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 influence of pair-status on the breeding behaviour of the Kittiwake Rissa tridactyla before egg-laying

The breeding behaviour of individually marked Kittiwakes Rissa tridactyla that retained mates from the previous year (SAME) was compared, over the period from pair-formation to egg-laying, with those that changed mates (CHANGE). Position of nest-site in colony and breeding experience did not differ in the two groups studied. Hirds in CHANGE pairs attended the nest-site more frequently and as a result probably had less 'off-duty' time during which to forage. Rates of greeting were higher in CHANGE pairs. Differences between the two groups were usually greatest during the first two weeks after pair-formation an decreased toward egg-laying. CHANGE pairs copulated over a longer period before egg- laying, but stopped copulating sooner than SAME pairs. Copulation rates were higher in SAME pairs immediately before egg-laying. The causation of the behavioural differences between the two groups is discussed in terms of mate familiarity. It is suggested that the recorded differences explain the lower reproductive success in CHANGE pairs reported by other workers.     

Use of cloacal sexing techniques in mark-recapture estimates of breeding population size in Wilson's stormpetrel Oceanites oceanicus at South Georgia

Summary Many stormpetrel species breed in habitats where their populations cannot be estimated by direct counts of burrows or birds; mark-recapture experiments have been confounded by the presence of many wandering non-breeders. With a population of Wilson's Stormpetrel Oceanites oceanicus at Bird Island, South Georgia, we tried to estimate the proportion of breeding females in samples obtained during a mark-recapture experiment. These were identified by measurements of the cloaca, which greatly enlarges at egg-laying. A concurrent experiment with individually marked birds determined that breeding females could be discriminated from males and non-breeders for c 30 days after laying. The technique is probably applicable to other petrels, though it will work best with those that lay most synchronously. The overall population estimate was 4841–5515 birds (SE 856–1417); estimates of breeding females gave a population of 2300 paris early in the incubation period and 1400 pairs near hatching.     

FACTORS AFFECTING THE AGE OF FIRST BREEDING OF THE KITTIWAKE RISSA TRIDACTYLA

At a Kittiwake colony in Northumberland, 80% of those birds which returned to their natal colony to breed were males and these supplied 52% of all male recruits. More females breed away from their natal colony than males. There was no differences in the proportions of young fledged from sites in the centre or at the edge of the colony, or by parents of different experience, which returned to breed. Kittiwakes breed for the first time at ages from 3 to 8 years, but most at 4 or S years old. Males arrive back at the colony at an earlier age than females and breed for the first time one year earlier. Males obtaining sites at the centre of the colony first breed at an earlier age than those at the edges. Neither the age nor the area of first breeding appear to be transmitted from parent to offspring. Males breeding first aged 4 years or younger produced more young than those which first bred aged 5 years or older, despite their partners laying smaller clutches. This difference was most marked among those males recruited to sites in the centre of the colony. The advantage of this earlier breeding is counteracted by a lower survival rate among those males which start to breed at the younger ages. In all breeding Kittiwakes, annual reproductive output increases with experience while annual survival rates decrease. Once they had started to breed, many birds failed to breed in one subsequent season. Nearly 60% of these cases of intermittent breeding occurred in the year following first breeding. Intermittent breeding was most frequent among young birds and among females. It is suggested that each breeding involves a cost to the individual in terms of reduced survival, and that deferred and intermittent breeding are means of guarding survival. A model is proposed whereby the age at which a bird starts to breed, the nesting site which it obtains, and its subsequent breeding strategy result in each individual producing an optimal number of reproducing offspring in its lifetime, relative to its quality.     

BREEDING BIOLOGY OF THE MANX SHEARWATER PUFFINUS PUFFINUS

Studies on the breeding biology of Puffinus puffinus were carried out in 1963 and 1964 at the large colony on Skokholm, Wales. During the six weeks before laying the birds spent up to a quarter of the days in the burrows, but the ten days immediately prior to laying were normally spent at sea. There is a prolonged laying period, with a marked peak in the first half of May. Details are given of a second egg being laid when the first was deserted immediately after being laid. The male took the first incubation spell. The incubation spells ranged from one to 26 days and averaged six. The incubation period was about 51 days. The frequency of visits to land by breeding birds, unlike those by non-breeders, was not affected by the moon. On hatching, the chicks grew rapidly and reached maximum weights of between 505 and 755 gm. sometime between 39 and 61 days. There was a variable desertion period, usually eight or nine days, before the chicks left the island about 70 days after hatching. During the feeding period the chick received about two feeds every three days. There is evidence that adults visited the chicks more frequently than this. There was no correlation between growth of the chicks, their feeding rates or fledging weights and the time of laying. There was a high survival (about 95 %) of chicks during the fledging period but some eggs were lost in disputes for burrows. Nine pairs in 1964 were unable to raise two young simultaneously. Parents altered their feeding rhythms to try to feed two young but did not themselves lose weight. It is suggested that the critical factor in the production of young is the availability of food for the young immediately after they leave the colonies.     

Dynamics of food availability, body condition and physiological stress response in breeding Black-legged Kittiwakes

1. The seasonal dynamics of body condition (BC), circulating corticosterone levels (baseline, BL) and the adrenocortical response to acute stress (SR) were examined in long-lived Black-legged Kittiwakes, Rissa tridactyla , breeding at Duck (food-poor colony) and Gull (food-rich colony) Islands in lower Cook Inlet, Alaska. It was tested whether the dynamics of corticosterone levels reflect a seasonal change in bird physiological condition due to reproduction and/or variation in foraging conditions.
2. BC declined seasonally, and the decline was more pronounced in birds at the food-poor colony. BL and SR levels of corticosterone rose steadily through the reproductive season, and BL levels were significantly higher in birds on Duck Island compared with those on Gull Island. During the egg-laying and chick-rearing stages, birds had lower SR on Duck Island than on Gull Island.
3. The results suggest that, in addition to a seasonal change in bird physiology during reproduction, local ecological factors such as food availability affect circulating levels of corticosterone and adrenal response to acute stress.     

THE BREEDING BIOLOGY OF THE GANNET SULA BASSANA ON THE BASS ROCK, SCOTLAND

The Bass colony is increasing—in 1962 there were 5,350–5,700 breeding pairs; 1,340–1,430 pairs of non-breeders with nests or sites (mainly pairs in their season before first breeding) and 2,000–2,500 “club” birds without nest or site. 15% of nests were occupied by both birds of a pair at the time of the count. Oldest males return to the colony in January, followed by experienced females, considerably later by young adult-plumaged birds, immature birds later still, and the few one year-olds that return usually not until May and June. Mid-cliff sites are the first to be re-colonized each year. Gannets usually breed in their fifth year and there is some evidence that females breed earlier than males. The characteristics of Gannet nests and sites are described. Nests function in raising the egg and young above the morass of the breeding colony, and reach a density of about 2.3 per square metre. Nests are demolished and their positions changed more often than might be suspected. The extremely strong social tendency which causes Gannets to establish their sites amongst or very close to existing breeders probably is the factor ensuring high density and this contributes to synchronization of laying. Egg laying is analysed. Experienced pairs forming an isolated group of 20 nests began laying later and showed less synchronization than two other groups of the same size but from the middle of a dense mass, probably due to the greater social stimulation experienced by the non-isolated groups. The date for first and median eggs was also earlier in larger than smaller groups in the same year. The effect of density as distinct from group size is also discussed. Early eggs are mainly laid on cliff or cliff-edge sites and in large nests. Different groups within the colony produced the median egg within 2–3 days of the end of April each year. In the fullest documented group the mean date was also constant from year to year and closely approached the median, implying a considerable degree of synchronization within the gannetry as a whole. Laying in the observation colony became progressively earlier in successive years, probably due to recovery from previous disturbance. Nevertheless, individual females tended to lay in a fixed position each year with relation to the mean for the group. Increasing age of the female causes earlier egg laying and heavier eggs for up to at least five years. It is suggested that the survival value of seasonally synchronized laying in the Gannet is maximum utilization of a seasonally dependable and abundant food supply for the production of young with the optimal chance of post-fledging survival. The spread of laying acts as an insurance against possible adverse conditions. There is a considerable reserve of unutilized breeding capability within the colony (adult non-breeders, a pre-maturity period longer than physiologically necessary for egg production, and a one-egg clutch when in fact two young can easily be reared). The mean of 393 Gannet eggs was 104.5 gm. (range 81–130). Eggs constitute about 3.4% of adult female weight and lose 9–13% in weight during incubation. Replacement laying of the invariable one-egg clutch takes 6–32 days. The mean incubation period was 43.6 days. Male incubation spells averaged 35.6 hours; female's 32.0 hours. Copulation ceases immediately after egg laying. During three seasons, 82% of eggs laid in the observation colony hatched. Inexperienced pairs hatched 62.6% of eggs laid; experienced pairs hatched 86%. Some of the processes of incubation and chick rearing depend on the maturation of innate abilities and not on experience Inexperienced breeders do not seem inferior to experienced ones in finding enough food for their young. Parental care of the new chick is described; the pterylosis of the chick is figured. A summarized account of plumage development is given. Food fish and chick feeding are described. The average frequency of feeds throughout a continuous two-day watch was 2.7 feeding bouts per chick per day. Adult fishing trips usually took 7–13 hours and the estimated fishing range is over 100 miles, and possibly up to 400 miles, from the breeding colony. Despite this, 15% of daylight hours are spent by the pair together at the nest in addition to the constant guarding of the chick by one or other. Gannet young have a very compressed growth period compared with boobies and fledge at 3,100–4,100 gm., after a steady growth uninterrupted by periods of starvation or arrested development after an average 90 days. 92.3% of all eggs which hatched in 500 nests in the observation colony during three years of the study gave fledged young. Excluding inexperienced birds, there was no difference in the fledging success of eggs laid at different dates in the breeding season, in accordance with the proved abund- ance of food. However, post-fledging survival is probably higher among young fledging at the peak period (first half of September) than later and the few relevant ringing returns tend to support this. Breeding success at the small colony at Bempton was less compared with groups from the Bass for the years 1961–3. Causes of chick loss before and during fledging are discussed. They are unimportant compared with the great mortality in the first year of life after fledging. The adaptive significance of black plumage in the juvenile Gannet probably lies in reduced attack-releasing qualities of such plumage on the male parent. The Gannet alone in the Sulidae produces young which leave the nest with large fat deposits, and which are not fed at all by the parents after fledging. This is possibly another result of adaptation to using a seasonally abundant food supply to the maximum. The present Gannet population increase is discussed in relation to cessation of human predation, the possible impetus given by a temporary but large increase in pelagic fish during the war, but also the overall steady downward trend in fishing returns since the early part of this century. One cannot explain the steady and considerable rise in Gannet numbers only in terms of increased food supply. The fact that, at a time of population expansion and obviously favourable conditions, Gannets are still far from utilizing their full recruiting powers, needs investigating further. It may be partly due to the relative slowness of evolutionary change in a long-lived species with slow population turn-over, if the Gannet has evolved its characteristics in response to an environment different from the present one.     

Patterns of colony attendance in the Cape Petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica

The annual cycle of colony attendance of the Cape Petrel Daption capense at Nelson Island, Antarctica, covered about 9 months from the start of September until the end of May. Large numbers of birds attended the colony continuously for 6 months from arrival until fledging, except for a 2-week incomplete pre-laying exodus when attendance was reduced to 10% of that during incubation. After 1 month (March) of post-fledging absence, birds returned to the colony for 2 months in numbers comparable to the level in the breeding season. This was followed by a winter absence that lasted only 3 months and was interrupted by two short visits by a small number of birds. Mean attendance was highest during incubation, but short peaks of maximum attendance occurred during the pre-breeding period. Except for the incubation and brooding periods, colony attendance showed high variability both within and between days. Large numbers of visiting birds occurred on the sea and on islets in front of the colony until the end of October. Unemployed birds constituted an almost constant proportion of up to 50% of colony-attending birds during the incubation period. Diurnal variability did not show a consistent pattern but may represent preferential feeding during early morning hours, with additional short departures depending on the length of the daylight period. Colony counts were the most consistent during the second half of incubation, when the lowest counts gave an estimate of the number of breeding pairs and peak numbers an impression of the additional number of attending nonbreeders. Pre-breeding counts revealed the maximum numbers of birds associated with a particular colony. However, for both pre- and post-breeding surveys, several counts a day over a period of at least a week are recommended because of variability in attendance.     

Prerequisites for recruitment of Kittiwakes Rissa tridactyla

Observations at the North Shields Kittiwake colony between 1982 and 1984 showed over 100 potential recruits (birds of breeding age) associated with the colony, only about 40% of which were actually recruited. In addition, there were younger birds which visited the colony. The origins, age structure and intercolony movements of members of the pool of prospectors have been examined. Porter & Coulson (1987) demonstrated that recruits are heavier than prospectors, implying that they are in better body condition. Three further prerequisites for breeding in Kittiwakes are recognized: the age of recruits is at least three years; all birds which recruit are present at the colony at least one year prior to breeding; and recruits show a high degree of attachment to colony as illustrated by their colony and nest site tenacity, attendance and early arrival. The results suggest that there is considerable selection of individuals at the time of recruitment; the mechanism for this has yet to be examined.     

The establishment and growth of a new Fulmar colony on sand dunes

The Fulmar continues to extend its breeding range in Britain. This paper records the growth of a new Scottish colony. atypically on sand dunes, from initial prospecting to eventual breeding. It also examines the danger of roofless enclosures to prospecting birds.     

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.     

Aspects of the biology of Glaciopsyllus antarcticus (Siphonaptera: Ceratophyllidae) during the breeding season of a host (Fulmarus glacialoides)

Summary The breeding period of the Antarctic flea, Glaciopsyllus antarcticus (Smit and Dunnet), was synchronised with the breeding period of the host, Southern Fulmar (Fulmarus glacialoides Smith). Although eggs were laid in the host nest, larvae developed amongst the down (particularly on the belly) of host chicks. Larvae were blood feeders and pupated amongst the down of host chicks. The development of pupae was arrested by ambient temperatures (mean temperature of +2.5°C in January), but recommenced when pupae were warmed. Female fleas comprised 55.8% of a collection of 1988 adults. Low numbers of adult fleas were found in nests prior to host breeding and subsequent to host fledging in comparison to numbers on the host; adults are therefore presumed to overwinter on the host, remote from the nest.     

Movements of radio-tagged Grey Herons Ardea cinerea during the breeding season in a large pond area

This paper describes the strategies of resource utilization in the course of the breeding season by five radio-tagged Grey Herons Ardea cinerea. The seasonal changes in exploitation of the environment by two breeding adults, one non-breeding adult and two non-breeding first-year birds were studied from March to August 1982, near Zonhoven in Belgium. Two adult breeding birds could be followed continuously from the end of March until the middle of June. During the first month they explored an extended area all around the colony, but each concentrated its search in a specific direction. From the end of April until the beginning of June, most probably from egg-hatching until the end of breeding activities, each bird spent a very large proportion of its time at a particular feeding site, from which other herons were actively excluded. In the first part of June they again visited different sites, each maintaining its preferred direction. From the middle of June onwards they seemed to have left the fish-pond area. The pattern of movements of the first-year birds differed markedly from that of the breeding adults. In April, although both non-breeding and breeding birds explored large areas, only the areas used by non-breeders were centred on the colony. From the end of April onwards, probably after general egg-hatching in the colony, the non-breeders very rarely revisited the colony, and from May till August their ranges became more and more restricted to very small areas at an increasing distance from the colony. They were never observed defending particular sites. The results are discussed with regard to recent speculations about the evolution of colonies as an adaptation for the exploitation of food resources. Breeding herons seem to explore a large part of the environment during incubation and defend a particular site while feeding their young. Choice of feeding site by non-breeding birds may be influenced by the site defence of the breeding birds. Non-breeding birds exploit a large area when breeding birds occupy feeding territories. Perhaps they are forced to forage in less suitable places at this time. Colonies might have evolved as a strategy to minimize effort in resource esploitation as, especially at the beginning of the breeding season, the colony could act as an assembly point in the exploration of the environment. However, its importance as an assembly point diminishes in the course of the season, as non-breeding birds no longer visit the colony and adults defend territories.     

Nest site characteristics, duration of use and breeding success in the Guillemot Uria aalge

The fate of all eggs laid by Guillemots Uria aalge in seven study areas on the Isle of May was recorded in 12 consecutive seasons. The average success at each site was related to its physical characteristics, bird density, position on the cliff and availability of suitable habitat for ticks. The numbers of neighbours and walls, type of site, slope where the egg was incubated and distance from the top of the cliff all had a significant effect on both hatching and breeding success. Potential tick habitat had no significant effect. Much variation in hatching and breeding success remained unexplained. Sites used when the colony was less than half its current size and most frequently in the period 1984–1995 were the most successful. Since an average Guillemot pair remains together at a site for only 3–4 years. the consistently high success of many sites over a period of 10–12 years suggests that the most used sites were occupied by a succession of high-quality birds.     

Breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in high-arctic northeastern Greenland, 1993

The breeding ecology of the Fulmar Fulmarus glacialis and the Kittiwake Rissa tridactyla in the high Arctic was studied in relation to the occurrence of the northeast water polynya in northeasternmost Greenland (80̀N). Mean laying dates were 31 May in the Fulmar and 18 June in the Kittiwake; the total nesting season for the Fulmar just matched the time window of the polynya opening period. Fulmar colony attendance fluctuated within a period of 11.6 days because of variation in nonbreeding prospectors but showed no clear diurnal variation. Fulmar incubation shifts, on average, lasted 6.1 days (range 1–13 days), which is significantly longer than elsewhere, and the average chick-guard period of 10.9 days (range 1–17 days) was significantly shorter than in other studies. Egg neglect occurred in 18% of Fulmar nests or 0.7% of nests per day. Overall breeding success (chicks fledged per egg laid) was 0.56 in the Fulmar and 0.67 in the Kittiwake; the latter produced 1.4 young per active nest or 1.2 per completed nest. Mean Kittiwake clutch size was 2.03; larger clutches were laid early. Nest site characteristics (presumably reflecting nest predation risk) and breeding behaviour affected breeding success. in the Fulmar, hatching success was negatively correlated with laying date and the proportion of egg neglect, while overall breeding success was correlated negatively with distance to nearest neighbouring site and positively with the length of the chick-guard period. Kittiwake breeding success was negatively correlated with laying date. Using seabirds as indicators of marine food supply, breeding success in both species suggested moderate to good food supply in the northeast water polynya in 1993, although at least in the Fulmar the high reproductive output appeared partly maintained by behavioural buffering; long incubation shifts, egg neglect and short chick-guard periods were symptoms of foraging constraints.     

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.     

Negative indirect effects of an avian insectivore on the fruit set of an insect-pollinated herb

Though an abundance of research has focused on direct interactions between birds and plants, relatively few studies have reported on indirect interactions. Of those reports, all have focused on positive indirect effects of birds on plants through predation of plant natural enemies. We conducted an observational study along the Middle Rio Grande in New Mexico to determine if avian aerial insectivores had a negative, indirect impact on insect-pollinated plants through predation of insect pollinators. We found considerable taxonomic overlap, at the order and family level, between insects visiting sweet clover ( Melilotus officinalis ) and those eaten by cliff swallows ( Hirundo pyrrhonota ). We found a significant negative relationship between proximity of sweet clover to cliff swallow breeding colonies and sweet clover fruit set during the cliff swallow nestling period. The apparent effect of cliff swallows was strongest within 200 m of breeding colonies (approximately 50% reduction in fruit set) and decreased nonlinearly to a distance of approximately 400 m. Finally, we found that the clover fruit set gradient disappeared after the nestling period, when chicks had fledged and the colony was abandoned.     

Seasonal and Diurnal Changes in Aggression and Territory Size in the Black-headed Gull (Larus ridibundus L.) on Islands in the Middle Reaches of the Vistula River

Seasonal and diurnal changes in territory size and frequency of aggressive interactions of various intensity were examined in colonies of the black-headed gull (Larus ridibundus) on river islands and ponds of central Poland. The highest frequency of the total aggression and the largest territories were recorded before the egg-laying stage, this being related to the establishing of territories, defence of the mate, and defence of nest material. Later in the season both aggression and territory size declined. Unlike the large species of gulls, black-headed gulls did not increase their aggression and territory size during hatching of the young. High-intensity aggressive behaviours (choking, attacking and fighting) were not so frequently displayed as low-intensity aggressive postures (upright, long call, forward) in each phase of the breeding cycle. Presumably this was because the former were more costly in terms of energy. Peak territorial activity occurred in the late afternoon. An increase in the frequency of intense aggressive behaviours (attack, fight) at that time was combined with an increased mobility of birds (departures to and arrivals from foraging sites). A reduced frequency of high-intensity aggressive displays in the period when the largest number of birds was present in the colony was likely to be adaptive.     

BREEDING BIOLOGY AND BEHAVIOUR OF THE GREY PHALAROPE PHALAROPUS FULCARIUS IN EAST SIBERIA

Studies were made in 1970 in the Chukotski Peninsula, in 1971 in the delta of the Indigirka river and in 1972 in the delta of the Yana river. Grey Phalaropes inhabit polygonal and tussocky moss-sedge tundra rich in swamps, lakes and (in June) temporary ponds. Population density in favourable habitats may reach 1–2 pairs ha^-1. Data on breeding chronology are presented, and various aggressive and courtship displays described. Most phalaropes seem to keep within a home range, sometimes large, during courtship time, but no defended territories and no forms of territorial behaviour exist. Many birds, both local nesters and wanderers, can feed on any pond. Sexual dimorphism is described. In 1970, three non-breeding cock-plumaged females were taken. Pairs are formed both before arrival and on the nesting grounds. All courtship displays are wholly or mostly initiated by females. In 1971, in the Indigirka delta, all the Grey Phalaropes were paired by 12 June, and stayed in pairs until the end of egg-laying. In 1970, on the northern Chukotsk, phalaropes seemed to form no (or very few) permanent pairs. Throughout June, most birds occurred in mixed flocks constantly moving between lakes or ponds. Copulation seemed to be promiscuous within the local population; polyandry cannot be excluded in some cases. Pairs appeared to be created only for the time of egglaying; probably, the only biological role of pair-formation is to find a male for incubating. Thus, a definite social system is not a species-specific feature; it can vary depending on local and yearly situations, including probably sex ratio. Nests are usually situated in very wet places, sometimes on the water edge. They can be found as little as 3 m apart, but are usually 40–80 m apart, or further. Incubation begins after the second or third egg. After the end of egg-laying, males drive females away from the nests, and pairs break up. Females and non-breeders gather in flocks and move onto the lakes of maritime tundra, and later on to the sea. The composition of the flocks is not constant: they often join together or part. Brooding males feed near their nests, sometimes in groups; not unfrequently they join flocks of females and non-breeders for some time. The normal average clutch-size is c. 4 eggs; when nesting was delayed (in the central Indigirka delta in 1971) the average was 3–61. The loss of nests was great in 1971; numbers of young on 1–3 August was 10 times lower than adult numbers in June.     

山西省代县小鸊鷉繁殖生物学的研究

１９８５—１９９４年的３—１１月,在山西省代县对小鸊鷉的繁殖生物学进行了研究。该鸟在山西省为夏候鸟,每年最早３月６日迁来,最晚１１月２４日迁离,居留期为２５５天左右。种群密度在繁殖前的４月为１．３７,繁殖后的９月为１．９５（只／ｈａ）。产卵始期为５月２７日,日产１枚,窝卵数５—８枚,产出第二枚卵时开始孵卵。观察到最早孵卵期为５月２８日,孵卵期２０－２２ｄ。雏鸟出壳先后顺序与产卵先后顺序相一致,孵化率为９５．４５％,雏鸟为早成鸟。