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 success and chronology of Wood Storks Mycteria americana in northern and central Florida, U.S.A.

The breeding success and chronology of Wood Storks Mycteria americana were studied at eight colonies in northern and central Florida during 1981–1985. Mean ± s.d. clutch size for all colony-years was 3.07 ± 0.56 (n = 2694 nests), with three-egg clutches (72%) most frequent. Mean clutch size among all colonies and years ranged from 2.73 ± 0.55 to 3.41 ± 0.61. Many colonies exhibited significant negative trends in clutch size with, hatching date because of a proportional decrease in four-egg clutches later in the season. Mean colony clutch size was not correlated with nest numbers, nesting density or mean hatching date within most years. Mean ± s.d. number of fledglings for all colonies and years was 1.29 ± 1.16 fledglings per nest (n = 2812 nests). Mean annual fledging rates in colonies ranged from 0 (colony failed) to 2.66 fledglings per nest. Most breeding failure occurred prior to egg hatching, and the second highest mortality occurred between hatching and 2 weeks of age. Four-egg clutches fledged more storks than three-egg clutches, which in turn were more successful than two-egg clutches. However, all clutch sizes showed similar fledgling per egg rates. The seasonal decline in productivity was associated proportionally with smaller clutch sizes later in the breeding season. An increase in mean hatching date was correlated with an increase in latitude. There was greater within-year breeding synchrony among colonies than interyear breeding synchrony within each colony. Breeding synchrony was not correlated with mean hatching date, latitude, longitude, nest numbers or nesting density.     

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.     

Reproductive performance of the Bald Ibis Geronticus calvus in relation to rainfall and grass-burning

Reproductive performance of the Bald Ibis Geronticus calvus was studied during three consecutive nesting seasons (1978–80) at a single large breeding colony in Natal, South Africa. A focal sample of 22 nest sites (out of a total of 60 in the colony) was inspected at frequent intervals to determine clutch sizes, egg survival, hatching success, nestling mortality and fledgling production in each year. Mean clutch size was slightly lower in 1980, compared to the two previous years, although survival and hatching success of eggs were similar in all three years. However, nestling mortality was significantly higher in 1980, largely as a result of parental nest desertion and nestling starvation, so that the proportion of breeding pairs producing at least one fledged offspring, and the average brood size at fledging, were lower compared to 1978 and 1979.
The 1978 and 1979breeding seasons were both preceded by 12 months of above average rainfall, and Bald Ibises fed extensively on small invertebrates obtained in burnt grassland in the study area during the nesting period. By contrast, the onset of nesting in 1980 followed 11 consecutive months of sub-normal rainfall. As a result of the drought, and the consequent lack of controlled burning by landowners in the study area in 1980, very little burnt grassland was available during the Bald Ibis' breeding season, and the birds fed primarily in cultivated pastures during the customary burning period. The lower reproductive performance of Bald Ibises at the study site in 1980 is discussed in relation to the possible effects of drought on the abundance of invertebrate prey and the extent of controlled burning in the species' grassland biotope.     

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.     

Age-specific variation in reproduction is largely explained by the timing of territory establishment in the New Zealand stitchbird Notiomystis cincta

1. Using data from 327 nests over a consecutive 8-year period we examined age-specific variation in reproduction in a population of stitchbirds (or hihi) Notiomystis cincta and related how differences in reproductive performance were linked to the timing of territory establishment and breeding. 2. Across the population all reproductive parameters showed a quadratic relationship with an increase mainly between the first and second breeding season and a decline after the fourth year. A longitudinal analysis showed evidence of senescence by the sixth year in the numbers of chicks fledged and recruited. 3. Reproductive increases between years 1 and 2 were the result of poor-quality females dying after their first breeding season (differential selection hypothesis) in combination with surviving females showing improvements in reproduction in their second year (individual improvement/constraint hypothesis). 4. There was no effect of mate experience or territory quality on improvements in breeding between years. 5. The key variable influencing reproductive output was the timing of breeding. Birds that started laying earlier were more likely to lay multiple clutches in any given season. This was the main difference between first-year and older birds; generally first-year birds initiated egg laying later and consequently laid fewer clutches. 6. Approximately half of all first-year birds did not establish their territory until after the breeding season had begun. This delay in territory establishment resulted in these birds delaying breeding, which resulted in them having a lower reproductive output relative to all other birds. First-year birds that managed to establish their territory before breeding commenced, had similar rates of reproduction as older birds. 7. There was a positive relationship between the timing of territory establishment during a female's first year and her hatching date in the previous breeding season. We hypothesize that this was because late-hatched females were less able to effectively compete for territories against earlier-hatched members of their cohort, and this delayed their establishment and breeding in their first year. Thus, this social constraint is likely to be a major factor driving age-specific reproductive variation in this population.     

Prospectors' colony attendance is sex-specific and increases future recruitment chances in a seabird

In most long-lived vertebrates, including seabirds, young non-breeders often attend potential breeding sites. In seabird colonies, this prospecting behaviour has nearly never been studied with respect to potential sex-specific benefits, and its fitness consequences are largely unknown. We compared attendance patterns of sexed common tern prospectors at six subcolonies with future breeding status and nesting site choice. We also tested for potential effects of population density. Birds that arrived earlier at the colony were recorded more often along the season. This arrival effect was stronger in males, which generally spent more time at the colony. Birds prospecting for two consecutive years attended the colony more intensively in the second year. A high colony attendance enhanced recruitment probability in both sexes, but only in females, it was linked with a higher probability to return. Attendance at a preferred subcolony increased during the season. For first breeding, individuals favoured the subcolony where they had prospected most intensively in the previous season. In males, this subcolony fidelity was stronger and increased simultaneously to breeding pair density. We conclude that prospecting is a process of integration into the community of breeders, and that benefits are higher for males, the more territorial sex in this species.     

Hatching date influences age at first reproduction in the black-headed gull

In long-lived colonial birds, age at recruitment is an important life-history character. Variation in this parameter may reflect differences in several factors, including competitive ability and breeding strategies. Further, these differences may be due to timing of hatching (for instance through differences in competitive ability). We investigated the age of first-time breeders in relation to hatching date in a black-headed gull Larus ridibundus colony situated in central France, from 1979 to 1993. Age at first breeding was estimated for four groups of individuals (total n=550) according to their hatching date, using a recent capture-recapture methodology which allowed us to estimate recruitment rate without the limiting assumptions of methods relying on simple return rates. The age at first breeding was negatively correlated with the hatching date of individuals: individuals hatched earlier in the season started breeding at a younger age than individuals born later. Proportionally more 2-year-old late-hatched individuals were seen breeding on small peripheral colonies than young early-hatched individuals. This difference disappeared after age 3 years. These results strongly suggest that individuals hatched late in the season start to breed on peripheral colonies before recruiting to their natal colony. A difference of few weeks in hatching date has consequences which can last for several years.     

Parental care in Semipalmated Sandpipers Calidris pusilla: brood desertion by females

Although Semipalmated Sandpipers Calidris pusilla are monogamous, with biparental incubation, most females (86–97%) deserted their broods to the care of their mates, 0–11 days (average 6) after their eggs hatched. Males left the brood an average of 8 days later, shortly before or after the chicks fledged. In several instances, females that deserted in one year remained with the chicks the next year, and vice versa. Females deserted chicks at nests that hatched later in the season at an earlier age than those had hatched earlier ( r ²> o.6). Since females appeared to have an energy deficit in at least some years, and suffered higher mortality rates than males during breeding, it is possible that females deserted broods in order to take advantage of better feeding conditions at migratory stopovers in northeastern North America early in the season. There was little evidence of higher nesting success or earlier hatching date in reuniting pairs, although if both members of a pair returned to the breeding area, 80% reunited. Increased survival of their mate may be most advantageous to males in ensuring that they obtain a female the following year.     

Seasonal changes in the distribution and composition of common seal (Phoca vitulina) haul-out groups

Seasonal changes in the distribution and composition of common seal haul–out groups were followed in a study area in Orkney, Scotland. A marking programme was also undertaken, using both conventional and radio–tags, to study individual movements between sites and seasonal changes in site–use. Certain haul–out sites were used only in the breeding season, while others were used during the winter. Seals were seen at one site all year round and at another during only the pre–pupping and moult period. On one island where two sites were used during the summer, there were significant differences in the sex ratio of groups at the two sites: at one site males predominated and few pups were seen; on another, nearby, mothers and pups were regularly seen, although the site was also used by males. There was also evidence for segregation of the sexes outside the breeding season. Repeated observations of marked seals showed that seals used several different haul–out sites throughout the year, and that the seasonal changes in abundance at different sites resulted from individual changes in site–use. These changes in site–use are discussed in relation to feeding movements, breeding requirements and the physical characteristics of different sites.     

Spatial differences in breeding success in the pied avocet Recurvirostra avosetta: effects of habitat on hatching success and chick survival

I studied the breeding biology of pied avocets Recurvirostra avosetta in natural habitats (alkaline lakes), and in semi-natural sites (dry fishpond, reconstructed wetlands) in Hungary to relate habitat selection patterns to spatial and temporal variation in breeding success. Colonies were initiated earlier in semi-natural sites than in natural habitats, and earlier on islands than on the mainland. Hatching success was higher on islands than on the mainland, in semi-natural sites than in natural habitats, in colonies of at least 15 pairs than in smaller colonies, and for nests initiated earlier than later within a colony. Fledging success was higher in the wet years (1999–2000) than in the dry year (1998), decreased strongly by season in both habitats and increased with average daily temperature in the first week post-hatch in 1999–2000. Most pairs hatching young in semi-natural sites attempted to lead their chicks to feeding areas in natural habitats, whereas no such movement occurred in the opposite direction. Chick mortality due to predation was high during brood movements and only 23% of the pairs moving their young produced fledglings, compared to 43% for pairs remaining in semi-natural sites and 68% for pairs hatching and rearing young in natural habitats (total n=192 broods). These results suggest that semi-natural sites were more suitable for nesting, whereas natural habitats were more suitable for chick-rearing. The opposing trends in habitat-related breeding success between nesting and chick-rearing suggest sub-optimal habitat selection by Pied Avocets due to an incorrect assessment of the potential for successful reproduction of semi-natural sites, which may thus function as ecological traps.     

The influence of food supply on the breeding ecology of Kittiwakes Rissa tridactyla in Shetland

We measured the breeding performance, body condition, time budgets and foraging ranges of Kittiwakes Rissa tridactyla at Sumburgh Head, Shetland, in two years of contrasting food availability. Kittiwakes in Shetland generally feed their young almost entirely on sandeels, and fisheries data indicated that stocks of sandeels in Shetland waters were at least ten times higher in 1991 than in 1990. Fledging success of Kittiwakes was nil in 1990 and 68% of eggs laid in 1991, although clutch-size and hatching success were no different between years. Post-hatching foraging trips in 1991 were of comparable duration to those recorded at other colonies in conditions of good food supply (2–3 h), while trips recorded during incubation or post-hatching in 1990 were approximately three times longer on average than at corresponding stages of the breeding season in 1991. Radio-tracking data indicated that adults generally stayed within 5 km of the colony in 1991 but flew more than 40 km from the colony on each trip in 1990. Eggs were apparently not left unattended in either year, despite the fact that this required adults to incubate for periods in excess of 44 h in 1990. The extent to which adults were able to increase trip durations, foraging ranges and incubation shift lengths between years, while maintaining hatching success, indicates the degree to which Kittiwakes are normally buffered against adverse feeding conditions during incubation. Reduced nest attendance and lower body-condition of adults post-hatching in 1990, in conjunction with complete post-hatching breeding failure, indicate that adults were beyond the limits of their buffering capacity during chick-rearing in 1990.     

The breeding biology and aspects of the feeding ecology of brown noddies Anous stolidus nesting near Culebra, Puerto Rico, 1985–1989

We have studied brown noddies Anous stolidus breeding on Cayo Noroeste in the Culebra National Wildlife Refuge for the past five years (1985–1989). Daily visits during our residence there each year permitted collection of precise data on arrival chronology, egg-laying and incubation patterns, egg morphometries, chick growth rates, food delivered to chicks and breeding success. Over 150 adults are now individually colour-banded, and each annual chick cohort has been uniquely marked. Some features of the breeding biology of Culebran brown noddies were similar to those reported for pairs at Atlantic and Pacific Ocean colonies; these included duration of egg development, mass of newly hatched chicks, chick growth rates and fledging periods. Other features appear unique to Culebra. These include a tightly synchronous arrival (range of first egg dates: 29 April-4 May) and short egg-laying period (about six weeks), a consistently high breeding success each year (average 84% hatching success, 88% chicks fledged from eggs hatched, 0.79 chicks fledged per pair), and an unusually narrow range of food items (two fish species) taken by adults for their own needs and those of chicks. Observations of the direction taken by adults departing the colony suggest a predictable and productive foraging area along a prominent east-west shelf-break located c. 20 km to the north. We conclude that during these five years, brown noddies on Cayo Noroeste were not limited by food during the breeding season.     

Dispersion and site fidelity of breeding female grey seals (Halichoerus grypus) on North Rona, Scotland

Female grey seals ( Halichoerus grypus ) formed breeding aggregations on the island of North Rona, Scotland. Aggregations of females were associated particularly with gullies leading from the sea, leaving large areas of available space unoccupied. Changes in the degree of aggregation of females during the breeding season were similar in 1987, 1988 and 1989. Pronounced aggregations occurred in the early and late parts of each breeding season.
Of 67 breeding females marked in 1985, 62 (93%)) returned to N. Rona to breed in at least one season up to 1989, but 18 (27%) were present in all five years. Females came ashore up to 14 days before giving birth and 82% were observed first in the vicinity of their subsequent pupping site. Between 1985 and 1989, marked females which returned were faithful to their previous pupping sites, even when the previous pup had died. There was no evidence of a gradual change in the location of individual pupping sites over time. This pupping site fidelity may generate aggregations whose location, timing and composition is predictable.     

Annual productivity and individual female reproductive success in a Great Bustard Otis tarda population

The reproductive success of Great Bustards Otis tarda in north-western Spain was studied between 1987 and 1998, both at the population ( c . 700 adult females breeding in our study area) and the individual level (sample of 32 marked females). Overall productivity was low, with a population mean of only 0.14 chicks reared per adult female, and an average breeding success of 0.15 chicks per year in the sample of marked females, but interannual variability was high (0.04–0.29). Population productivity was positively correlated with winter (October–March) precipitation prior to each breeding season, and negatively correlated with the number of days of rain during the hatching period. High annual productivity resulted from a high proportion of females rearing two chicks. Reproductive success was higher in females older than 6 years than in younger birds. The proportion of females in the marked sample that failed in breeding after having bred successfully the previous season was significantly higher than the proportion of those that did not. Finally, females with a higher than average breeding success tended to breed successfully in years of both low and high population productivity, whereas those with lower than average breeding success did so only in years of high productivity.     

Cooperative enhancement of reproductive success in white-winged choughs

Summary White-winged choughs are a cooperatively breeding species which provide parental care to their young over an entire year. I traced the reproductive success of groups of white-winged choughs from the start of one breeding season to the next over 3 years. I examined the effect of helper number on timing of breeding, the success of each effort, the number of efforts made in a season, and the final reproductive success at the end of each year. Timing of commencement of breeding varied between years but was not related to group size. Early broods were not more successful than late broods. Nest building (July–September) commenced earlier in years which had high rainfall in July; choughs rely on rainfall for supplies of mud for nest construction. Most nest failures occurred gradually and were attributed to starvation of nestlings, although some sudden failures were attributed to predation. Large groups have more young by the beginning of the following season; this is due to higher fledging success and a greater likelihood of having second broods. Disappearance of young after fledging and during the transition to independence was not dependent on group size. Only groups of seven and above produce more than one young on average over the entire year; choughs provide one of the most marked cases for helpers enhancing the reproductive success of breeders. Large groups are virtually guaranteed of reproductive success over the whole year and grow more quickly than small groups. These results highlight the need to consider the effect of helpers over the entire period of reproduction and care of young, rather than just at fledging.     

Temporal and spatial variation in dispersal in the greater flamingo (Phoenicopterus ruber roseus)

We studied movements of individually marked greater flamingos (Phoenicopterus ruber roseus) born in the Camargue, southern France, between their two most important breeding colonies in the western Mediterranean (Camargue and Fuente de Piedra, Spain) from 1986 to 1992. The two sites differ in the frequency with which they offer suitable conditions for breeding. Flamingos have bred each year in the Camargue since 1974, but in only 12 of the past 22 years at Fuente de Piedra. Higher colony fidelity is thus expected in the less variable environment (Camargue), but if dispersal occurs competition might be an important factor causing this dispersal. Following years during which breeding birds in the Camrgue were disturbed (1988 and 1990) a higher proportion of adults changed colonies between breeding attempts (= breeding dispersal, 12.4%), while only 0.4% of flamingos breeding in the Camargue dispersed in the other years. As expected, flamingos breeding at Fuente de Piedra showed a higher rate of breeding dispersal (8.14%). No differences were observed between males and females. The importance of breeding failure as a factor causing breeding dispersal in flamingos was also confirmed by the movements of individual birds. The proportion of young flamingos that moved from their natal colony to start breeding at Fuente de Piedra (= natal dispersal) was independent of sex and age, but increased when breeding access to the Camargue colony was more difficult. However, natal dispersal was also higher in 1988 and 1990 (40.5%) than in the remaining years (1.2%), as was breeding dispersal. We discuss possible ways in which the increased natal dispersal among inexperienced birds could be linked with the increased breeding dispersal of adults in the same year.     

5. LIVINGSTONE IN NORTH-EASTERN RHODESIA

Sugg, M. St. J. 1974. Mensural and moult data from a breeding colony of Pied Kingfishers. Ostrich 45: 227–234.

Pied Kingfishers were ringed over two breeding seasons at a breeding colony on the Kenyan shore of Lake Victoria. Data were collected on weight, wing and bill length, injury and moult. Bill abrasion and breakage from nest excavation was found in both sexes and regeneration of worn and broken bills was recorded.

Adult birds of both sexes returned to the colony to breed but no juvenile was recaptured the year after hatching. Females are slightly larger than males (wing length) and both sexes showed a weight increase in the evening prior to roosting. Juveniles had shorter wings and bills than adults but their weights were similar. Their bills were short, soft, weak and the gape was salmon pink. No juveniles showed any moult. Adult moult records support the sequence and duration of moult suggested by Douthwaite (1971) though no overall decrease of moult activity was recorded during the breeding season.     

On the biology of the catfish Chrysichthys auratus (Geoffory) in the man-made Tiga Lake in Northern Nigeria

SUMMARY. (1) A study was made of the biology of Chrysichthys auratus , a bottom-feeding omnivore, in Tiga Lake, a man-made lake in Northern Nigeria.
(2) Ripe females were present from April to September, coinciding with the hot season and rise in water level. First spawning took place at age II in females. No ripe males were found, and this was attributed to their spatial separation. Absolute fecundity was related to length and ranged from 327 to 1466 in forty-six females.
(3) Hyaline rings on the centra of lumbar vertebrae were counted to age 235 fish. The first ring is laid down at the end of the second year, after which the rings were formed annually, probably during the breeding season. The fish live for up to 6 years. There is a good fit of the growth curve to the von Bertalanffy equation. Growth is slow and a smaller maximum size is attained than in rivers.     

Intermittent breeding in the short-tailed shearwater Puffinus tenuirostris

1.  At one colony of short-tailed shearwaters in Bass Strait, Australia, all birds breeding have been recorded individually each year for over 50 years. Among individuals known to be alive and to have bred before, 14% of each sex were not present at their breeding colony, on average, in any one year.
2.  A further 15% of males and 13% of females (a significant difference), known to have bred before, were present but not associated with an egg in any one year.
3.  Intermittent breeding was associated with pair bond breakdown and with a reduced ability to raise offspring, even when years of absence were allowed for.
4.  The frequency of attendance, laying and successful rearing of progeny to fledging increased with age in the early years of breeding. However, this was followed by a decrease, except in the case of laying frequency, which continued to increase throughout life.
5.  Our analysis indicated that intermittent breeding in short-tailed shearwaters did not result from individuals implementing trade-offs between the effort required for breeding success and their breeding life span. Rather, we suggest that individuals of higher quality are able to breed more frequently than others without any compensatory reduction in either their annual breeding success or their overall breeding life span.