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.     

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.     

THE BIOLOGY OF THE WAVED ALBATROSS DIOMEDEA IRRORATA OF HOOD ISLAND, GALAPAGOS

As a nesting species, the Waved Albatross Diomedea irrorata is restricted to Hood Island in the Galapagos archipelago where 12,000 pairs bred in 1971. Outside the islands the species occurs over the northern parts of the Humboldt Current. Two colonies were studied in detail (1970–1971). At the start of a season, males returned first to the colonies and defended a small territory. Copulation occurred without any elaborate ceremony and the female spent little time on land before laying. There was no fixed nest-site, even within a season, and birds moved their eggs considerable distances. This resulted in heavy egg losses. Younger birds bred later than older birds and laid longer but narrower eggs. The average incubation spell varied from four to five days at the extremes of the incubation period to 19 days in the middle. The average incubation and fledging periods were 60 and 167 days respectively. Pairs which lost an egg sometimes adopted the abandoned egg of another bird and successfully reared the chick. Most pairs nested in both seasons. Nesting success was extremely variable, both between years and between colonies. Between 1961 and 1971 at Punta Suarez, virtually no young were reared in four seasons. Even in 1970–71, where nesting success was good, some groups of birds deserted their eggs en masse whereas in neighbouring areas up to 80% of the pairs reared young. The main foods of the young were squid and fish. Birds did not moult wing and tail feathers at the breeding colonies, and about 50% retained some primaries for more than one season, suggesting that successful pairs had difficulty in fitting in a complete moult between breeding attempts. Old feathers were normally found among the inner primaries and at the next moult were preferentially replaced, though adjacent newer feathers were sometimes retained for another season. Some birds bred in their fourth years, but most not until a year or two older. Immatures were present at the colonies late in the breeding cycle, the youngest returning latest and remaining until the last young fledged. Survival of adults and young averaged at least 95% and 93% per annum over many years. Adults and young ringed in 1961 survived equally well. The significance of the timing of the return of immatures and of the large-scale desertion of eggs, apparently not due to food shortage or disturbance, is discussed.     

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.     

Effects of supplementary food on density-reduced breeding in an African eagle: adaptive restraint or ecological constraint?

Increased population density often reduces reproductive output in breeding birds, but the underlying mechanisms (adaptive restraint v reduced food resources) behind decreased productivity are poorly understood. Here I correlatively and experimentally investigated the roles of food, breeding density, latitude, altitude and rainfall in limiting productivity of Wahlberg's Eagles Aquila wahlbergi throughout Africa. Breeding success in equatorial and subtropical Africa (0°–30°S) was highly density-dependent but showed no latitudinal or rainfall-related trends. Pairs in dense populations produced half as many young annually as pairs in low-density populations. Density (but not rainfall or latitude) also explained much of the geographic variation in the mean proportion of pairs attempting to breed each year and the incidence of two-egg clutches.
Breeding within populations was consistent with these density-dependent trends: incidence of two-egg clutches increased in a declining population, and productivity was inversely related to breeding density and rainfall combined. To determine if reduced food resources accounted for reduced output in dense populations, eight pairs were food supplemented: supplementary food failed to induce nonbreeding pairs to breed: nor did it induce earlier laying or increase egg size or clutch-size. Population density itself was unrelated to two correlates of food resources, rainfall and latitude. I conclude that population density influences most aspects of breeding in Wahlberg's Eagles, and reduced food resources do not appear to explain these trends. Hence, adaptive restraint may account for decreased annual reproduction in this species.     

The effect of food on laying date and clutch-size in Tengmalm's Owl Aegolius funereus

The breeding of Tengmalm's Owl Aegolius funereus was studied at Umeå, Sweden, during the 1984–85. Mean clutch-size was one egg larger in 1984 than in 1985 despite the later laying in 1984. The difference in clutch-size was related to a better food supply in 1984. Daily weight increase of females during the prelaying period showed a high negative correlation with laying date in 1985, and a high positive correlation with clutch-size independently of laying date in 1984–85. This suggested that food eaten before and during laying had a great and direct influence on both laying date and clutch-size. Many females increased in weight during laying and most others decreased only moderately (relative to egg weight), suggesting that body reserves were not a main source for egg production.
Late breeding females were provided with extra food during the prelaying and laying periods in 1985. Fed females weighed more, bred eight days earlier and laid one more egg than controls. At the same laying dates in mid season, and after heavy snow-fall, clutch-size and female weight were larger in the fed birds than in controls, but this was not so near the end of the laying season. Although the earliest of the fed late breeders weighed more, and probably were less restricted by food availability just before or during laying, they did not lay more eggs than did early breeders. This result suggested some limitation on clutch-size that could not be overcome by the supplementary feeding. Weights of females during laying did not show any consistent relationship with clutch-size during successive laying date intervals, suggesting that clutch-size was not directly related to body condition.     

Individual quality and food availability determine yolk and egg mass and egg composition in tree swallows Tachycineta bicolor

Variation in egg size and composition can have important consequences for the quality of offspring. We investigated the factors influencing the yolk mass and egg mass of tree swallows breeding in Ithaca, NY. Using a nondestructive technique to estimate yolk mass via standardized digital-candler photographs, we compared yolk size and egg size of tree swallows Tachycineta bicolor in response to variation in food availability and individual quality. Insect availability one to three days prior to laying, but not four to six days, predicted yolk mass, while insect availability two to three days prior to laying predicted total egg mass. This suggests that, while eggs are formed over longer periods, food availability closer to time of laying has the greatest influence on egg size. These results were supported by collected eggs, as yolk rings revealed that tree swallow eggs are formed over 5–6 days. There was an influence of female quality as well, with early laying birds, independent of food availability, laying larger eggs. Eggs laid later in the laying sequence had larger yolks and greater egg mass. Overall, variation in egg quality appears to be due to a combination of environmental conditions, reflected in food resources, individual quality, and allocation tradeoffs during the laying period.     

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.     

Testing the relationship between clutch size and brood size in the Coot (Fulica atra)

The time between egg laying and chick fledging is of crucial importance for the survival of young birds. I analyzed breeding output at consecutive phases of growth of young Coots (Fulica atra) relative to the clutch size and laying date. Considering the specific breeding biology of the Coot, I tested whether chick survival reveals clutch size-dependent variability. Clutch size did not affect hatching success; it only affected brood size, and that merely temporarily. During the first 20 days after hatching, i.e. during the time of the highest chick mortality, birds with larger clutches lost chicks at a higher rate. As a result, the number of fledged chicks was independent of the initial number of chicks, and pairs with different clutch sizes had a similar number of fledglings. The laying date had no effect. This pattern of age-related chick survival points to the greater role of the type of chick growth (semi-precocial) and behavior in their survival.     

Larval sensory abilities and mechanisms of habitat selection of a coral reef fish during settlement

In most animals, siblings from a given reproductive event emerge over a very short period of time. In contrast, many species of birds hatch their young asynchronously over a period of days or weeks, handicapping last-hatched chicks with an age and size disadvantage. Numerous studies have examined the adaptive significance of this atypical hatching pattern, but few have attempted to explain the considerable intrapopulation variation that exists in hatching asynchrony. I explored proximate determinants of hatching asynchrony by monitoring 112 Burrowing Owl (Athene cunicularia) nests in the grasslands of southern Saskatchewan, Canada, over 4 years. Age disparities between first- and last-hatched siblings (i.e., hatching spans) varied considerably, ranging between 1 and 7 days (mode = 4 days). These hatching spans increased with increased hatching success. Hatching spans also increased with larger clutches, but the increase was less than predicted given the increased time required to lay more eggs. Hatching span was unrelated to number of prey cached in the nest during egg laying (an index of food availability), and was unaltered by a year of super-abundant prey. Furthermore, pairs given extra food during laying had hatching spans equal to those of unsupplemented control pairs. These results were inconsistent with both the energy constraint and facultative manipulation hypotheses, which predict that hatching asynchrony should vary with the level of food during laying, when incubation onset is determined. Burrowing Owls were apparently free of food limitation early in breeding, yet may not have been able to optimize hatching spans because food conditions during laying were largely unrelated to food conditions during brooding. Thus, one of the premises for facultative manipulation of hatching asynchrony—that laying females are able to forecast post-hatch food conditions—may not have been met for this population of Burrowing Owls.     

BREEDING PERFORMANCE OF PUFFINS FRATERCULA ARCTICA IN RELATION TO NEST DENSITY, LAYING DATE AND YEAR

The paper presents data on the breeding and predation of Puffins in two areas of different nest density within a single colony on Dun, St Kilda group, Outer Hebrides in 1973-78.
Within a season birds laying early had a slightly higher nesting success than birds laying late, but laying date had little influence on the peak and fledging weights of young. The main disadvantage in late laying was a reduced chance of relaying if the first egg was lost.
Breeding success and chick weights varied from year to year. The 1974 season was the least successful with the lowest nesting success, lowest frequency of feeds, lowest calorific value of feeds, lightest chicks and slowest growth. Overall breeding performance was not related to the annual mean laying dates.
In all years pairs nesting in the area of high nest density did better than pairs nesting at low density. The effect is attributed to differential predation and disturbance by predatory gulls. At least 4.2% of adult Puffins breeding in the area of low burrow density were killed by gulls each breeding season; this is higher than the total annual mortality found in three other studies. Only 0.9% of adults from the high density area were found killed. The subpopulation in the low density area cannot survive without much immigration, yet there is no evidence that this happens.     

Seasonal declines in reproductive success of the common tern Sterna hirundo: timing or parental quality?

Reproductive success declines over the course of the breeding season in many bird species. Two categories of hypothesis have been evoked to explain this decline. The “timing” hypothesis suggests that seasonal declines in breeding success are attributable to the date of laying. The “parental quality” hypothesis suggests that seasonal declines result from the fact that young, inexperienced, or low quality birds breed later in the season. To evaluate the relative importance of timing and parental quality, egg exchanges and removals were used to manipulate hatching dates of common terns Sterna hirundo. Indices of quality, attendance, provisioning rates, and reproductive success of birds in three experimental groups (delayed hatch pairs, advanced hatch pairs, and pairs induced to relay) were compared to those of date‐matched controls. Pairs that hatched chicks early raised more chicks than pairs hatching chicks late in the season, regardless of initial laying date. This suggests that hatching chicks early is advantageous in itself. Our results, however, also support the parental quality hypothesis. There was a significant negative relationship between natural laying date and fledging success, independent of hatching date. Differences in chick growth and survival suggest that higher quality adults may be able to compensate for the disadvantages of late hatching dates and achieve similar reproductive success to that of pairs hatching chicks early. We found that pairs hatching chicks late in the season were subject to more incidents of kleptoparasitism than those hatching chicks early. This may be a proximate factor contributing to seasonal declines in reproductive success for common terns, although such a mechanism would not be likely in non‐colonial species. Failure to control for egg quality may have biased the results of some prior egg exchange experiments. Additionally, altered cost of incubation may be an unavoidable confounding factor in studies designed to manipulate timing of breeding.     

Components of productivity in black-legged kittiwakes Rissa tridactyla: response to supplemental feeding

In contrast to the high productivity of black-legged kittiwakes in Britain, kittiwakes at many colonies in Alaska have failed chronically to reproduce since the mid 1970s. To determine if food is limiting productivity and, if so, at what stages of nesting food shortages are most severe, in 1996 and 1997 we supplementally fed kittiwakes nesting on an abandoned building. The effects of feeding were stronger in 1997 than in 1996, possibly because naturally occurring prey were of poorer quality in 1997. Consumption of supplemental herring declined as egg laying approached then increased slowly during incubation and more rapidly after hatching. All of the six components of productivity we studied were improved by supplemental feeding to some degree. Supplemental food did not significantly alter laying success in either year, although fed pairs bred at slightly higher rates than unfed pairs in 1997, the poorer food year. In 1996 and 1997, extra food noticeably increased clutch size and hatching success, but significantly so only in 1997. Fledging success and productivity were substantially augmented by feeding in both years. Fed pairs fledged twice as many chicks per nest as did unfed pairs in 1996 and three times as many in 1997. Fed and unfed pairs lost most of their potential productivity through the inability to hatch eggs, and secondarily because of their poor success at raising chicks. The benefits of supplemental feeding did not carry over from one stage of breeding to another. Pairs cut off from supplemental food after laying or hatching performed similarly to pairs that had not been previously fed. This study provides benchmark values of breeding performance attainable by kittiwakes in Alaska under optimal conditions. These values are comparable to highly productive colonies in Britain and suggest that differences in life-history characteristics between Pacific and Atlantic breeding populations are primarily controlled by food supply.     

Annual and seasonal reproductive trends in the Lesser Spotted Woodpecker Dendrocopos minor

Annual and seasonal variation in reproductive timing and performance were studied in a population of the Lesser Spotted Woodpecker Dendrocopos minor over 10 years in southern Sweden. The median laying date of the first egg varied by up to 17 days between years, being generally larger than the variation of laying dates within years. Neither clutch size, brood size in successful nests, fledging success in successful nests nor mean nestling weight differed significantly between years. There was no trend for mean clutch size to vary between early and late years. In spite of a more than threefold variation in population size, no reproductive variable demonstrated an apparent density-dependence. Within the season, clutch size declined steeply with increasing clutch initiation date, whereas fledging success and nesting success did not, leading to a trend in brood size almost identical to the trend in clutch size. The survival prospects of fledged young declined with increasing clutch initiation date, and it is argued that the clutch size laid is a strategic adjustment to laying date. Out of 124 breeding attempts, 34% did not produce fledged young. In 9% of the breeding attempts, pairs laid no eggs. At least 20% of the breeding attempts failed after egg-laying. The most common cause of breeding failure was loss of the breeding partner followed by nest abandonment (40% of the failures). Only 16–28% of the failures were due to predation on the nest. Most complete failures, and also partial losses from nests, occurred at the early breeding stages. It is argued that the early nestling phase may be a critical stage, which the woodpeckers adjust to coincide with the seasonal food peak, explaining the strikingly late breeding season compared with other non-migrant species.     

THE BREEDING OF THE GREATER FLAMINGO AND GREAT WHITE PELICAN IN EAST AFRICA

This paper brings together observations on the breeding of the Greater Flamingo Phoenicopterus ruber and Great White Pelican Pelecanus onocrotalus, mainly at Lake Elmenteita, Kenya, 1951–1971. The Greater Flamingo bred at Lakes Elmenteita and Nakuru in 11/21 observed years and at Lakes Natron and Magadi in 5/12 observed years. On average, it breeds about every second year, but a succession of breeding years is followed by several years in which no breeding occurs. A history of 21 years' breeding at Lakes Nakuru and Elmenteita is given. At Elmenteita three sites have been used, the main site in every breeding year, the others less often. The number of pairs breeding in any year has varied from 500–9,250, but in 1968 flamingos bred three times, involving perhaps 8,500 pairs which made about 15,700 nests, some pairs perhaps laying twice or even thrice in a year. Losses of eggs (38.2% overall) were caused by rising water (16.2%), competition for nest space with Great White Pelicans (6.9%, after 1968 only), human interference (3.5%), Marabou Stork predation (1.8%) and other natural causes (9.8%). Losses among chicks totalled 68.3% overall and were mainly due to Marabou Storks (36.5%), undiagnosed disease (8.6%), and rising water (6.6%). Disease caused serious loss only in 1966, and after 1968 losses from Marabous rose from 2.7% to 76.5%, resulting in an increase in overall mortality from 48.7 to 92%. This was perhaps associated with the establishment of a fish factory at Lake Naivasha. When attacking flamingo colonies Marabous did not actually eat many eggs or chicks, but simply caused wholesale desertion by alarming the flamingos. In 1968 total desertion of a colony of 4,500 pairs was caused between 18 and 26 March by a maximum of 17 Marabous, and similar wholesale desertion was caused in later years. The overall breeding success among Greater Flamingos at Elmenteita was about 19% of eggs laid, but without the excessive post-1968 Marabou predation would have been about 30%. At such a rate Greater Flamingos require at least 24 years of adult life to replace themselves, but if the mortality caused by Marabous since 1968 continues they will require about 58 years, and the population will inevitably decline. Breeding success at Lakes Magadi and Natron has been higher, about 44% of eggs laid; but figures available are much more approximate than at Elmenteita. Some new data on display, nest-site selection, laying dates, clutch-size, hatching and creche behaviour are given for the Greater Flamingo. The Great White Pelican bred at Lake Elmenteita from 1968 to 1971 without a break, some birds laying in every month, but with reduced laying November–December. They bred on the same islands as, and in association with the Greater Flamingo, and caused heavy losses among the latter, not through aggressiveness, but simply because of their superior size and weight. Although food supply must ultimately have controlled the pelican's ability to breed, an adequate food supply was available for 6 years before they did and continued after they had ceased. Their breeding was finally triggered by the Greater Flamingo colonies, with which the pelicans associated. When a flamingo colony was deserted because of Marabou Storks the pelicans, unafraid themselves of the Marabous, also deserted. They also associated with, and wiped out, a colony of Sacred Ibis. From July 1968 to June 1969 about 2,600 pairs of pelicans bred at Elmenteita, rearing about 2,200 young to the flying stage. The breeding colony apparently comprised most of the adults from Lake Nakuru and Lake Naivasha, the main feeding areas. From July 1968 to January 1971 certainly 7,200 and probably 8,000 pairs of pelicans bred at Elmenteita. Some pairs may have bred twice or thrice in this period. Breeding ceased suddenly in January 1971, eggs, and small and large young being alike abandoned for no established reason, although food supply was certainly still plentiful. Additional information on pair formation, incubation and fledging periods, nest-relief, etc. is given. The best available record of the incubation period is 35–36 days. Nest relief takes place on average about once every 48 hrs, and is dependent on thermal activity enabling the pelicans to soar. At Elmenteita large young ate quantities of putrefying matter, including the corpses of other young pelicans. They also ate living young hatching from eggs, and up to 14 days old. Touch probably plays an important part in helping them to locate possible food in opaque water.     

Reproductive assessment of the great hornbill (Buceros bicornis) by fecal hormone analysis

The population of great hornbills (Buceros bicornis) in the United States is rapidly aging, and captive breeding efforts have not met population managers' expectations for a sustainable captive group. Little is known about the reproductive physiology of these birds. This study reports the first data on the re‐productive endocrinology of the great hornbill. The hormone profiles of the only pair of these birds that hatched a chick in the 1999–2000 breeding season are compared to the profiles of six other pairs of hornbills, from different institutions in the United States, that did not reproduce successfully that season. The study investigates the estradiol, corticosterone, and testosterone profiles of these seven pairs of birds, establishing a base of knowledge from which endocrine data may be used to improve the success of captive breeding programs. The estradiol profiles from this study indicate a difference in hormonal patterns between laying and non‐laying female great hornbills. Egg‐laying females had significantly higher estradiol concentrations during the breeding season than the non‐laying females (P<0.003). Testosterone concentrations of the males were not significantly different between the mates of egg‐laying and non‐egg‐laying females. The corticosterone concentrations tended to be lower in the females that laid eggs vs. the non‐egg‐laying group. The males of the egg‐laying pairs showed a significantly lower (P<0.036) corticosterone concentration than the non‐egg‐laying male pairs. This, combined with the extremely low corticosterone levels (compared to the other birds in the study) of the pair of hornbills that hatched a chick in the 1999–2000 breeding season, suggests that adrenal activity may play a role in the reproductive failure of some captive great hornbills. Zoo Biol 22:135–145, 2003. © 2003 Wiley‐Liss, Inc.     

Effects of egg size, parental quality and hatch-date on growth and survival of Common Tern Sterna hirundo chicks

We examined the relative contributions of egg size, parental quality and hatch-date to growth and survival of second-hatched chicks (those chicks making the greatest contribution to differences in productivity among pairs) by exchanging clutches among nests of Common Terns Sterna hirundo matched for lay-date (range 13 May to 9 June). The mass of a second-laid egg in an exchanged clutch ranged from 17.70 to 23.80 g. Growth and survival were studied during three periods: early (days 0–3), middle (days 3–12) and late (days 12–25). Both egg mass and hatch-date were important predictors of hatchling mass (positive relationships), although there was no seasonal trend in egg mass. During the middle period, hatch-date was a significant predictor of mass gain and survival (inverse relationships). After controlling for hatch-date, other indices of parental quality made only small contributions to chick mass gain and survival. Our results suggest that although breeding early generally leads to greater overall survival of chicks, several important interactions among egg 'quality', parental quality and early laying may affect breeding success under specific conditions.     

Increased survival and breeding performance of double breeders in little penguins Eudyptula minor,New Zealand: evidence for individual bird quality?

The little penguin Eudyptula minor is unique among penguin species in being able to fledge chicks from two clutches in one breeding season. Pairs laying two clutches in a given season make a higher reproductive investment, and may be rewarded by a higher reproductive success as they may raise twice as many chicks as pairs laying one clutch. The higher effort made by pairs laying two clutches could correlate negatively with survival, future reproductive performance or offspring survival, indicating a cost of reproduction. Conversely, a positive relationship between the number of clutches produced in a given breeding season and survival, future reproductive performance or offspring survival would indicate that birds laying two clutches belonged to a category of birds with higher fitness, compared to birds laying only one clutch in the season. In this study we used a long‐term data set taken from an increasing population of little penguins in Otago, SE New Zealand. We modelled the relationship between the number of clutches laid in a breeding season and survival probability, reproductive performance in the next breeding season and first year survival of offspring using capture‐recapture modelling.
Birds laying two clutches produced 1.7 times more fledglings during a breeding season than pairs laying one clutch. We found that birds laying two clutches had a higher probability of breeding in the following breeding season, a higher probability of laying two clutches in the following breeding season and a higher survival probability. There was no overall difference in post‐fledging survival between the young of birds producing one clutch and the young of birds producing two clutches. However, the survival of young of single clutch breeders declined with laying date, whereas the young of double clutch breeders had the same survival rate irrespective of laying date. For a subset of data with birds of known age, we found evidence that the probability of laying two clutches increased with age. However, there were also indications for differences among birds in the tendency to lay two clutches that could not be attributed to age. We tentatively interpret our results as evidence of quality difference among little penguin breeders.     

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

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

Supplementary feeding increases Common Buzzard Buteo buteo productivity but only in poor‐quality habitat

Temporal heterogeneity in the effects of food supply during the breeding season on the productivity of the Common Buzzard Buteo buteo was investigated in a supplementary feeding experiment. Pairs were fed artificially (1) before egg‐laying, (2) after chicks hatched and (3) continuously throughout the season, and compared with (4) unfed controls. Pairs fed before egg‐laying had marginally larger clutches than those not fed, but lay date, egg volume and weight, brood size and hatching success were unaffected. Territorial quality had far greater effects, with pairs nesting in low‐quality habitats (bog, scrub and semi‐natural grassland) laying later and having lower hatching success, smaller broods and fewer fledglings than those in more productive agricultural landscapes. Supplementary feeding after egg hatching neutralized the negative effect of poor habitat, resulting in fed birds having significantly more fledglings. This study emphasizes the importance of food availability when provisioning chicks in suboptimal habitats and has implications for the success of diversionary feeding in reducing game losses to Buzzards.