Behaviour of young Ringed Plovers Charadrius hiaticula and its relationship to growth and survival to reproductive age

The behaviour, growth and survival of Ringed Plover chicks were studied in 1974 at Mestersvig, northeast Greenland, and from 1973 to 1976 at Lindisfarne, northeast England. Post-fledging survival, dispersal and recruitment were also investigated for the Lindisfarne population. Growth rates were similar in both study areas, and to those reported from other sites. A diurnal rhythm in feeding activity was more evident in the Arctic than at temperate latitudes, despite continuous daylight in the former area. This was correlated with low nocturnal temperatures and prey availability. It is concluded that food supply did not limit growth or chick survival in either study area. Brooding decreased progressively during the pre-fledging period; the relationship of brooding period to age, environmental conditions and area was investigated. The adaptations of feeding and brooding behaviour for arctic and temperate breeding Ringed Plovers are discussed. Survival from hatching to fledging varied between 40% and 60% in different areas and years, and at least 59% of birds survived from fledging to about one year old. Minimum survival from one to two years old was only 57%. but year-to-year survival of older birds was higher, that of breeding adults being at least so?;. Most birds returned to the breeding area and attempted to breed when one year old. The extent of dispersal from Lindisfarne and the wintering areas of the Lindisfarne-breeding birds are investigated.     

Annual variation in breeding biology of gentoo penguins, Pygoscelis papua, at Bird Island, South Georgia

The breeding biology of the gentoo penguin, Pygoscelis papua , was studied over a three-year period (1986–1988) at Bird Island, South Georgia, with particular reference to birds of known age or breeding experience. Laying date varied significantly between all three years, being three weeks later in 1987, when the breeding population decreased markedly. Factors involved in the timing of breeding are discussed. Within years egg-laying was highly synchronous: 95% of clutches were initiated in 14·5 days or less. The incubation period was 35 days and the laying interval, between the two eggs, 3·3–3·4 days. Chicks creched when 25–30 days old, and this varied between years, possibly related to food supply and chick growth. Chicks left the colony for the first time between 75 and 85 days of age. The breeding population at Bird Island decreased by 20% and increased by 84% in successive years during the study period. Breeding success (chicks fledged per egg laid) varied between 0·33 and 0·65 within colonies, but for the whole island was very consistent over the three years: 0·45, 0·51 and 0·47. Overall, colony differences were not correlated between years. Disturbance from Antarctic fur seals, Arctocephalus gazella , is suggested as the cause of consistently lower breeding success at one colony. Mean egg weight varied annually, and with age of the breeding bird, nest location and, in one year, with laying date. Young, first-time breeders laid smaller eggs and had lower breeding success compared to older, experienced birds, similar to other seabirds. However, they differed from other species in laying on average earlier than older birds. The relationship between age, egg weight, laying date and breeding success is discussed in relation to predation and seasonal food supply.     

Senescence effects in an extremely long-lived bird: the grey-headed albatross Thalassarche chrysostoma

Studies attempting to document reproductive or other pre-lethal senescence effects in wild birds typically face an array of problems, including flaws in statistical analyses, non-adaptive philopatry to deteriorating environments, confounding effects arising from cohort heterogeneity and differential death rates of phenotypes and the frequent pairing of old birds to younger mates. Furthermore, recent studies suggest that birds could maintain a high level of physical fitness until old age, before being struck by a catastrophic illness leading quickly to their demise. The presence of terminally ill individuals in most datasets (and their greater incidence in older age categories) may therefore provide a false impression of progressive senescence in cross-sectional analyses. This study was designed explicitly to avoid all the known pitfalls linked to the demonstration of progressive senescence in wild populations, and involved one of the very longest-lived bird species. We show that, during incubation, old (aged 35 years and over) male grey-headed albatrosses Thalassarche chrysostoma make longer foraging trips, and have lower daily mass gains, than experienced mid-aged individuals (aged up to 28 years). This is, to our knowledge, the first report documenting reduced foraging performance with old age. Hatching and breeding success of pairs composed of two old individuals were reduced in comparison to mid-aged pairs. Overall results were very similar when analyses were repeated using only individuals known to have survived 1 or 2 years beyond field measurements (hence probably not suffering from the effects of an advanced terminal illness). We conclude that extremely long-lived individuals usually experience some degree of general physical deterioration, leading to reduced foraging and breeding performance, long before their final demise.     

Influence of different photoperiods on development of gonad, cloacal gland and circulating thyroid hormones in male Japanese quail Coturnix coturnix japonica.

One day old chicks of Japanese quail were exposed to different photoperiods (LD, 8:16, 13.5:10.5, 16:8 and LL) and observations (testes weight, cloacal gland size, body weight and circulating thyroxine and triiodothyronine) were taken at the age of 3, 5, 7, 9 and 16 weeks. Results indicate that immediate reproductive development occurred in birds exposed to long photoperiods (greater than 12 hr). Growth under LD 8:16, was not apparent till 7th week and by 16 weeks, degree of gonadal development was similar in all the birds, irrespective of photoperiodic treatment. Whereas body weight of the intermediate and long day (LD 13.5:10.5, 16:8 and LL) treated birds increased upto 5th week and remained constant thereafter. But the chicks maintained under short day length (LD 8:16), showed spontaneous increase till the end of the study and birds were much heavier compared to all other groups. Plasma T4 concentration increased with increasing age till 9th week and remained unaltered thereafter. On the other hand T3 level did not change till 7th week followed by a decline. It is suggested that the initiation and degree of gonadal growth in quail depends on the availability of daily photoperiod, until the achievement of full breeding condition. Peak level of T4 observed in 9 week old birds may be involved in the development of photorefractoriness at that age.     

ASPECTS OF THE POPULATION DYNAMICS OF CAPE VULTURES IN THE CAPE PROVINCE

Robertson, A. S. 1984. Aspects of the population dynamics of Cape Vultures in the Cape Province. Ostrich 55: 196–206.

Information gathered in 1981 and 1982 and collated from previous records on the numbers, spatial distribution, proportion of age classes, age and frequency of breeding, breeding success and causes of breeding failure, and the survival of immature and adult Cape Vultures Gyps coprotheres in the southern and southwestern areas of the Cape Province, South Africa, is presented. This sub-population of about 75 birds is apparently isolated from conspecifics in the rest of southern Africa; the implications of this are discussed. At the Potberg colony in both years an average of 85% of birds 5 years and older were involved in breeding attempts. The age of first breeding was 4–6 years. Nest sites were active for about two in every three years. Between 1975 and 1982, 0,51-0,67 nestlings were reared per active nest site (n=165). Four (possible maximum six) of 21 immatures were resighted one year after they had flown. Of 123 birds that had been ringed at Potberg to 1980, 14 (11%) were sighted in 1981; only four of 48(8%) colour-ringed birds 5 years old and older were breeding in 1981.     

Body mass changes in Brünnich's guillemots Uria lomvia with age and breeding stage

Body mass of Brünnich's guillemots Uria lomvia breeding at Coats Island, Canada, was measured during incubation and chick-rearing in 1988–2001. In most years, mass increased during incubation and fell after hatching, leveling off by the time chicks were 18 d old, close to the age at which chicks departed. Mass during incubation increased with age up to about 12 yr, but the mass of birds brooding chicks was not related to age. The trend towards increasing mass during incubation was mainly a consequence of mass increases of young breeders as older birds maintained a constant mass. The variation in adult mass with age during incubation seems likely to reflect age-related variation in foraging ability, but the loss of mass after hatching, being greater for older birds, appears best explained as a response to the demands of provisioning chicks, with older birds transferring their accumulated reserves to their chicks via higher provisioning rates.     

Acquisition of foraging skills by Heron Island Silvereyes Zosterops later alis chlorocephala

Age-related differences in foraging efficiency and behaviour were investigated in a population of colour-ringed Silvereyes Zosterops lateralis . First-year birds were less efficient foragers than older birds with second-year birds being intermediate. First- and second-year birds had lower success rates than older birds overall, and for most capture techniques and substrates. First-year birds never attempted breeding while about half of the second-year birds and all of the older birds did. Both learning and selection effects may have been involved in causing the age-related differences in foraging behaviour. Foraging skills appear to improve during the first 2 years of life and selection could remove the least efficient foragers from the population in winter when food is short. That second-year birds, some of which had already attempted breeding, were not significantly more efficient foragers than first-year birds suggests that reproduction is not delayed until adult levels of foraging efficiency have been attained.     

BIRDS OF THE UNDERSTORY OF LAKE-SHORE FORESTS ON THE ENTEBBE PENINSULA, UGANDA

Monthly mist-netting of low-flying forest birds was conducted in three lake-shore forests in southern Uganda from September 1970 to July 1972. It was found that the edge of the forest formed a distinct ecological barrier that was rarely crossed by species on either side of it. Catches were concentrated in the hours of daylight, in two peaks, a morning peak at 09.00 hrs and an afternoon peak at 16.00 hrs (sun-time) for all birds combined, although there were slight species differences. Most bird species were found to be sedentary in a fixed home-range, and covered lateral distances of up to c. 300 m. No movement between the different forests was recorded.
The most common birds such as bulbuls were often caught at heights between 0·6 and 1·8 m above the ground, but the upper limit of their vertical range was not determined. It is suggested that the bi-modality of flight activity may be intrinsic, even though it may be correlated with such extrinsic factors as light and radiation. On a month to month basis, the numbers of birds caught reflected breeding and moult activity. This was best shown by the Pygmy Kingfisher, taken in higher numbers towards the end of the first rainy season when a majority of the birds caught were young and many birds were moulting the remiges. On the other hand higher numbers of Olive Sunbird were found during the second rainy season, again at a time corresponding to a general moulting of remiges. The bulbuls showed fluctuations in numbers caught that were matched by an equally fluctuating pattern of moult. The role of food as a possible limiting factor in bird breeding is discussed.     

Body mass changes in Brünnich's guillemots Uria lomvia with age and breeding stage

Body mass of Brünnich's guillemots Uria lomvia breeding at Coats Island, Canada, was measured during incubation and chick‐rearing in 1988–2001. In most years, mass increased during incubation and fell after hatching, leveling off by the time chicks were 18 d old, close to the age at which chicks departed. Mass during incubation increased with age up to about 12 yr, but the mass of birds brooding chicks was not related to age. The trend towards increasing mass during incubation was mainly a consequence of mass increases of young breeders as older birds maintained a constant mass. The variation in adult mass with age during incubation seems likely to reflect age‐related variation in foraging ability, but the loss of mass after hatching, being greater for older birds, appears best explained as a response to the demands of provisioning chicks, with older birds transferring their accumulated reserves to their chicks via higher provisioning rates.     

Reproductive endocrinology of the Wandering Albatross Diomedea exulans in relation to biennial breeding and deferred sexual maturity

The reproductive endocrinology of the Wandering Albatross Diomedea exulans was studied at South Georgia to investigate the potential endocrine correlates of biennial breeding and of the acquisition of sexual maturity. Gonads of breeding birds and of known-age immature birds of both sexes were examined by laparoscopy throughout the period that they were at the nest site. Blood samples, subsequently analysed to determine concentrations of luteinizing hormone (LH), prolactin, progesterone, testosterone and oestradiol-17/i, were obtained from samples of breeding birds of both sexes at regular intervals from first arrival until the chicks fledged nearly a year later. Before laying in December, breeding birds had mature testes and ovarian follicles and high concentrations of LH, prolactin and sex steroids. Gonadal regression and a rapid drop in hormone levels (except for LH in females) occurred in early incubation (January). Testes (and follicles to a lesser extent) enlarged in mid-incubation, coinciding with high levels of LH and increases in prolactin and testosterone. Gonads finally regressed completely near hatching time. LH, prolactin and testosterone remained at low levels throughout chick rearing (April to November), but females had several periods of active progesterone and oestradiol secretion, and progesterone was detectable in males only late in the chick-rearing period. Although some changes in hormone levels are difficult to explain, the patterns are fairly typical of temperate birds. The persistence of progesterone secretion in both female breeders and non-breeding ‘immature’ birds is viewed as part of a mechanism inhibiting an ovary from becoming vitellogenic. Although testis size and testosterone concentrations increased with age in immature males (of ages 4–10 years), birds of 5 years and older are probably physiologically mature, even though breeding does not start until they are 7 years of age and only half an age group has bred by an age of 11 years. Immature females (of age 4–7 years) had undeveloped follicles, very low oestradiol concentrations but high progesterone levels, providing further support for the role of this hormone in inhibiting gonadotropin secretion. The condition of the female is therefore probably decisive in determining when a pair first attempts to breed but it is unknown what factors initiate normal ovarian development.     

Feeding ecology of the tufted puffin (<Emphasis Type="Italic">Lunda cirrhata</Emphasis>) and the horned puffin (<Emphasis Type="Italic">Fratercula corniculata</Emphasis>) in the northern Sea of Okhotsk

The feeding of tufted puffin and horned puffin nestlings was studied on two islands of Tauyskaya Bay of the Sea of Okhotsk: at Talan Island in 1999–2004 and 2006 and Umara Island in 1994 and 1996–1997. The composition of the puffin diet during the breeding season was determined. The main fish species in the diet were determined and the seasonal and annual dynamics of the occurrence of the items was analyzed. Comparative data on biological characteristics, such as the size, weight and age of the fish caught by these birds, as well as the size and composition of food samples, were collected. Differences between the age groups of fish in the diets of the two puffin species were revealed. Conditions for forming of seabird feeding base in Tauyskaya Bay were described.     

Non-breeding behaviour of Magenta Petrels Pterodroma magentae at Chatham Island, New Zealand

Magenta Petrels Pterodroma magentae were caught at light-attraction stations on southwest Chatham Island, New Zealand, and most were fitted with transmitters. Of 52 captured since 1993, 71% were males, and all 36 tracked adequately proved to be non-breeders in the breeding season of capture. Our data indicated no sex bias in their probability of being captured at lights. Males provided 86% of trackings, and 87% of trackings of birds flying over the breeding area were males. Males landed 118 times; females 13 times. Only males were found on the ground, by night and day, apparently unassociated with burrows (three with and ten without transmitters), but subsequently digging burrows ( n  = 8). Of 19 birds banded as fledglings up to 2000, males were first recaptured nearing 4 years old (at lights and on the ground) and a female nearing 6 years old (in burrow). Among 37 fledglings, the sex ratio was even. Nine tracked males occupied burrows, as did two females, but the latter were older recaptures (10+ and 25+ years old). It appears that only males claimed existing, or dug new, burrows. They then attracted a mate to the burrow by means unknown, but from among females frequenting an inshore courtship area near the colony, or occasionally flying over the colony, at night. Females established in burrows, but then losing their mate, were able to re-mate there, by calling from near the burrow or by attracting a mate in flight or from the postulated inshore courtship area. Both sexes sometimes took years to pair or re-mate, possibly reflecting the dearth of available mates.     

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.     

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.     

Spatial distribution and behaviour of laying hens housed in an alternative system

The spatial distribution and behaviour of perchery housed laying hens were compared at a constant stocking density (18.5 birds/m(2)) in eight pens with colonies of five different sizes (323 birds (N=1), 374 birds (N=2), 431 birds (N=2), 572 birds (N=1) and 912 birds (N=2)). The birds were placed in the perchery when they were 12 weeks old. Observations began when they were 26 weeks old and continued at 8 week intervals until 61 weeks of age. Colony size did not appear to affect the spatial distribution of birds, but more standing behaviour and less feeding behaviour were observed in the smallest and largest colony sizes. Older birds spent more time on the floor areas and less time on perches. Young birds (26-28 weeks) spent more time feeding, foraging, drinking and preening, and less time standing idle than older birds. In the afternoons, there were fewer birds on the perches and more on the floor levels, corresponding with less time spent resting and more time spent performing active behaviours. Birds did not distribute themselves evenly throughout their pens: within specific areas of pens densities varied between 9 and 41 birds/m(2). This variation, which reflects the flux of birds from one part of the pen to another, was greatest for the larger colony sizes, and may have adverse implications for welfare in terms of crowding and hysteria.     

Annual cycles in Jamaican forest birds

The annual cycles of forest birds in Jamaica were found to be very similar to those at higher latitudes. Most species bred between March and September, though a few possibly breed throughout the year, especially in cultivated areas. Primary moult followed immediately after breeding, and in some species was apparently arrested to allow a further breeding attempt. Several species were fatter outside the breeding season than during it, and this is interpreted as "winter fattening" comparable to that found in many birds at higher latitudes. Weights varied little but individuals retrapped were usually heavier outside the breeding season. In some species the first complete moult took place at the end of the first year, implying that the birds do not breed until at least two years old.     

Patterns of Haemoproteus beckeri parasitism in the gray catbird (Dumatella carolinensis) during the breeding season

We determined the prevalence and intensity of blood parasites in breeding gray catbirds (Dumatella carolinensis) at Killbuck Wildlife Area in Wayne and Holmes Counties, Ohio (USA) from June through August 2000. Of 98 catbirds sampled, 40 (40.8%) had detectable infections of Haemoproteus beckeri. Overall prevalence of H. beckeri in this population is high relative to that reported in earlier blood parasite surveys of both breeding and migrant catbirds. Mean intensity of H. beckeri infection did not vary significantly between young and old birds or among sampling periods. We found no effect of age on prevalence or intensity of H. beckeri infection. Older birds were not more likely to be infected than younger birds, despite longer exposure to arthropod vectors. Prevalence varied significantly with season and was highest in June and lowest in August. This pattern also was observed in older birds sampled repeatedly. This seasonal variation may reflect both newly acquired infections and chronic infections relapsing in response to hormonal changes associated with breeding. Evidence of transmission was observed in the single hatching year bird that lacked detectable infection in early summer, but demonstrated a very high intensity infection in late summer. These observations provide supportive evidence that hematozoa infections are acquired on the breeding grounds during the first year of life and relapse during the breeding season in subsequent years.     

Identification and breeding of yearling Piping Plovers

ABSTRACT Age of first breeding is an important life history trait. Many Piping Plovers (Charadrius melodus) do not breed as yearlings, but little information is available concerning the age of first breeding. From 2000 to 2006, we marked 991 chicks in three areas in Saskatchewan, Canada, and subsequently determined when 102 (49 females and 53 males) first bred. Females bred significantly earlier, on average, than males. More females (68%) bred as yearlings than did males (41%; P= 0.04), with most others first nesting when 2‐yr old (29% of females and 50% of males). As expected from differences between the sexes in age of first breeding, younger females were more likely to pair with older males than were younger males with older females. Chicks that hatched early in the breeding season did not breed at an earlier age than those that hatched late in the year. Unlike older birds, juvenile Piping Plovers do not replace flight feathers during their first winter. As a result, 18 of 27 yearlings (67%) had worn outer primaries, whereas only one of 123 (1%) older birds had worn primaries. In addition, whereas 20 of 24 yearlings (83%) retained a few buff‐tipped median coverts, none of 119 known older birds had such coverts. As a result, we were able to identify all yearlings by their worn primaries, buff‐tipped median wing coverts, or both. Wing lengths of yearling Piping Plovers were 3% shorter than those of older birds, presumably due to wear. Because there is no evidence of differences in adult survival rates between the sexes and breeding habitat is available, we speculate that fewer yearling males than females breed because primary wear may reduce the ability of yearling males to perform aerial breeding displays.     

OBSERVATIONS ON NOCTULE BATS (NYCTALUS NOCTULA) CAPTURED WHILE FEEDING

In the summer and early autumn of 1960, 1961 and 1962, noctule bats flying low, taking house crickets as these flew from a municipal rubbish tip, were captured in mist nets, ringed, released and in many cases recaptured a number of times. The flying bats showed no fear of human beings or predatory birds and did not learn to avoid the net. In June and July of each year the majority of bats caught were adult females, the flying young of the year first appearing in August though some did not fly until September and October. Young males did not reach sexual maturity in the year of their birth, though five out of fourteen females recaptured at a year old did. There was a considerable movement of adult males in the late summer, adult bats being captured in approximately equal numbers of both sexes during August and September. In October the females seemed to disappear, the majority of the bats caught during that month being males: by November the crickets had ceased to fly and no more bats could be captured though a few wero still flying on warm nights. There was a marked difference in feeding behaviour over these three years, the bats concentrating more on crickets in 1960 than in 1961 and 1962. Though the differences are not statistically significant there were indications of an increase in body weight between July and October in the years when less cricket feeding was occurring. About 50 per cent of the females captured in each of the years 1960 and 1961 were recaptured feeding on the same site in the following year: the recovery rate of males was about 30 per cent in 1961, 60 per cent in 1962.     

Survival in a long-lived territorial migrant: effects of life-history traits and ecological conditions in wintering and breeding areas

Despite its key role in population dynamics and evolutionary ecology, little is known about factors shaping survival in long‐lived territorial species. Here, we assessed several hypotheses that might explain variability in survival in a migratory Spanish population of a long‐lived territorial species, the Egyptian vulture Neophron percnopterus, using a 16‐year monitoring period and live‐encounter histories of 835 individually marked birds. Cormack‐Jolly‐Seber capture–recapture models showed no evidence for effects of sex or nestling body condition on survival. However, the normalized difference vegetation index (NDVI; an indicator of primary productivity) of natal territories had positive effects on juvenile survival, indicating that environmental conditions experienced early in life can determine survival prospects. Survival increased with age (0.73±0.02 in the first 2 years to 0.78±0.03 in years 3 and 4) to later decrease when birds were five years old (0.60±0.05), the age at which they acquire the adult plumage, abandon the communal lifestyle of juveniles, and may look for a breeding territory. At older ages, survival was higher for non‐breeding (0.75±0.02) and breeding adults (0.83±0.02). Among the latter, birds that recruited into better territories had higher survival prospects. Age‐specific variation in survival in this species may be related to behavioural changes linked to dispersal and recruitment into the breeding population, while survival prospects of adult birds strongly depend on breeding territory selection. These results suggest a tradeoff between recruiting soon, and thus reducing mortality costs of a long and extensive dispersal period, and trying to recruit into a good quality territory. Finally, annual survival rates for birds of all age classes were positively related with the NDVI in their African wintering grounds. Although this relationship was probably mediated by food availability, further research is needed to properly identify the limiting factors that are affecting trans‐Saharan migrants, especially in light of global climate change.