The social organization of non-breeding Magpies Pica pica

The social behaviour of non-breeding individuals in a colour-marked population of Magpies was studied. In early autumn most non-breeders began to forage in a common area, the 'Non-breeding Flock Area'. A few individuals remained on their natal territories away from the flock area as solitary non-breeders. Most non-breeders were first-year birds, but some were second years or adults which no longer held a territory. Birds foraged in groups, their food intake rate differing with both group size and location. It appeared that the area in which birds foraged had a significant effect on food intake rate; foraging groups tended to form at sites rich in food. There was a dominance hierarchy amongst non-breeders; an individual's foraging behaviour, survivorship and chances of breeding were status dependent, with subordinates feeding less in groups and being less likely to survive and breed. Solitary individuals' chances of breeding were similar to those of high status birds, although their survivorship to breeding age may have been lower. Non-breeding Magpies are compared with non-breeders of other species and the factors which may influence their social behaviour are discussed. It is suggested that remaining as a solitary non-breeder is a viable alternative to becoming a low-status flock member for some birds.     

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.     

Spatial distribution and time budget of radio-tagged grey herons, Ardea cinerea, during the breeding season

Spatial distribution of foraging grey herons during the breeding season seemed to be determined by a trade-off between costs and benefits of area sampling versus site fidelity. Breeding birds with little experience of the area spent more time exploring than birds with more experience in the early stages of reproduction. Non-breeding herons and first-year breeders showed little site fidelity. Time budgets appeared to be affected mainly by daily changes in food availability and, for breeding birds, by demands of the brood.     

Colony and nest site fidelity of the rock shag ( Phalacrocorax magellanicus )

The rock shag (Phalacrocorax magellanicus) is an endemic species from southern Argentina and Chile. In Argentina, the population is distributed along the Patagonian coast in 143 small colonies (5–377 pairs). We studied colony and nest site fidelity of rock shags at Punta Loma, Patagonia, Argentina. We banded 37 adults from December 2001 to June 2003, during two breeding and two non-breeding seasons. Nest site fidelity was extremely high during consecutive breeding and also non-breeding seasons. The high percentage of banded birds residing in the colony in consecutive seasons suggests that rock shags are highly faithful to breeding sites and supports the idea of low post-breeding dispersion.     

SHOREBIRDS OF COASTAL BUENOS AIRES PROVINCE, ARGENTINA

The Pampas shorebird community is comprised of species from diverse taxonomic and faunal origin. Four major groups can be recognized: North American migrants residing in coastal Buenos Aires Province during their non-breeding season, Patagonian species wintering locally but breeding further south, Pampean species breeding locally, and Brazil–Paraguayan species present during their breeding season. Because of differences among these groups in the timing and migration patterns, community composition shows marked seasonal variation. Both nesting and non-breeding birds use a diversity of habitat types ranging from littoral sandy beaches to upland grasslands. Throughout the year, however, and in all habitat types, non-breeders predominate in species number and abundance. Intra- and inter-specific patterns of aggression suggest both that resources are limiting, and that non-breeding migrants may play an important role in determining breeding community composition.     

Attendance patterns and behaviour in relation to experience and pair-bond formation in the Wandering Albatross Diomedea exulans at South Georgia

Recruitment of Wandering Albatrosses Diomedea exulans to the breeding population at South Georgia took between 2 and 8 years after they first returned to their natal colony. In successive seasons, from first return to pairing, the date of arrival became earlier and the number of days spent ashore and the time spent interacting with other birds increased. Pairing birds arrived earlier and spent more time ashore than birds of similar experience which did not pair in that season. In the season following pairing they returned at the same time as breeding birds, but most did not breed; when ashore they spent much of their time alone or with their partner at the nest site. They left in mid-season before other non-breeders and bred the following season. Some birds accomplished this process by spending much time (50–60 days) ashore in two or three seasons but most birds spent a similar total time ashore spread over more seasons. Until the season prior to breeding, the number of birds of the opposite sex with whom interactions occurred was proportional to the amount of time spent ashore. There was, therefore, considerable scope for inter-individual assessment of potential partners before interactions were confined essentially to a single partner in the season before the first breeding attempt.     

Temporal variations in the feeding habits of the Purple Heron Ardea purpurea during the breeding season

The foraging strategies of adult Purple Herons Ardea purpurea from an increasing colony in northern Spain were studied in the feeding areas (nonagricultural habitats) during the breeding season (May-July). In this area, herons were mainly piscivorous, and there was no seasonal variation in prey type. Small fish (<12.5 cm) were more abundant in the diet in June and medium-sized fish (12.5–25 cm) in May and July. Foraging strategies varied seasonally: (1) More time was spent by the herons in the feeding areas in June than in May or July, whereas foraging time was only significantly greater in June than in July. (2) The biomass captured by the adults in each foraging session was similar in May and June but greater in July. (3) Feeding attempts were more numerous (but not more successful) in shallow water in June and July. (4) Aggression rates between conspecifics were higher in June than in both May and July. Probably the hatching of chicks (June) was the cause of these changes in diet and foraging strategies as well as increased intraspecies competition for foraging habitat.     

Behavioural and hormonal effects of social isolation and neophobia in a gregarious bird species, the European starling (Sturnus vulgaris)

Separating gregarious individuals from their group members often results in behavioural and physiological changes, like increased levels of corticosterone. Testosterone and corticosterone, in particular, have been implicated in the response of mammals to novelty. Data in birds are, however, rare. The presence or absence of group members may also influence an individual's response to novel stimuli. We assessed the behaviour and hormonal response of European starlings (Sturnus vulgaris) to a novel object in two different situations and seasons: each starling was tested when separated and when in contact with its group members in May/June (breeding season) and again in September/October (non-breeding season). Starlings are gregarious throughout the year, but as foraging flocks are small during the breeding season and large during the non-breeding season, we assumed that non-breeding starlings would be more affected by social isolation. Overall, starlings had higher levels of corticosterone, lost more body mass, and were more active when they were separated from their group. Isolated individuals, however, did not show a greater neophobic response than individuals in the presence of their group members in either season. Circulating levels of testosterone and corticosterone were higher after a test with novel object than after a test with only the familiar feeding dish in both sexes and seasons. However, control tests for handling effects confirmed only the increase in testosterone. Our study shows that social isolation is stressful for unrelated and unpaired members of a wild flocking bird species and demonstrates that novelty can lead to a rise in testosterone in birds.     

Life history varies with migratory distance in western sandpipers Calidris mauri

The propensity of migratory waders to remain on the non-breeding grounds during the arctic breeding season ("oversummer") in their first biological year of life ("juveniles") may be latitude, and thus migratory distance dependent. We compared the extent of preparation for northward migration of western sandpipers Calidris mauri spending the non-breeding season in México and Panamá during 1995–1998. During winter residency and premigratory periods, we measured body mass and scored the extent of dull basic versus bright alternate breeding plumage of captured juveniles and adults (second biological year or older), and obtained additional plumage scores from observations of uniquely colour banded birds. Nearly all western sandpipers in México prepared for northward migration by increasing body mass and moulting into breeding plumage. In Panamá, most adults prepared for migration, but few, if any, juveniles did so. Patterns of body mass and breeding plumage development do not generally support the hypothesis that oversummering by juveniles results directly from less efficient foraging or from resource competition with adults. We suggest instead that costs directly associated with migratory distance per se influence the life history strategies of sandpipers spending the non-breeding seasons at different latitudes. This latitudinal difference should interact with the well documented sex-ratio cline in non-breeding distribution (male western sandpipers predominating in northern parts of the range and females in southern parts). This suggests that females have more conservative life histories, prioritizing first year survivorship, relative to males that instead weight first-year breeding opportunities.     

NOTES ON THE ECOLOGY OF THE CATTLE EGRET ARDEOLA IBIS AT ROKUPR, SIERRA LEONE

SUMMARY
The Cattle Egret is a dry season visitor to Rokupr in Sierra Leone during the bird's non-breeding season. There are no cattle in the area to attract the herons and they obtain nearly all their food from small isolated areas of vegetable gardens which are associated with all the villages. The number of herons associated with each garden is closely correlated with the area of that garden and is about two birds per acre.
The daily life of the birds is discussed, in particular the pre-roosting assemblies and the flight to roost, and it is shown that each flock leaves for the roost at the same time each day, about 30–35 min. before dusk.
Observations made at a particular roost showed that all the Cattle Egrets reached the roost before dusk and from these observations, combined with those made on the feeding grounds, it is concluded that the number of Cattle Egrets using the roost is determined by the number and area of the feeding grounds that lie within 30–35 minutes flying time (about 12 miles) of the roost.
Other species using the roost are briefly discussed.     

THE EFFECT OF AGE ON THE BREEDING BIOLOGY OF THE KITTTWAKE RISSA TRIDACTYLA

1. From 1954 to 1956 inclusive, the biology of individual marked Kittiwakes was studied at North Shields, Northumberland.
2. It was concluded that older Kittiwakes reacted to the breeding stimulus earlier, more intensively and with greater success than younger breeding birds.
3. Birds with previous breeding experience returned to the colony before birds breeding for the first time and these before non-breeders.
4. Before breeding started, birds which had bred previously spent more of their time at the colony than those about to breed for the first time.
5. Birds breeding for at least the second time laid the first egg 7·5 days earlier than those breeding for the first time.
6. Breeding started one day later for every four days the return to the colony was postponed.
7. Older breeding birds showed greater nest-site tenacity, laid larger clutches and had greater breeding success than younger birds.
8. The chicks in broods of two (but not of one) increased in weight more rapidly where the parents had previous experience.
9. Breeding Kittiwakes showed strong colony tenacity, but 24% of the marked non-breeding birds were subsequently seen in other colonies.
10. Over half the birds retained the same mate as in the previous year.     

THE PELAGIC DISTRIBUTION OF THE SOOTY SHEARWATER PROCELLARIA GRISEA

The breeding distribution of the Sooty Shearwater and the dates of its breeding season are briefly reviewed.
Records from the Southern Ocean are summarised. It is concluded that they are accounted for by non-breeding birds feeding off the ice-edge in the Australian sector of the Antarctic during the latter part of the breeding season.
The great majority of Sooty Shearwaters spend the southern winter in the Pacific Ocean. Birds from the New Zealand colonies are probably mostly found on the feeding grounds off Japan: some may make a "circular" migration around the Pacific. Birds from the Cape Horn colonies spend the southern winter in the rich areas off the western coast of North America.
Some Sooty Shearwaters spend the southern winter off the South African coasts. A number of non-breeding birds remain there throughout the summer.
The Sooty Shearwaters "wintering" in the North Atlantic are probably numbered in tens of thousands only. The distribution of records is reviewed month by month and the records from Atlantic transects are summarised. The migrations in the North Atlantic are discussed in relation to prevailing meteorological and oceanographic conditions, and food supply. It is concluded that most Sooty Shearwaters in the North Atlantic make a "circular" migration, crossing the North Atlantic from west to east in June and July, and re-crossing about September further south in the northeast trades.     

ECOLOGY OF THE COMMON BABBLER TURDOIDES CAUDATUS

Some aspects of the ecology and behaviour of the Common Babbler are described. The species resembles most other members of the genus Turdoides in being group territorial and exhibiting cooperative breeding. Females remain with their natal group until 6–9 months old and then disperse. Males, in most cases, remain indefinitely in their natal group so that these essentially constitute male clans. In half of the groups observed only a single pair bred per season, but in some large groups two pairs attempted to breed, one always being unsuccessful, and in other cases one male bred successively with two different females. Overall breeding success was highest for groups in which successive polygyny occurred. Male non-breeders fed the nestlings more often than female non-breeders which were sometimes driven off by the breeding female. A positive correlation was found between the number of non-breeders feeding the nestlings at day 9–11 and the size of the brood. This probably did not indicate a causal relationship. Breeders in large groups continued to moult while breeding, while those in small groups arrested their primary moult. This difference suggests that assistance from non-breeders may have had some effect in reducing the physical strain on the breeding pair. Observed overall survival of adult birds was 63% per annum, but that of the breeding males was 88% per annum. The population declined by 19% during the period of the study, suggesting that observed rates of survival may have been lower than normal.     

Attendance off breeders and prospectors reflects the quality off colonies in the Kittiwake Rissa tridactyla

Attendance of breeders and prospectors (prebreeders, failed breeders and non-breeding adults) was studied from the time of hatching to the end of the breeding season in parts of two neighbouring Kittiwake Rissa tridactyla colonies in Brittany (France). Many clutches in one subcolony had been predated by a Carrion Crow Corvus corone , leading to a lower breeding success. Attendance of breeders and prospecting activity were greater in the more successful colony, and the number of breeding pairs increased in the following year in this colony only. Local reproductive success and social behaviour contributed to the attraction exerted by a colony. The demographic trend was explained by a performance-based conspecific attraction mechanism, acting through the effect of local reproductive success on recruitment, non-breeding and fidelity. Study of breeding success and prospecting activity allowed both evaluation of the quality of a colony and prediction of local demographic trends for the following year.     

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.     

Pair territoriality in the longnose filefish,oxymonacanthus longirostris

The longnose filefish,Oxymonacanthus longirostris, usually lives in heterosexual pairs, the male and female swimming together and sharing the same territory. Pair territoriality in the species was examined in detail in relation to sexual differences in territorial defense activities. Rigorous pair territoriality was maintained only during the breeding season, although pairs used their home ranges exclusively to a certain extent, during the non-breeding season. The frequency of aggression against other conspecific pairs in the breeding season was higher than in the non-breeding season. Agonistic interactions appear to be over both mates and food resources, the strict pair territoriality in the breeding season possibly being due to mutual mate guarding. In intraspecific aggressive interactions, males usually led their partner females when attacking intruders. The feeding frequency of males was much lower than that of females in the breeding season. Mate removal experiments indicated that females could not defend their original territories solitarily and their feeding frequency decreased. Conversely, males could defend territories solitarily without a decrease in feeding frequency. These results suggest that males contribute most to the defense of the pair territory, with females benefiting from territorial pair-swimming with their partner males.     

Seasonal changes in testosterone and corticosterone levels in four social classes of a desert dwelling sociable rodent

Animals have to adjust their physiology to seasonal changes, in response to variation in food availability, social tactics and reproduction. I compared basal corticosterone and testosterone levels in free ranging striped mouse from a desert habitat, comparing between the sexes, breeding and philopatric non-breeding individuals, and between the breeding and the non-breeding season. I expected differences between breeders and non-breeders and between seasons with high and low food availability. Basal serum corticosterone was measured from 132 different individuals and serum testosterone from 176 different individuals of free living striped mice. Corticosterone and testosterone levels were independent of age, body weight and not influenced by carrying a transmitter. The levels of corticosterone and testosterone declined by approximately 50% from the breeding to the non-breeding season in breeding females as well as non-breeding males and females. In contrast, breeding males showed much lower corticosterone levels during the breeding season than all other classes, and were the only class that showed an increase of corticosterone from the breeding to the non-breeding season. As a result, breeding males had similar corticosterone levels as other social classes during the non-breeding season. During the breeding season, breeding males had much higher testosterone levels than other classes, which decreased significantly from the breeding to the non-breeding season. My results support the prediction that corticosterone decreases during periods of low food abundance. Variation in the pattern of hormonal secretion in striped mice might assist them to cope with seasonal changes in energy demand in a desert habitat.     

A RINGING STUDY OF MIGRATORY BARN SWALLOWS IN WEST MALAYSIA

The Barn Swallow is a non-breeding winter visitor to West Malaysia (Malaya), abundant in season, by day feeding aerially over a wide range of habitats and by night normally roosting gregariously in trees, reed-beds or on service wires in towns. Records of ringed birds have demonstrated that those reaching Malaya breed in the Palaearctic region from 108°E eastwards between 37° and 51°N. Recoveries south of the breeding range suggested that migrating birds may follow either a continental route or a more easterly track through the Philippines and Borneo. Counts at roost sites in a reed-bed and in towns demonstrated a seasonal increase in numbers from late July to a peak in November, followed by a decline of about 20% to a level maintained until mid-February when departure commenced. Most birds had left by early May, but a few lingered and possibly overlapped with the first returning migrants in June. There was no evidence that any individuals remained in Malaya through the nuptial period. Repeats during winter at three regularly sampled urban roosts indicated that many birds on passage were present until November and again in late March–early April; from December to February the winter population was relatively stable and comparatively sedentary. Although the distances between towns were small in relation to the demonstrated foraging range of Barn Swallows, only 17% of 1,955 repeats of ringed birds represented a shift in roost site. Most shifts were towards the centrally situated and most populous roost of the three; interchanges between the outer pair of towns were few. A complete moult occurred on the wintering grounds, during which young of the year acquired adult plumage. Replacement of the primaries extended virtually throughout the moulting period, at an average rate of 2.4 feathers per month in the proximal part of the tract and 1.3 feathers per month in the distal part. Adults on average moulted slightly earlier than juveniles, but there was a wide scatter in timing between individuals of both age groups. There was no evidence that the initiation of moult was related to the dates of post-nuptial migration. The date of departure on prenuptial migration, however, was normally delayed until primary moult was complete. Large weight gains in March and April occurred only in swallows which had completed the moult. At this period the mean weight of birds in fresh plumage was about 30% above the lowest winter mean, and was significantly higher than that of contemporary samples of birds in which moult was continuing. In final samples in late April and early May mean weights showed a decline, indicating that late birds departed with reduced deposits of metabolic reserves. The gonads of adults of both sexes among passage and arriving birds in July and August had largely completed post-nuptial regeneration, and subsequently remained quiescent. Preliminary stages of recrudescence were observed in females from February onwards, and in males from March. Recrudescence was most advance in specimens which had completed the moult, but did not approach breeding condition in any bird before departure. Returning birds tended to be conservative in their choice of winter roost. Among 1,276 records, 82% were recaptured in the town of original ringing. Again shifts towards the centrally situated roost were more numerous than between the peripheral pair. The frequency of returns varied significantly with the month of ringing, being higher for December-March, lower for July-November and April-May. Survival rates, calculated from returns after one and two breeding seasons, indicated an annual mortality of 60–72%, higher among juveniles than adults. Comparison of results of successive years suggested that unfavourable conditions in 1967 resulted in lower survival of juveniles in particular than in 1966. There was no evidence of mortality at the roost sites, and it is argued that heavy losses probably occur during the migratory journeys.     

Phenology of migration and use of wintering sites by the increasing population of dark-bellied brent geese Branta bernicla bernicla

During the recovery of the population of dark-bellied brent geese Branta bernicla bernicla , traditional wintering sites in Britain have experienced increases in numbers and new sites have been colonized. Goose counts were used to describe the migration phenology and winter use of Scolt Head, north Norfolk (a traditional site) and the Burry Inlet, South Wales (a relatively new site) over the period of re-colonization and colonization, respectively. During the 1950s, only a few hundred birds wintered at Scolt Head, and most stayed for only a short period. By the late 1980s/early 1990s, numbers had reached several thousands, their period of stay was longer and there was little variation between years in median arrival (28 October) and departure (11 March) dates. A similar process was observed during the entire colonization phase at the Burry Inlet; initially birds arrived late and departed early, but arrived progressively earlier and departed later as wintering numbers increased. The autumn arrival rates at both sites were slower than the spring departure rates. During years of good breeding productivity, those adults with young arrived slightly later in Norfolk than failed breeders and non-breeders, and a higher proportion of young birds remained in spring after the main departure. However, the overall pattern of arrival and departure did not vary according to whether there had been a good or poor breeding season. Some adults of breeding age also remained until May. The effects of food depletion and use of novel feeding habitats in relation to migration phenology and winter use of sites are discussed.     

Weak migratory connectivity,loop migration and multiple non-breeding site use in British breeding Whinchats Saxicola rubetra

Determining the links between breeding populations and the pressures, threats and conditions they experience presents a challenge for the conservation of migratory birds which can use multiple sites separated by hundreds to thousands of kilometres. Furthermore, migratory connectivity – the connections made by migrating individuals between networks of breeding and non-breeding sites – has important implications for population dynamics. The Whinchat Saxicola rubetra is declining across its range, and tracking data from a single African non-breeding site implies high migratory spread. We used geolocators to describe the migration routes and non-breeding areas of 20 Whinchats from three British breeding populations. As expected, migratory spread was high, with birds from the three populations overlapping across a wide area of West Africa. On average, in non-breeding areas, British breeding Whinchats were located 652 km apart from one another, with some likely to share non-breeding areas with individuals from breeding populations as far east as Russia. Four males made a direct non-breeding season movement to a second, more westerly, non-breeding location in January. Autumn migration was through Iberia and around the western edge of the Sahara Desert, whereas spring migration was more direct, indicating an anticlockwise loop migration. Weak migratory connectivity implies that Whinchat populations are somewhat buffered against local changes in non-breeding conditions. If non-breeding season processes have played a role in the species’ decline, then large-scale drivers are likely to be the cause, although processes operating on migration, or interactions between breeding and non-breeding processes, cannot be ruled out.