds. We found no differences in the breeding success of control and tagged pairs during the 2007 breeding season, but tagged pairs had greater fledgling mortality in the following breeding season. Furthermore, nestlings of tagged individuals had higher triglyceride and uric acid concentrations in their blood than control nestlings during the breeding season following tagging. As for return rates, 75% of tagged birds came back to the colony after the nonbreeding period, a proportion similar to that reported in previous studies. Although back‐mounts are slightly heavier and require more skill to attach, we recommend their use on small migratory raptors because most leg‐mounted geolocators in this study were damaged or rendered useless by dirt obscuring the light sensor.     

A comparison of Herring Gull Larus argentatus and Lesser Black-backed Gull Larus fuscus nest sites: their characteristics and relationships with breeding success

Nest sites used by Herring Gulls and Lesser Black-backed Gulls in a mixed colony were compared. Correlations between features surrounding a nest and breeding success at different stages of the breeding cycle are presented. Lesser Black-backed Gulls nested on more vegetated and flatter areas than did Herring Gulls, even though the latter species had a lower hatching success at less vegetated sites. The difference in the general topography of nest sites between the species suggests that the Lesser Black-backed Gull, through an individual defence strategy, may be better adapted to use sites which are accessible to ground predators than the Herring Gull. The fledging success of Lesser Black-backed Gulls tended to increase with an increased proportion of relatively tall vegetation close to the nest. The Lesser Black-backed Gull may therefore be able to leave its young unattended and rely on their concealment for protection against predators. In contrast, the Herring Gull may rely more on parental vigilance to protect young. More frequent attendance by adult Herring Gulls at the nest site during chick rearing compared with Lesser Black-backed Gulls supports this hypothesis.     

Radio‐transmitters do not affect seasonal mass change or annual survival of wintering Bicknell's Thrushes

ABSTRACT Studies of the effects of transmitters on passerines have provided mixed results, but many have revealed no negative impacts. Most such studies have been conducted during the breeding season and, as a result, little is known about the possible effects of transmitters on wintering birds. We examined the effects of transmitters on Bicknell's Thrushes (Catharus bicknelli) wintering in the Dominican Republic. We used long‐term mark‐recapture data to compare birds with radio‐transmitters (N= 5 years, 64 individuals) to birds banded, but not radio‐tagged (N= 10 years, 164 individuals). For a subset of birds in each category, we measured the change in mass between early‐winter and late‐winter and, for all birds, we calculated return rates and modeled annual survival by sex and age classes. Return rates of radio‐tagged (16%) and banded‐only individuals (17%) did not differ, and survival models including the effect of transmitter attachment were poorly supported. We also found no sex or age‐specific effects of transmitter attachment. Radio‐tagged and banded‐only birds did not differ either in the proportion of birds gaining mass during the winter or amount of mass gained. Our results indicate that attaching light‐weight transmitters (<5% of body mass) to wintering Bicknell's Thrushes did not adversely affect either body condition or annual survival.     

Skuas (Stercorarius spp.) moult body feathers during both the breeding and inter‐breeding periods: implications for stable isotope investigations in seabirds

Seabirds are mostly thought to moult during the inter‐breeding period and the isotopic values of their feathers are often therefore assumed to relate to their assimilated diet during such periods. We observed Brown Skuas Stercorarius antarcticus lonnbergi and South Polar Skuas Stercorarius maccormicki moulting on a breeding site at King George Island, Antarctica. This raises concerns about the reliability of using stable isotopes in feathers to infer feeding localities of birds during the inter‐breeding period. We analysed the δ¹³C and δ¹⁵N values of growing and fully grown body feathers collected from the same individuals. For both species, δ¹³C values of growing feathers indicated feeding areas in the Antarctic zone (breeding grounds), whereas most fully grown feathers (100% for South Polar Skuas and 93.3% for Brown Skuas) could be assigned to northern latitudes (non‐breeding grounds). However, a few fully grown body feathers of Brown Skuas (6.7% of the feathers, belonging to two birds) showed isotopic values that indicated moult in the Antarctic zone. As the growth period of those feathers was unknown, they could not be used with confidence to depict the foraging behaviour of the birds during the non‐breeding period. Although precautions must be taken when inferring dietary information from feathers in seabirds where the moulting pattern is unknown, this study shows that if the development stage of a feather (growing/fully grown) is identified, then dietary information from both breeding and non‐breeding seasons can be obtained on the same individual birds.     

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.     

Rapid long‐distance migration in Norwegian Lesser Black‐backed Gulls Larus fuscus fuscus along their eastern flyway

We studied the long‐distance migration of Lesser Black‐backed Gulls Larus fuscus fuscus breeding in northern Norway along their eastern flyway using geolocators in 2009 and 2010. The majority of birds wintered in lakes in East Africa and the southeast Mediterranean was the most important stopover area. Larus f. fuscus along the eastern flyway travelled at a net travel speed of 399 and 177 km/day during the autumn and spring migration, respectively, higher than published travel speeds for Dutch Larus fuscus migrating along the western flyway. The results suggest that the long‐distance migratory Norwegian L. f. fuscus seek to minimize time spent in transit, whereas lower travel speed during northerly spring migration may reflect differences in wind patterns or food conditions between spring and autumn.     

Influence of rodent density on nesting associations involving the Bar-tailed Godwit Limosa lapponica

The Bar-tailed Godwit Limosa lapponica rarely defends its nest aggressively against predators, but rather associates with other more aggressive species for protection. I studied the distribution of Bar-tailed Godwit nests relative to nests of the aggressive Longtailed Skua Stercorarius longicaudus, as well as to nests of two slightly less aggressive alternative 'protective umbrella' species, the Whimbrel Numenius phaeopus and the Grey Plover Pluvialis squatarola. Rodents are the main prey of the Long-tailed Skua but, in years when numbers of rodents are low, Long-tailed Skuas also eat eggs and chicks of other birds. Data on rodent densities and nest distribution were collected on Finnmarksvidda, Northern Norway, in 1987, 1989, 1990, 1992 and 1997, and on Taimyr, Siberia, in 1994 and 1995. It is concluded that the Bar-tailed Godwit is able to evaluate the probability of nest predation from Long-tailed Skuas, based on direct or indirect information about the status of local rodent populations. This information is used to choose between Long-tailed Skuas and other potential species for protection against nest predators.     

The effect of age and year on the survival of breeding adult Great Skuas Catharacta skua in Shetland

The survival rates of breeding adult Great Skuas Catharacta skua were examined at Foula, the largest colony in the world, where numbers have been declining since the late 1970s. Resightings of colour-ringed breeding adults over a 12-year period were analysed using Cormack-Jolly Seber models to estimate survival rates. Annual survival rates averaged 0.89 but varied among years between 0.82 and 0.93, with annual variations being temporally associated with variations in sandeel abundance during the breeding season. Most birds appeared to die outside the breeding season and so it is possible that nutritional stress and reproductive costs of breeding in years of poor food supply affect survivorship on migration or in the wintering range. Survival rates of adult Great Skuas were affected by their age according to a quadratic equation, with survival increasing significantly with age from 0.73 in 5-year-old-birds to between 0.85 and 0.96 in birds from 7 to 22 years old, with a sharp decline to between 0.75 and 0.87 in birds over 22 years old. Year effects were evident when controlling for age, indicating that annual variations in survival rates are not explained by changes in age-composition of the marked population among years.     

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.     

Absence from the nest due to human disturbance induces higher nest predation risk than natural recesses in Common Eiders Somateria mollissima

Human disturbance of nesting birds may cause reduced breeding success. It is therefore necessary to assess the impact of disturbance to identify steps that minimize negative impacts. We carried out a study of nesting success at two adjacent colonies of Common Eider Somateria mollissima on the islands of Grindøya and Håkøya in northern Norway between 2006 and 2011. Over the study period, nesting success was consistently higher on Håkøya (69–82%) than on Grindøya (35–60%). Between 2009 and 2011 we used camera monitoring of individual nests to identify determinants of nest survival and predation, focusing in particular on the effect of departures from the nest due to human disturbance, which differed between the colonies due to a long‐term research project on Grindøya. Overall, absence of Common Eiders from nests due to disturbance increased the predation risk by a factor of 6.42 for an increase of one additional daily disturbance. In contrast, absence due to natural recesses did not increase nest losses. Under high levels of human disturbance, camera monitoring indicated that the main cause of breeding failure was predation, primarily by Hooded Crows Corvus cornix, but also to some extent Great Black‐backed Gulls Larus marinus. The presence of cameras did not increase the predation risk. Both the presence of researchers and the sight of Common Eider females conspicuously departing from nests are likely to have provided cues to these predators. We suggest management trials to reduce nesting disturbance through the guarding of unoccupied nests to mitigate the effects of human disturbance on reproductive success.     

Carry‐over effects provide linkages across the annual cycle of a Neotropical migratory bird,the Louisiana Waterthrush Parkesia motacilla

Population limitation models of migratory birds have sought to include impacts from events across the full annual cycle. Previous work has shown that events occurring in winter result in some individuals transitioning to the breeding grounds earlier or in better physical condition than others, thereby affecting reproductive success (carry‐over effects). However, evidence for carry‐over effects from breeding to wintering grounds has been shown less often. We used feather corticosterone (CORT_f) levels of the migratory Louisiana Waterthrush Parkesia motacilla as a measure of the physiological state of birds at the time of moult on the breeding territory to investigate whether carry‐over effects provide linkages across the annual cycle of this stream‐obligate bird. We show that birds arriving on wintering grounds with lower CORT_f scores, indicating reduced energetic challenges or stressors at the time of moult, occupied higher quality territories, and that these birds then achieved a better body condition during the overwinter period. Body condition, in turn, was important in determining whether adult birds returned the following winter, with birds in better condition returning at higher rates. Together these data suggest a carry‐over effect from the breeding grounds to the wintering grounds that is further extended with respect to annual return rates. Very few other studies have linked conditions during the previous breeding season with latent effects during the subsequent overwintering period or with annual survival. This study shows that the effects of variation in energetic challenges or stressors can potentially carry over from the natal stream and accumulate over more than one life‐history period before being manifested in reduced survival. This is of particular relevance to models of population limitation in migratory birds.     

Assessing the short‐term effects of capture,handling and tagging of sandgrouse

Capturing and marking free‐living birds permits the study of important aspects of their biology but may have undesirable effects. Bird welfare should be a primary concern, so it is necessary to evaluate and minimize any adverse effects of procedures used. We assess short‐term effects associated with the capture, handling and tagging with backpack‐mounted transmitters of Pin‐tailed Pterocles alchata and Black‐bellied Pterocles orientalis Sandgrouse, steppe birds of conservation concern. There was a significantly higher mortality (15%) during the first week after capture than during the following weeks (< 2.5%) in Pin‐tailed Sandgrouse, but no significant temporal mortality pattern in Black‐bellied Sandgrouse. In Pin‐tailed Sandgrouse, mortality rate during the first week increased with increasing relative transmitter and harness weight regardless of season, and with increasing handling time during the breeding season. There were no significant differences in mortality rate between study areas, type of tag, sex or age or an effect of restraint time. These results suggest the use of lighter transmitters (< 3% of the bird's weight) and a reduction of handling time (< 20 min), particularly during the breeding season, as essential improvements in procedure to reduce the mortality risk associated with the capture, handling and tagging of these vulnerable species.     

Recent changes in colony sizes of two seabird species in Scotland are not strongly influenced by Birds Directive status of the colony

Special Protection Areas (SPAs) designated under the EU Birds Directive have improved the conservation status of many terrestrial bird species in Europe, but protecting breeding sites may be less effective for highly mobile birds such as seabirds. Colony census data for Great Skuas Stercorarius skua and Great Black-backed Gulls Larus marinus in Scotland show that breeding numbers have fared no better in sites where these species are SPA breeding features and, counter-intuitively, the evidence indicates better performance in non-SPA colonies, most likely because non-SPA colonies are generally smaller so are less subject to density-dependent competition. The main drivers of population change are widespread rather than colony-based in these two species with recent reductions in carrying capacity. Many other seabird species are vulnerable to similar widespread pressures so seabird conservation strategy needs to focus on mitigating these pressures, as designation of seabird breeding sites as SPAs is not enough to ensure effective seabird conservation.