Rates and consequences of relaying in Cassin's auklets Ptychoramphus aleuticus and rhinoceros auklets Cerorhinca monocerata breeding in a seasonal environment

We removed first eggs from early‐laying females to measure rates and consequences of relaying in Cassin's auklets Ptychoramphus aleuticus and rhinoceros auklets Cerorhinca monocerata at Triangle Island, British Columbia, Canada. Based on egg size and composition, the investment that Cassin's auklets made in first eggs was very close to that predicted from adult body mass, whereas rhinoceros auklets invested more. In both species, a high percentage of females relaid (90% of Cassin's and 87% of rhinoceros auklets). Breeding success declined weakly with later laying among control Cassin's auklet pairs, but pairs that we induced to relay bred more successfully than naturally late pairs, and similar to values predicted from laying dates of their first eggs. Their chicks also fledged heavier and younger than late control chicks, and similar to values in early control chicks, but followed the population‐wide seasonal decline in wing length at fledging. Nestling diets were dominated by Neocalanus copepods until late in the season, a sign that feeding conditions remained favourable until late. In contrast, rhinoceros auklet pairs induced to relay followed the population‐wide seasonal decline in breeding success, which was driven by a decline in hatching success. Pacific sandlance Ammodytes hexapterus, thought to be a preferred prey species, virtually disappeared from nestling diets in mid‐to‐late season, yet there was no seasonal decline in fledging mass. However, chicks from replacement eggs followed the declines among control chicks in both age and wing length at fledging. Despite the female having produced a replacement egg, and despite delayed breeding, there appeared to be little immediate consequence associated with relaying for Cassin's auklets, except for a tendency for their chicks to fledge with short wings. Consequences were more marked in rhinoceros auklets (greatly reduced hatching success, and having their chicks fledge with short wings), and this may have been due to the large investment made in eggs, and/or to delayed breeding. Results of this study show that attributes of Cassin's and rhinoceros auklets that lay at different times in the season can be important in driving seasonal declines in breeding performance, as found in studies on other Alcidae. They also show how decisions taken during the egg stage can have variable yet potentially important implications for fitness, even in relatively long‐lived species that lay single‐egg clutches.     

Seasonal variation in reproductive measures of tropical Roseate Terns Sterna dougallih previously undescribed breeding patterns in a seabird

Seasonal variation in egg-laying, egg size, hatching success, hatchling mass, fledging success and chick growth of Roseate Terms Sterna dougallii breeding on Aride Island (Seychelles), Indian Ocean, were studied in 1997 and 1998. I investigated to what extent two patterns, common in a range of species, were followed by tropical Roseate Terns: (a) seasonal decrease in clutch size, egg size and breeding success and (b) an increase in breeding success with increasing egg weight. In 1997 (a poor year), the earliest nesting birds laid significantly smaller eggs, and chicks were lighter at hatching than those of peak nesting birds. The mean clutch size, of 1.04 eggs, showed no seasonal variation and no 'b'-eggs hatched. In 1998 (a good year) the earliest nesting birds laid eggs of similar size and their chicks were of similar weight to those of peak nesting birds. Mean clutch size, of 1.25 eggs, increased significantly through the season and about 60% of the 'b'-eggs hatched. In 1997, hatching success was 57% whereas in 1998 it was 80%. In both years, breeding success declined significantly through the season. The fact that the earliest breeding birds laid smaller eggs in a poor year and smaller clutches in a good year is in marked contrast to a range of other species, and to temperate-nesting Roseate Terns. Egg volume explained about half of the variance in hatchling mass in both years, but only 15% of the variation in linear growth rate. Hatching date was the only variable with a significant effect on fledging success. Roseate Terns on Aride seemed to sacrifice egg size and clutch size for earliness of laying. Presumably it is a strategy of older birds to lay as early as possible and may be regarded as a response of tropical Roseate Terns to breeding under relatively poor, and seasonally declining, food conditions.     

Using logistic regression models to predict breeding success in male Adélie penguins (Pygoscelis adeliae)

Measures of breeding success are traditionally derived using the proportion of total nests that successfully reach strategic stages across a breeding season, such as pair formation, egg laying or fledging chicks. The use of logistic regression has recently become a popular tool in avian literature for identifying influential factors that predict nest and/or individual breeding success. In this study, we use logistic regression models to assess the importance of a range of factors affecting male Adélie penguin (Pygoscelis adeliae) breeding success during the 2002–2003 austral summer, when the presence of icebergs and extensive sea-ice meant overall breeding success of Adélie penguins was low (16/51 focal males had chicks reach fledging point). Logistic regression models for the early breeding season showed that good/average nest quality and central/middle nest location within the sub-colony were the best predictors of successful pair formation. Later, during incubation, the most successful males were those that not only returned earlier but were also heavier upon arrival and built nests of at least average quality. During the final stage when chicks had begun to fledge, the combined parameters of heavier male weight, early arrival time and good nest quality were the best predictors of breeding success. The logistic regression approach used here showed that the predictive ability of these parameters varied as the season progressed. However, most importantly, our logistic models fit the data well across all breeding stages.     

Egg size, composition and offspring quality in some Alcidae (Aves: Charadriiformes)

The composition of fresh and fully developed (pipping) eggs of four alcids, Razorbill, Common guillemot, Brünnich's guillemot and Atlantic puffin was examined. There were no differences in egg composition between the semi-precocial Atlantic puffin and the three species with "intermediate" developmental patterns. The absolute amount of yolk increased with egg size in fresh eggs, and the relative amount remained constant with egg size for Common guillemot and Razorbill, but decreased in the Atlantic puffin. In fully developed eggs chick weight and egg weight were closely correlated, and this was due mainly to larger eggs producing chicks with larger yolk reserves. Under some conditions chicks from larger eggs do better than those from smaller eggs. Several factors influence egg size; a comparison of first and replacement eggs laid by the same females showed that a maternal effect accounted for 60–90% of the variance in egg-size and that laying date accounted for most of the remaining variance.     

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.     

Great tits provided with ad libitum food lay larger eggs when exposed to colder temperatures

The amount of nutrients deposited into a bird egg varies both between and within clutches of the same female. Larger eggs enhance offspring traits, but as a tradeoff, laying large eggs also infers energetic costs to the female. Income breeders usually lay larger eggs later in the season, when temperatures and food availability are higher. Egg size is thus affected by the daily amount of energy available to produce an egg under cold conditions, but it is less well known in how far temperature exerts direct effects on egg size. We show that great tit females Parus major with access to ad libitum food and breeding in climate‐controlled aviaries varied their egg investments. The size of an individual egg was best predicted by mean temperatures one week pre‐laying, with females laying larger, rather than smaller, eggs under colder conditions. Eggs increased in size over the season, but not significantly over the laying sequence. The degree of daily temperature fluctuation did not influence egg size. In addition to a substantial between‐female variation, sisters were more similar to each other than unrelated females, showing that egg size does also reflect heritable intrinsic female properties. Natural variation in egg size is thus not only determined by energy‐limitation, but also due to females allocating more resources to eggs laid in colder environments, thus increasing early survival of the chicks. That the positive correlation between temperature and egg investments that is found in a natural population is reversed under ad libitum food conditions demonstrates that wild great tits tradeoff own condition with survival prospects of their chicks as a function of available food, not ambient temperature.     

Reproductive tactics of the ringed plover Charadrius hiaticula

Reproductive tactics of ringed plovers Charadrius hiaticula were studied at three localities in SW Sweden during five seasons. The usual clutch size is four, but removal experiments showed that females can produce five eggs in succession, with similar intervals between all eggs. High predation made mean breeding success per clutch low, 6.3% of eggs resulting in fledged young. Replacement clutches were common, and many pairs laid again after rearing their first brood to fledging. Egg laying spanned three months, much longer than for other waders in this region. Between years, reproductive success varied unpredictably with time of the season, but averaged over several years, the expected success was low and similar for the different parts of the breeding season. Chicks from late clutches had similar survival and recruitment as others. Because of the long breeding season and high rate of nest failure a female may produce up to five clutches of four eggs per season, containing in total about 3.7 times her own mass. Yearly local survival of adult females and males was estimated to 84.6 and 88.6%, respectively. Ability to produce many clutches with similar expected success throughout the season favours a long reproductive period, sometimes leading to double-brooding. Possible life-history trade-offs are discussed.     

Experimental manipulation of female reproduction reveals an intraspecific egg size-clutch size trade-off

A negative relationship, or trade-off, between egg size and clutch size is a central and long-standing component of life-history theory, yet there is little empirical evidence for such a trade-off, especially at the intraspecific level. Here, I show that female zebra finches (Taeniopygia guttata) treated chronically during egg formation with the anti-oestrogen tamoxifen lay smaller eggs (by 8%) but produce larger clutches (on average two eggs more) than controls. Decreased egg mass in tamoxifen-treated females was associated with a 50% decrease in plasma levels of the two yolk precursors, vitellogenin and very-low-density lipoprotein. Although tamoxifen-treated females laid more, smaller eggs (and had a higher total expenditure in their clutch), they did not differ from controls in the number of chicks fledged, the mass or size of these chicks at fledging, or the chicks' egg-production performance at three months of age. However, tamoxifen-treated females had lower relative hatching success: they laid more eggs but hatched the same number of chicks. Among individual tamoxifen-treated females, birds that laid the smallest eggs early in their laying sequence laid the largest number of additional eggs, that is, there was a negative correlation, or trade-off, between egg size and clutch size.     

Breeding success in a Houbara Bustard Chlamydotis [undulata] macqueenii population on the eastern fringe of the Jungar Basin, People's Republic of China

Nesting success and chick survival of a migratory population of Houbara Bustard Chlamydotis [undulata] macqueenii were studied during three consecutive years (1998–2000) in the Xinjiang province of north‐west China. A total of 45 nests was monitored and 85 broods comprising 227 chicks were captured, of which 82 chicks were radio‐tracked. Start of laying varied between 6 and 17 April between years but the laying mode fell consistently between 26 and 30 April. Mean clutch size was 4.0 (sd = 0.8) (range 2–6) for early clutches and 3.3 (sd = 1.1) for late clutches (range 2–5). The average nesting success was 0.588 (sd = 0.270) but great variations were observed between years –0.882 in 1998, 0.530 in 1999 and 0.351 in 2000. This was related to increased predation in 1999 and 2000, which is reflected by increased predator density (chiefly Corsac Fox Vulpes corsac and Long‐legged Buzzards Buteo rufinus). The overall hatchability, defined as the proportion of eggs hatched in successful nests was 0.839 sd = 0.238). The average brood size at hatching varied from 2.9 (sd = 0.8) to 3.3 (sd = 0.9) according to years, and no significant decrease in brood size was observed in the first 5 days post‐hatching. In 1999 and 2000 the brood size diminished sharply (14% and 27%, respectively) in prefledging chicks. A further severe decrease (37%) was observed in fledglings in 2000, probably due to predation by raptors. For the 3 years of the study, a successful female Houbara would bring on average 2.3 (sd = 0.9) chicks to fledging and would have lost 30.2% (sd = 14.9%) of its brood to adversity during the rearing process. The proportion of females that lost their entire brood was 0.181 in 1998, 0.708 in 1999 and 0.453 in 2000. For the 3 years of the study, only 55.3% (sd = 26.3%) of the females hatching eggs brought chicks to fledging. The overall chick production was 0.827 per breeding female per year and the probability of an egg laid producing a fledgling of 8 weeks old was 0.190.     

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.     

Differential predation of eggs in clutches of Glaucous-winged Gulls Larus glaucescens

Differential predation of eggs of Glaucous-winged Gulls Larus glaucescens was studied on Mitlenatch Island, British Columbia. The predation of single eggs from 3-egg clutches by conspecifics and Northwestern Crows Corvus caurinus was recorded to assess whether the predators selected eggs at random or showed a preference for a specific egg in the laying sequence. It was found that small and light-coloured eggs, characteristically the last-laid egg in the clutch, were taken significantly more often than the other two eggs in the clutch. As chicks derived from this third egg are least likely to fledge because of their small size and the effects of asynchronous hatching, the lightness of the third egg is adaptive in that it helps to focus attention on the small egg.     

Time-dependent reproductive decisions in the blue tit

Many breeding attempts in birds do not result in any fledged young due to predation on eggs or young. Consequently, the influence of time constraints on reproductive decisions are integrated parts of the reproductive behaviour of birds breeding within short, seasonal climate zones. In this study, I mimicked nest predation by removing blue tit (Parus caeruleus) clutches shortly after completion. Around 75% of the removed clutches were followed by a repeat clutch. Females producing their first clutch early in the season and females with an early onset of incubation in the laying sequence (an indication of high parental or territory quality) were most likely to initiate a repeat clutch. A trade‐off between the benefits of a repeat clutch and survival likely stopped late females in bad condition from investing more in the current reproductive season. Females producing a repeat clutch laid fewer eggs, had an earlier onset of incubation in the laying sequence and produced larger eggs than they did when producing their original first clutch. Eggs produced after the onset of incubation were especially large in the repeat clutches. Since food availability was presumably higher when the female produced her repeat clutch compared with her first clutch, females made a strategical decision when reducing clutch size, whereas onset of incubation and egg size may have been energetically constrained when producing the first clutch. Females that produced a relatively large clutch, had a relatively early onset of incubation, and laid relatively large eggs in their first clutch also did so when producing a repeat clutch, indicating that some of the variation in breeding parameters are due to differences in parental or territory quality. Differences between years in the temperature‐dependent development rate of caterpillars seem to affect the time constraints on breeding. A year with a predicted early seasonal decline in caterpillars resulted in short intervals between removal and relaying, small clutches and an early onset of incubation.     

BREEDING BIOLOGY OF THE GREY-FACED PETREL PTERODROMA MACROPTERA GOVLDI

The Grey-faced Petrel is a non-migratory winter breeder whose reproductive season occupies 9–10 months. Males spend more time in the burrows than females during the courtship period. Some females keep company with strange males, and may be fertilized by them, but subsequently share incubation with their mate of the previous year. The duration of the pre-laying absence of females is about two months, and of the pre-incubation absence of males about seven weeks. Since copulation is presumed to occur before this absence, these petrels seem to have evolved prolonged viability of the spermatozoa, though ovulation may take place some time before laying. Eggs are laid in late June or July but chicks are rarely reared from eggs laid after 14 July; effective laying thus lasts three weeks. The single egg is about 15·5% of the female's weight; she may be able to exert slight control over timing of oviposition. She may be required to incubate, if capable, for up to 14 days from laying but the male takes over, on average, after four days. There are three main incubation spells of 17 days' average duration, two by the male. These are of a duration such that there is usually a change-over near hatching. Incubation lasts about 55 days. There is competition for burrows, resulting in two-egg nests. Norway Rats take unattended eggs and young chicks and scavenge, but their predation (less than 10–35% of chicks per year) is not considered to be endangering the population. After initially more frequent feeds, chicks are fed approximately once a week by each parent. They do not become much heavier than adults and the growth rate is slow: about 120 days to departure. The ability to begin breeding in winter, atypical of petrels in this region, may be facilitated by three factors: improved availability of food resulting from longer nocturnal feeding time and reduced inter-specific competition; the ability to lay fertile eggs two months or more after copulation; and the brevity of the non-breeding season due to the relative proximity of a sufficient food supply.     

Rates and consequences of relaying in little auks Alle alle breeding in the High Arctic an experimental study with egg removal

Most seabirds have a small clutch size. Thus, replacement of a clutch after loss can make important contributions to an individual’s lifetime reproductive success. However, in the condition of short polar summer, relaying propensity may be time‐constrained. In this study, we investigated rates and consequences of relaying in a small High Arctic seabird, the little auk Alle alle. We performed an experiment in which we removed the single egg from 20 nests of early‐laying breeders. We measured relaying rates, and compared chick body mass and breeding success between the experimental and control nests. Despite the narrow window of the Arctic summer and the closely synchronized breeding, 75% of females produced a replacement egg just 2.7% smaller in volume than the first egg. This indicates that in little auks, the demographic effects of disruptions to breeding attempts (by predators, adverse weather or human activity) may be mitigated to some extent by replacement clutches. However, peak body mass and fledging body mass were lower in the experimental than the control chicks. This effect was rather a consequence of late hatching – chicks from replacement clutches followed seasonal decline in peak body mass and fledging mass. Finally, breeding success and chick survival up to 20 d in the experimental nests were respectively 34 and 37% lower than in the control nests. Thus, the quality and post‐fledging survival of chicks from the replacement clutches were probably lower compared to the chicks hatched from the first‐laid eggs.     

FACTORS AFFECTING THE EGG SIZE OF RED-BILLED GULLS LARUS NOVAEHOLLANDIAE SCOPULINUS

The factors influencing the egg size of the Red-billed Gull Larus novaehollandiae scopulinus were studied at Kaikoura, New Zealand, between 1964 and 1972. In two- and three-egg clutches there was a trend for the eggs to become smaller in the sequence of laying. Length, breadth and volume of eggs of one-, two- and three-egg clutches declined significantly as the season progressed. The size of eggs from single-egg clutches tended to be smaller than eggs from two-egg clutches laid at the same time. There were correlations between the proportions of one-egg and of three-egg clutches being laid at a given period and the mean egg volume of two-egg clutches. When the mean egg volume of two-egg clutches increased there was a corresponding increase in the proportion of two- and three-egg clutches laid. When the mean egg volume of two-egg clutches decreased there was an increase in the proportion of single-egg clutches laid. The egg size of the Red-billed Gull showed no direct correlation with the abundance or availability of food; the largest eggs were produced early in the season when food was in short supply. In spite of an increase in the food supply in the middle of the breeding season, birds laying at this time produced smaller eggs than birds which laid earlier in the season. However, early breeders which relayed at the peak in food abundance on average produced a larger replacement clutch than originals laid early in the season. It is suggested that the birds nesting early in the season are able to produce the largest eggs because they are the most efficient foragers for food, and those which nest later in the season produce smaller eggs, even at peak food abundance, because of their inefficiency or inexperience. Early breeders laying replacement clutches tended to lay larger eggs and larger clutches than birds which are producing their first clutches at the same time. Two-year-old females laid eggs which were significantly shorter than older aged birds while the breadth and volume of the egg increased with the age of the female up to the fifth year. There was a trend for females to lay larger eggs when mated with older rather than younger males. No statistical differences in egg size were detected between females changing or retaining the partner of the previous season. Female body weight and egg volume were positively correlated in females weighing less than 275 g but not for heavier females. It is suggested that the seasonal decline in egg size and clutch size results from a decrease in the availability of food and the ability of the individual to exploit the resource.     

Patterns of variation in size of Boat-tailed Grackle Quiscalus major eggs

Boat-tailed Grackle Quiscalus major eggs averaged 8^.09 g wet weight. Mean egg weight represented 8 05 % of female body weight. Based on egg weight, Grackle eggs have a shorter incubation period than the general passerine pattern.
Egg weight varied significantly with sequence of laying and between clutches in both two-egg and three-egg clutches. Last laid eggs weighed less than first laid eggs. The mean weight of the first two eggs laid in three-egg clutches did not differ from the mean egg weight of two-egg clutches. The average clutch of two and three eggs weighed 16.33 g and 24.16 g, respectively.
Mean egg weight varied between study colonies and with season. Grackles at East Lake laid eggs that weighed less than grackles at either Alligator Lake or North Lake. Eggs laid during March averaged less than eggs laid later in the season. The locality variation reflects the different timing of nesting between sites rather than food abundance. As nutrient provisioning increases with an increase in egg weight, the seasonal change in egg weight probably reflects improved feeding conditions for the female. This suggests that selection favours beginning laying before reserves of the female are sufficient to lay the largest egg possible.
The size of the young at hatching was correlated with egg size. Newly hate had young averaged 79.6% of fresh egg weight. Egg weight did not determine the probability of hatching or starvation of the young. Females do not appear to adjust egg size facultatively depending on the sex of the young. Eggs producing males and females did not vary significantly in weight, whether comparisons were made within or between clutches.     

Shiny Cowbird parasitism of a low quality host: effect of host traits on a parasite's reproductive success

ABSTRACT.   The reproductive success of parasitic cowbirds ( Molothrus spp.) varies among host species and is influenced by the degree of synchronization in timing of egg laying, the duration of parasite and host incubation periods, and the ability of hosts to incubate and rear parasite young. We studied the reproductive success of Shiny Cowbirds ( Molothrus bonariensis ) that parasitized the nests of Creamy-bellied Thrushes ( Turdus amaurochalinus ) in the Monte desert region of Argentina. Shiny Cowbirds frequently parasitized Creamy-bellied Thrush nests (60%), and most cowbirds synchronized egg laying with that of thrushes (79%). Most parasitic eggs (80%) hatched within 1 d of the hatching of the first host egg, and more than 91% of the eggs survived until the end of the incubation. However, only 60% of the cowbird eggs hatched and 52% of young survived. The proportion of Shiny Cowbirds eggs laid in Creamy-bellied Thrush nests that resulted in fledged young was 0.03, including eggs and young lost due to predation or desertion. Despite this low reproductive success, Creamy-bellied Thrushes were heavily parasitized by Shiny Cowbirds in our study area. Shiny Cowbirds may continue to parasitize these thrushes because of diffuse selection or because Shiny Cowbird chicks are more likely to fledge from Creamy-bellied Thrush nests in years or areas with greater food availability when brood reduction does not occur.     

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.     

Captive breeding for reintroduction: influence of management practices and biological factors on survival of captive kaki (black stilt)

An important component of the restoration strategy for the critically endangered kaki or black stilt (Himantopus novaezelandiae) is captive breeding for release. Since 1981 1,879 eggs were collected from wild and captive pairs, with birds laying up to four clutches. Eggs were incubated artificially and most chicks reared by hand until released as juveniles (about 60 days) or sub‐adults (9–10 months). Because survival in captivity is a significant determinant of the number of birds available for release, we wished to identify sources of variation in mortality to assess potential impacts of management on productivity. Hatchability was 78% for captive‐laid eggs and 91% for wild‐laid eggs. Survival of hatched eggs was 82% by 10 months of age for both wild and captive birds. Most egg mortality occurred early in incubation and around hatching: the timing of mortality was unaffected by whether birds were captive or wild, hybrid or pure kaki, or when eggs were laid. Heavier hatchlings showed higher initial survival, as did chicks from wild parents. Hatchlings from fourth‐laid eggs showed lowest survival, even though hatchling mass tended to increase with hatch order. Survival of chicks subjected to major health interventions was 69% after 4 months. No differences in survival were found between different genders, hybrids and pure kaki, hand‐reared or parent‐reared birds, chicks hatching early or late in the season, different seasons, different‐sized groups of chicks, chicks reared in different brooders, juveniles kept in different aviaries, and chicks from subsequent clutches. Birds subjected to minor health interventions were equally likely to survive as healthy chicks (82%). Survival was high despite aggressive management (quadruple clutching and collecting late in the season). Differences between captive and wild birds suggest further improvements could be made to captive diet. Wide variation in hatchability between parent pairs substantiates the practice of breaking up poorly performing pairs. Zoo Biol 0:1–16, 2005. © 2005 Wiley‐Liss, Inc.     

Aspects of the reproductive ecology of the Hamerkop Scopus umbretta in central Mali

In a 4-year study of Hamerkops in central Mali, territories of breeding pairs overlapped and most clutches were laid in the late rainy or early dry seasons. Clutches averaged 4–8 eggs. Different pairs laid eggs having, on average, different dimensions. Eggs laid in the late rainy and early dry seasons were larger than those laid at other times. Eggs weighed, at 27-8 g, about 6% of adult female weight. Hatching success was 79% with a fledging rate of eggs hatched of 53%. Successful nests fledged 27 young but for all completed clutches the fledging rate was 0'9. Young hatched early in the sequence had a higher survival rate than those hatched later. Mean age at fledging was 47 days. Pairs fledged between 0–5 and 0–9 young per year.