Reproductive biology and ecology of white‐winged trumpeters (Psophia leucoptera) and recommendations for the breeding of captive trumpeters

The reproductive biology and ecology of a wild population of white‐winged trumpeters (Psophia leucoptera) were studied in southeastern Peru from 1983 to 1987. Because little information is available about any of the trumpeter species and because trumpeters have proven difficult to breed in captivity, information relevant to breeding and management of captive trumpeters is reported in this paper. White‐winged trumpeters lived in territorial social groups that ranged in size from four to 13 individuals. A typical territorial group contained three adult males, two adult females, and several sexually immature offspring, but smaller temporary groups sometimes formed for the duration of the breeding season. Only the dominant female contributed eggs to the clutch, and all adult males in the group competed to obtain copulations with her. Eggs were laid in elevated nesting cavities and no nest was constructed. The average clutch size was three eggs and incubation was not begun until the final egg was laid. The dominant male and female shared most of the incubation duties, but subordinate males covered approximately 15% of the incubation shifts. Eggs hatched approximately 27 days after incubation was begun and chicks left the nesting cavity the day after they hatched. Chicks were completely dependent on older birds to feed them for their first 3 weeks and then gradually began to feed themselves more and more food. The subordinate adult males fed chicks the most food, the dominant male and female and older offspring fed chicks an intermediate amount, and the subordinate adult female fed chicks the least. Young chicks behaved aggressively toward each other but were separated by adults before they injured each other. If at least one chick from the clutch survived, trumpeters did not breed again until the beginning of the next breeding season the following year. Zoo Biol 19:65–84, 2000. © 2000 Wiley‐Liss, Inc.     

ASPECTS OF THE BREEDING BIOLOGY OF THE FIERYNECKED NIGHTJAR

Jackson H. D. 1985. Aspects of the breeding biology of the Fierynecked Nightjar. Ostrich 56: 263–276.

A marked population of nightjars in Zimbabwe was studied intensively for four breeding seasons. This paper covers certain aspects of the breeding biology of the Fierynecked Nightjar Caprimulgus pectoralis. The male shows strong site fidelity during the breeding season (September to December), singing, feeding and breeding within an area of about 5,8 ha. There is some evidence of site defence by the male. The female shows strong mate fidelity, resulting in a pair bond for life. Egg laying starts with full moon in September and is further stimulated by the next two full moon periods. The eggs are laid directly on dense leaf litter at a site overhung by foliage. The normal clutch is two eggs (12S % are one egg) laid on successive days during the afternoon. Incubation starts with the first egg and is by the male at night and the female by day. The incubation period is 18 days. The birds respond to undue disturbance by deserting the eggs and laying a replacement clutch. The chicks usually hatch on successive afternoons; they are mobile on the first day and will move to a parent if called. Both parents feed and brood the young during twilight and moonlight; the male broods them on dark nights and the female does so by day. The species is double-brooded when time permits, the female laying again once the first brood has reached independence; she may even lay a third clutch if the second one comes to grief. There is no evidence of adults transporting eggs or young.     

Proceedings of the New Zealand Society for Parasitology Inc.

Abstract

A study of the nesting habits and breeding biology of blue penguin Eudyptula minor was undertaken over the 1995–96 and 1996–97 breeding seasons on Matiu‐Somes Island, Wellington, New Zealand. Male and female blue penguins tended to be faithful to both mates and nest sites, although there was insufficient evidence to detect any association between a bird's breeding success in 1995 and a subsequent change of mate or nest in 1996. Over the 1995 and 1996 seasons the recorded hatching success (0.51 ±0.11 and 0.63 ± 0.10 respectively), fledging success (0.81 ±0.12 and 0.85 ±0.10 respectively) and reproductive success (0.41 + 0.11 and 0.54 ± 0.11 respectively) were similar each season. There was no significant difference between the proportion of eggs laid, or eggs hatched and chicks fledged, between the two seasons. The mean number of chicks raised over the two seasons was 0.94 ± 0.05 per nest. Replacement clutches were laid by 11 per cent of failed breeders in each season, but only in 1996 were they successful in fledging chicks.

No significant difference was found between the breeding success of the Matiu‐Somes Island blue penguin colony recorded during this study and a previous study undertaken on the island 40 years ago.     

BREEDING BEHAVIOUR OF OSPREYS PANDION HALIAETUS IN SCOTLAND

Ospreys Pandion haliaetus nested at a site near Loch Garten, Inverness-shire continuously from 1959 to 1973. Each year the Royal Society for the Protection of Birds has organized a continuous watch on the eyrie in the breeding season. The detailed records kept of the activities of Ospreys at the nest by those participating in the watch were analysed and the results presented here. Ospreys are migratory and arrived in the breeding area in early April. Nesting material was usually added to an existing eyrie platform. The male collected more material than the female. The female lined the nest cup. The extent of nest building activity and the frequencies of mating and other activities prior to laying varied markedly from year to year. These differences may have been related to changes in the identity of the nesting female, but the birds were not individually marked. Both sexes incubated but the female took the greater share and normally incubated at night. When the young hatched they were brooded by the female. The female stayed in the vicinity of the nest for most of the time until the young fledged at about 53 days old. The male Osprey caught almost all the fish eaten by his mate and young during the breeding season. The number of fish caught per day increased markedly after the young hatched. Pike Esox lucius and Trout Salmo trutta were the main species taken, and some Rainbow Trout Salmo gairdnerii were identified. There were seasonal and diurnal changes in the size and the species composition of the catch. The effects of weather conditions on hunting are examined. The occurrence of Ospreys other than the resident birds at the nest site is described. The behaviour of another pair of Ospreys which repeatedly failed to hatch eggs is described. There was an instance of egg eating in this pair, and some differences in behaviour were found between these birds and those at Loch Garten whose breeding success was good. The breeding biology of Ospreys is compared with that of other British diurnal birds of prey. In other species the female leaves the young unguarded at some stage in the nestling period and hunts food for them, whereas female Ospreys do not usually hunt in the nesting period.     

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.     

6. GENERAL NOTES

Stutterheim, C. J. 1982. Breeding biology of the Redbilled Oxpecker in the Kruger National Park. Ostrich 53:99-90.

The nest of the Redbilled Oxpecker Buphagus erythrorhynchus in the Kruger National Park is a natural hole in a tree where no excavation is required. No evidence of a territorial system WBS observed and only the nesting tree is defended. Mammal hair, dung, grass and rootlets are used for nesting material. The average clutch size was 2.8 eggs with a mean incubation period of 12,6 days. The average nestling period was 30 days. The Redbilled Oxpecker can raise three broods in a season of 176 days such as in the 1973/74 breeding season. The activity area of one breeding group was 7,0 km². The breeding unit consists of two to five birds with helpers of both sexes. All the birds in a group help to select a nest site, build the nest and feed the young. Only one male and one female participate in incubation. Post-hatching development was studied in 13 chicks.     

OBSERVATIONS ON THE BREEDING OF THE PINK-BACKED PELICAN PELECANUS RUFESCENS

This paper summarises observations on the breeding behaviour of the Pink-backed Pelican Pelecanus rufescens at Rakewa, Nyanza Province, Kenya, where the species has bred for at least 200 years. Observations covered most of one breeding season, November 1962 to April 1963. Of at least 250 nests, 35 were closely observed. The community consisted of about 815 pelicans of which about 540 were adults. The death rate is estimated at 13% per annum and the mean life-span at about seven and a half years. The breeding site, in trees above a small swamp, is 15 miles from the favoured feeding ground. The colony is protected by local Luo people. The pelicans feed and roost mainly at the Miriu Delta, 15 miles away, travelling between the two places so high up as to be unseen. They fish in the early morning, visiting the colony to feed young mainly between 09.00 and 13.00 hrs. Once the young are large both parents roost away from the colony at the Delta. The breeding season takes place from August, towards the end of the rains, to March, at the end of the dry season. The birds breed in synchronised groups, the breeding cycle for any group occupying five months. Nuptial display is performed on the nest trees, by single pairs or small groups. Two main displays are described, “pointing” and “bill-clapping”. Mating occurs on the nest, with little preliminary display. Nests are slight stick structures, repaired from year to year, and used by other pelicans if abandoned. The clutch is normally two eggs, occasionally three. Both sexes incubate, with infrequent change-overs, for 33–35 days. The chick is first brick-red, becoming covered with white down. Feathers break through at about 12 days and have covered much of the body by 30 days. At 40 days chicks can recognise their own parent. They fly at 70–75 days. Parents feed chicks by regurgitation, sometimes into the nest. They brood them closely at first, but after 10–12 days leave them much alone. Large chicks thrust the head far into the parental gullet, and injuries result from such feeding struggles. Feeding usually occurs before mid-day, each parent normally delivering two feeds with a rest between. Curious convulsive movements of the young are probably begging displays. Forty-two young hatched in 35 nests, an average of 0.6 chicks/egg laid. The heaviest mortality among young occurred between 10–30 days when 31% of all chicks died. Young which flew were produced at the rate of 0.47/egg hatched, 0.28/egg laid, and 0.57/pair.     

The breeding biology of the Namaqua Sandgrouse,Pterocles namaqua

The breeding biology of the Namaqua Sandgrouse, Pterocles namaqua, was studied and its nesting success determined through the observation of 278 nests over four consecutive breeding seasons at Droëgrond, Northern Cape Province, South Africa. The normal clutch of three eggs is laid over five days (±48-hour laying interval). The incomplete clutch is left unattended overnight, but is attended during the heat of the day by the female on days when an egg is laid and by the male on alternate days. After clutch completion, the pair share incubation, with the female relieving the male 151 min (±21 S.D.) after sunrise and the male relieving the female 105 min (±21 S.D.) before sunset. The incubation period is 21 days from clutch completion, and the three chicks normally hatch within 18 hours of one another. Nesting success ranged from 5.7% to 13.5% between seasons and averaged 8.2%. Predation, primarily by small mammals, was responsible for 96% of nest losses. Estimates of annual recruitment at Droëgrond ranged from minima of 3–10% to maxima of 6–20%, and are believed to be representative of a core area of the distribution of the Namaqua Sandgrouse in South Africa. These low estimates suggest that annual juvenile recruitment may be too low to maintain Namaqua Sandgrouse populations locally. Possible reasons for the sustained low level of breeding success are discussed.     

Breeding behaviour and performance of the Knysna Warbler Bradypterus sylvaticus on the Cape Peninsula,South Africa

Three pairs of Knysna Warblers were monitored on the south-eastern slopes of Table Mountain during the 2000 breeding season. Males displayed alone on territories until the second half of August, when females arrived. Nest-building (8 days) and incubation (16 days) were undertaken entirely by the female, who was not fed on the nest by the male. Chick provisioning was done mainly by the male. Arachnids and terrestrial amphipods were the most common prey brought to chicks. The fledging period was 12 days. Modal clutch size was three eggs, and depredation rates of eggs and chicks were high. After losses, replacement clutches were laid on average 19 days later, after a new nest was built. A maximum of three clutches per pair was recorded. Of 18 eggs monitored, 28% hatched and 17% fledged, equating to a production of one fledgling per pair per year. Ten days after fledging, the entire family leaves the territory, males probably returning once young are independent. The main threats to the local populations are clearing of riparian undergrowth and management practices that impact the predators of rodents.     

THE BIRDS OF BLYTHSWOOD AND SOME NOTES ON BIRDS OF THE DISTRICT

Wintle, C. C. &; Taylor, P. B. 1993. Sequential polyandry, behaviour and moult in captive Striped Crakes Aenigmatolimnas marginalis. Ostrich 64:115-122.

Captive Striped Crakes showed sequential polyandry, the female laying for a second male when the clutch of her first mate was about to hatch. Where aviary space permitted each male set up a breeding territory and each female defended a larger area encompassing the territories of one or two males. Non-territorial subordinate males and females did not breed. The female initiated breeding by attracting the male and soliciting copulation, and the male incubated the eggs and cared for the young. Incubation took 17–18 days, the chicks left the nest at 4–5 days of age and were fully grown and capable of flight at 46–53 days. Breeding occurred from September to March and males normally reared two broods per season. Territoriality was evident only during the breeding season. Juvenile plumage was a duller version of the sexually dimorphic adult plumage; post-juvenile moult bean at 13–15 weeks and was complete at 21 weeks. Remex moult was simultaneous and a complete moult regular1 occurred twice a year in adults, in December and April (males) and September and March/April (females).     

4. UNUSUAL EXPERIENCES WITH BIRDS

Earlé, R. A. 1989. Breeding biology of the Redbreasted Swallow Hirundo semirufa. Ostrich 60: 13–21.

The two races of the Redbreasted Swallow Hirundo semirufa seem to have separate breeding seasons with the northern race H. s. gordoni breeding April-July, while most records for the nominate race fall in October-February. All nests studied were in concrete culverts less than 1 m high. Eggs laid in second clutches by individual females weighed significantly less than eggs laid in first clutches. Eggs hatched on average 16,2 days after incubation started or 18–21 days after the eggs were laid. Only females incubated. Chicks fledged 23–25 days after hatching and reached a maximum body mass of about 31,5 g on day 18 before a steady decline in mass until fledging. Most nesting failures resulted from infertile eggs or starvation of young in the nest (16,2% of all young starved). Overall breeding success was 60,6%. In all, 81,8% of first clutches produced fledglings but only 44,4% of second clutches. Over a three year period 4,9 young were produced per pair breeding in the area (1,6 young/pair/breeding season).     

Incubation prior to clutch completion accelerates embryonic development and so hatch date for eggs laid earlier in a clutch in the great tit Parus major

The ability of parents to respond to changes in food supply within a season will have a large effect on fitness through the number and quality of chicks fledged. Great tits, Parus major, attempt to synchronise their production of chicks with a seasonal food peak, but when food supply fails, hatching asynchrony of chicks provides a mechanism by which some young can be fledged because more developed chicks outcompete their less developed siblings for the reduced parental food supply. We tested whether female great tits can potentially control the degree of hatching asynchrony by using incubation before clutch completion, so that early laid eggs develop faster and hatch sooner. The temperature of an artificial egg placed in 29 nests during the laying period was measured with data loggers, and nocturnal incubation of eggs similar to incubation post clutch completion was recorded in all nests. We then demonstrated that eggs removed from the nest for 72 hour periods prior to clutch completion hatched later than eggs remaining in the nest for the entirety of the laying period. Our results show that variable pre clutch completion incubation (which was mostly nocturnal) can lead to faster embryo development and earlier hatching, so potentially providing a mechanism for adaptive female control of degree of hatching asynchrony.     

4. A FIELD NATURALIST'S PEREGRINATIONS. BIRD NOTES FROM A NORTHERN SECTOR OF SOUTHERN RHODESIA—Part III

Boyer, H. J. 1988. Breeding biology of the Dune Lark. Ostrich 59:30-37.

The peak of the breeding season of the Dune Lark Mirafra erythrochlamys occurred in January and February and was not dependent on rainfall. Most nests were domed, although one undomed nest was recorded. Ninety-one percent of clutches were of two eggs (mean = 1,9; range 1–2; n = 11). The eggs are described and measurements given. Incubation, by the female only, began with the laying of the second egg, and hatching occurred after 13–14 days. Growth and development of nestlings are described. The young left the nest after 12–14 days, and post-nestling parental care lasted for approximately one month. Sixty-one percent of eggs hatched. and 28% produced young which successfully left the nest. Most losses of eggs and young were the result of predation.     

THE BREEDING BIOLOGY OF THE BLACK-BELLIED STORM-PETREL FREGETTA TROPICA

The breeding cycie and habits of the Black-bellied Storm-petrel are described from observations made over three seasons at Signy Island, South Orkney Islands. The species is strictly nocturnal on land and nests in stable scree slopes. With an estimated population of 100–200 pairs, Fregetta tropica is the rarest petrel breeding on the island. In general, the breeding cycle of F. tropica resembles that of Oceanites oceanicus. Birds usually arrive from mid-November onward and return to the same nest and mate in successive seasons. The female is absent from the nest for a week or more before the egg is laid, during which time the male continues to visit the site. From ten laying dates, egg laying appears normally to begin during the last week in December, but evidence is given that, in 1966-67, laying was delayed by heavy winter snow build-up coupled with a late melt. The egg comprises 26 % of the body weight of the female. Incubation is by both sexes in alternate spells of three days, the whole period lasting 38–44 days. The chick is left alone in the nest by the parents almost immediately after hatching. Chick growth is described briefly and the effects of drift snow on development are discussed. The fledging periods of two chicks were 65 and 71 days, departure from the nest taking place in mid-April. Measurements of 36 Signy Island birds show considerable variability but are similar to those from other breeding localities.     

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.     

EDITORIAL

Olver, M. D. &; Kuyper, M. A. 1978. Breeding biology of the Whitebreasted Cormorant in Natal. Ostrich 49:25-30.

From 1972–1975 Whitebreasted Cormorants Phalacrocorax carbo bred at the Cedara Dam, Natal, South Africa (29 32S, 30 17E), and from 1973 they fished for food at Midmar Dam, 5 km away and carried the food back to the nestlings. Breeding occurred from April to October and was preceded by a period of courtship. Nesting material was collected by the males and the nests built by the females. The mean clutch size for 1972–1973 was 3,1. Both parents incubated the eggs and guarded the nest and chicks. Growth of the chicks was studied in 1972–1973. The mean number of chicks reared was 1,6 per nest although seven nests contained three nestlings. At 28 days they left the nest when alarmed, but could not fly until 49 days old. The average flying age appeared to be about 53 days. The height of the nests above the ground seemed to determine the nest leaving age. Of the 186 eggs laid in the 60 nests observed over two years, 74% hatched. Fledging success was 52% of eggs laid and 69% of eggs hatched. Chick mortality seemed to be caused mainly by falling from the nests and dying of starvation.     

Cooperation and conflict among dunnocks, Prunella modularis, in a variable mating system

The mating system of a population of 90 breeding dunnocks (or hedge sparrow, Prunella modularis) included monogamy, polygyny, polyandry and polygynandry. Monogamous males guarded females during their fertile period to prevent neighbouring males from copulating. The most intense guarding occurred where two (unrelated) males shared a territory. Here, the alpha male tried to prevent the beta male from copulating with the female. Beta males were seen to copulate in only half the cases. They were more likely to succeed when the alpha male found it difficult to guard the female closely because her range was large, the vegetation was dense or there were other females breeding synchronously on the same territory. Close guarding and chasing by males reduced the female's feeding rate and was correlated with unhatched eggs in the nest. Females attempted to escape the alpha male's attentions and actively encouraged the beta male to mate. Beta males only helped to feed the young if they copulated with the female. Nestlings fed by two males and a female got more food and weighed more than those fed by just one male and a female. Indirect evidence suggested that when beta males failed to copulate, they destroyed eggs or young chicks. Females laid larger clutches when two males mated with them as opposed to one, thus adapting their clutch size to the amount of parental care they expected. The results of natural removal experiments and matched comparisons of the same female in different mating systems support these conclusions. For females, selection favours cooperative polyandry, whereas for males if favours polygyny; the variable mating system may reflect the different outcomes of this sexual conflict.     

OBSERVATIONS ON THE BREEDING OF THE PALILA PSITT1ROSTRA BA1LLEUI OF HAWAII

The behavioural ecology and breeding biology of the endangered Palila Psittirostra bailleui was studied from 1971 to 1975. The most intensive breeding occurred from June to August, and coincided with peak production of mamane Sophora chrysophylla seeds, the bir?s major food source. The Palila was able to make adjustments in its breeding to compensate for yearly differentiation in the timing and abundance of this food supply. Sexual chasing and courtship feeding were the most frequently encountered pre-nesting behaviours. Territory was a mate-defended area, which later in the nesting sequence was confined to the nest site. A total of 26 nests was found; most were placed on larger branches of mamane trees. Nest construction occurred primarily in the morning hours and lasted up to 20 days. Both sexes took part in nest construction, albeit the male role was minimal. Unless the nest was placed in the terminal fork of a tree, it usually contained a large stick base. The modal clutch size was two; eggs were laid early in the morning and in all cases one per day. Incubation sometimes began with the first egg and lasted 15–16 days. Only the female incubated, and she covered the eggs for about 75% of the daylight hours and throughout the night. Egg hatching was asynchronous, with the first young emerging early in the morning and the second not until later that same day. Only the female brooded, and the rate declined until day 15 when essentially it stopped. Both parents fed the young by regurgitation, and the number of feedings per hour decreased slightly over the nestling period. It is thought that insects and finely masticated plant material formed the bulk of the nestling diet until about day 5 when mamane seeds became important. Helpers were found at one nest. Young developed slowly and did not leave the nest until 21–27 days old. It is believed that these prolonged nestling periods were able to evolve because of the (former) absence of ground predators. After fledging, young remained with their parents for at least 30 days. Productivity was regulated by small clutch size, low population numbers and by the length of an individual nesting sequence (in that a pair could potentially raise only one brood each year). The primary reason for the endangered status of this bird appears to be the effect of habitat alteration upon a specialist, coupled with the fact that the small effective breeding population and low dis-persability of the species may have resulted in decreased genetic fitness.     

BREEDING ECOLOGY OF TURNSTONES ARENARIA INTERPRES AT HAZEN CAMP, ELLESMERE ISLAND, N.W.T.

Turnstones arrive on Ellesmere Island in late May or early June. Pair-formation takes place during migration, or after arrival in courtship groups along the beaches, or on the nest territory, depending on weather conditions. Pair-formation was not observed at Hazen Camp in 1966 as most birds were already paired when observations began on 3 June. The preferred nesting habitat was Dryas-hummocked tundra closely associated with a marsh, stream, or pond. A census area of 240 ha supported 13 breeding pairs, possibly 14; the total number of pairs breeding in the Hazen Camp study area was estimated to be 70 (3.04 pairs/km²). Egg-laying began on 10 June, with 46% of the first eggs laid 13–15 June. 62% of the sets were completed between 19 and 21 June. Both sexes incubated, the female regularly and the male sporadically. Hatching was also well synchronised; most clutches hatched between 7 and 14 July. Nest success was high (79%). After hatching territories dissolved and family groups moved freely over the tundra, concentrating at ponds where food was readily available for the young. Both adults attended the young during the pre-fledging period, but the females apparently departed long before the young fledged. Males left once the young could fly and the adult fall migration was complete by early August with the exception of late breeders. Most of the young departed in the second half of August. Fall migration is complete by late August or early September. The breeding season appears to be timed so that the young are raised when food is most abundant. Food supply (dipterous insects, especially adult chironomids) was highest from 8 to 12 July at the peak of the hatching period. While food supply for the young declined during the growing period, the early departure of half the adult population, and family movements over the tundra, appeared to reduce the food demand correspondingly. Food appeared to influence the distribution of breeding pairs markedly, restricting them to the vicinity of marshes, streams, and ponds. Territoriality displayed by Turnstones is believed to be closely associated with the protection of the nest against predators and adult food requirements during incubation; it also seems likely that territory has at least a local effect in regulating the number of breeding pairs.