Breeding biology of the Barn Owl Tyto alba in central Mali

Data were obtained on 178 clutches of African Barn Owls in central Mali from four breeding seasons during 1979–1983. Significantly more clutches were laid in 1979–1980 and significantly fewer in 1980– 1981 than the average for the 4 years and there were significantly more clutches laid in the middle period of the annual breeding season. The egg volume was significantly smaller at the beginning of the breeding season and significantly larger in the middle than the overall mean with eggs of second clutches being larger than those of first clutches. The clutch size was 605 eggs of which 479 hatched. The number of young fledged per successful nest was 319 and was 1 83 for all nesting attempts. The month was the only variable shown to affect significantly the clutch size, eggs hatched and fledging rate, the highest success rates being associated with the middle of the breeding period. The average interval between the hatching of eggs was 2–31 days. Survival rates (47'1%) to fledging were significantly affected by year (1981–1982 being the least) and month (mid-season birds the best). The order of hatching significantly affected age at death or disappearance, the first-hatched birds surviving the longest. The year significantly affected age at fledging, the young from the year in which most clutches were laid leaving the nest at the youngest age and those associated with the year having the least number of clutches remaining in the nest the longest. The month of hatching also affected fledging age, birds at the extremes of the breeding season fledging at older ages. The discussion compares these data with those from elsewhere.     

CLUTCH-SIZE, INCUBATION AND HATCHING SUCCESS IN THE HOUSE SPARROW AND TREE SPARROW PASSER SPP. AT OXFORD

Clutch-size, incubation and hatching success were studied in P. domesticus and P. montanus in 1961 and 1963–64 at Oxford. The most frequent clutch-size was four eggs in P. domesticus and five eggs in P. montanus . With one exception, colonies of P. domesticus showed no significant annual or local variations in its mean clutch-size; in P. montanus , however, there were significant annual variations in the mean clutch-size. Both species showed a seasonal increase followed by a decrease in their mean clutch-sizes.
Partial incubation occurred during the laying period of the clutch; sufficient incubation for continuous development of the embryo was apparently achieved when the last egg had been laid in clutches of two and three eggs in P. domesticus and in clutches of four eggs in P. montanus , but when the penultimate egg had been laid in larger clutches of both species. On average, hatching in P. domesticus occurred more or less synchronously in all eggs in clutches of two and three eggs, and in all eggs except the last one laid in larger clutches; the last egg in the larger clutches hatched up to a day after the others. It is suggested that this pattern of hatching was brought about by the pattern of incubation during the laying period.
P. domesticus had a lower hatching success than P. montanus , probably because fewer of its eggs were fertile.     

BREEDING NUMBERS AND REPRODUCTIVE RATE OF EIDERS AT THE SANDS OF FORVIE NATIONAL NATURE RESERVE, SCOTLAND

The total population of Eiders at the Sands of Forvie National Nature Reserve, Aberdeenshire, increased from about 3000 birds during 1961–63 to about 4800 birds in 1964–70. At the same time the estimated number of breeding pairs increased from about 1200 in 1961–63, to about 1800 in 1965–68 and to 2000 in 1970. The mean size of completed clutches was 4.4; the number of eggs in a clutch showed a significant decline during the season from an average of 4.7 to 3.4 in most years. The overall hatching success was 63%, but there was a decline with season as a result of increased losses to predators. In 1964 and 1969 laying was retarded, mean clutch-size reduced, and the number of birds attempting to lay dropped by half, due to inclement weather just at the onset of laying. In 1969 the eggs produced were smaller than in other years, and hatching success was lower. The female incubates without feeding, and is required to store sufficient energy, in the form of fat, to last her through the 26 days in addition to producing her clutch of eggs. An hypothesis is advanced which relates the feeding efficiency of the paired female immediately prior to laying to her egg production and incubating pattern, and which offers an explanation of the decline in clutch-size and hatching success evident during the season. In most years less than 5% of hatchlings survived to fledge; high survival was recorded in 1963, 1968 and 1970. Following the high production in 1963 the population increased in 1964 and the breeding stock in 1965 (females start to breed at two years old). The corresponding changes, following good breeding in 1968, were not so clearly detected because of unusually high adult mortality from oil pollution at the winter grounds in 1968 and the ‘late’ breeding year in 1969. The breeding stock in 1970, however, was higher than in any other year. This population continues to grow apparently as a result of its own reproductive output checked only by low productivity in most years and occasional high adult mortality.     

Diet dimorphism' within an Oystercatcher Haematopus ostralegus population– adaptive significance and effects on recent distribution dynamics

Of the 50 or so Oystercatcher pairs breeding on Skokholm in 1965 and 1966, 15% bred near the shore and 85% bred inland; 28% fed their young on limpets, and 72% on terrestrial arthropods and earthworms. 'Terrestrial-feeders' were three times more successful than limpet-feeders in fledging young, irrespective of variation in initial brood size and of hatching date but depending on differential losses to gull predation. This is because parents' vigilance is relatively low in limpet-feeders' territories, and limpet-fed young are on average undernourished and hence behave relatively carelessly. Since the 1930s the population of shore-breeding gulls has increased considerably on Skokholm. A certain increase of the Oystercatcher population during this period seems to have been associated with a shift inland, until the island became saturated at the late 40s. It is proposed that the increase in shore-breeding gulls has triggered a genetically based behavioural change in Oystercatchers, which has been responsible for the shift to inland habitats and to terrestrial food, which promotes a population increase. This interpretation for events on Skokholm may be applicable for the general trend of habitat shift and population increase of other British populations.     

The pair bond and divorce among Oystercatchers Haematopus ostralegus on Skokholm Island, Wales

Mate changes of Oystercatchers were followed over a 15-year period. In 8.0% of instances, the pair split up between breeding seasons even when both members bred the second season. The likelihood of divorce was highest for birds aged five years, the modal age of first breeding. After divorce birds did not select an older or younger mate when re-pairing. Low hatching success increased the chances of divorce. Divorce was not followed by significant changes in clutch-size, laying date or nesting success (although females tended to be more successful).     

Selection for synchronous breeding in the European starling

Colonial birds often demonstrate considerable breeding synchrony. In southern Sweden the semi-colonial European starling initiated the vast majority of clutches within one week. Laying dates were positively skewed so that many birds initiated clutches at similar dates early in the season. Breeding was further synchronised by a particularly strong clutch-size reduction equivalent to one third of an egg per day during the first part of the breeding season. The decline in clutch size with season also held true for separate age-classes of females, for individual females laying at different times at different years and for individual females laying at different times the same year. Trends in breeding success during nestling rearing were unlikely to explain the high degree of breeding synchrony or the seasonal decline in clutch size; nestling survival and growth were weakly related or unrelated to reproductive timing. In contrast recruitment success of fledged offspring declined sharply with season. Even within the synchronous laying period, defined as clutches initiated during the first week each year, local recruitment success declined. It is suggested that the early seasonal decline is caused by selection for synchronous fledging permitting the immediate formation of flocks after fledging, whereas the late seasonal trends may be caused by either population differences in female quality or deteriorating conditions for raising young.     

FACTORS AFFECTING BREEDING OF RAZORBILLS ALCA TORDA ON SKOKHOLM

A study of the breeding biology of the Razorbill was carried out on Skokholm (South Wales) during 1971-73. Birds ringed or colour ringed before the study began provided additional information upon the effects of age on breeding. Mean laying date was delayed in 1972, compared with 1971; the effect is attributed chiefly to stormy weather which upset colony attendance. Eggs were also smaller in 1972. A seasonal decline in egg size (volume) was noted in all three years, attributed mainly to the later laying of young birds. Egg size increased with age, at least up to the fifteenth year. Eggs lost totalled 30% of those laid; 73% of this total was due to predation by Herring Gulls and of Jackdaws. Most losses (45%) occurred during the first 10 days after laying. Of lost eggs, 25% were replaced, usually 14 days after the loss of the original; only eggs laid and lost early in the season could be replaced. Only 7% of the chicks which hatched failed to fledge. Most (62.5%) chick losses occurred in the first week of nestling life, when chick weight was related to egg size. Afterwards, both growth rate and fledging weight were independent of egg size. The chicks fledging early in the season were heavier than later chicks. Failure to fledge was mainly due to a breakdown in behaviour between parent and young, rather than to predation. Breeding success was highest for birds breeding early in the season, most of which were older, more experienced breeders. These laid early enough to replace an egg if it was lost; they produced large eggs, and their chicks were therefore both heavier than average during the critical first 7–10 days of life, and fledged at a high weight. Thus experience accumulated with age, and the ability to lay early in the season are important for successful breeding in the Razorbill.     

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.     

Determination of clutch-size in the Kentish Plover Charadrius alexandrinus

We studied the clutch-size distribution of Kentish Plover Charadrius alexandrinus in 1991 and 1992 and tested three hypotheses for determination of clutch-size: egg-formation ability of females, incubation ability of parents and nest predation. Variation in clutch-size was small: 71 out of 74 clutches had three eggs (coeff. var. = 7%). Females spent more time foraging (51 ± 6%) pre-laying and during egg-laying than their mates (39 ± 5%). However, we concluded that egg-formation was not constrained by food availability because 77% and 100% of clutches were initiated before the peaks of prey density and of prey mass, respectively. Furthermore, the number of clutches initiated and completed over time was unrelated to prey density. By experimentally reducing and enlarging clutches, we found that enlarged clutches of four eggs took longer to hatch (24.8 ± 0.9 days) than control clutches of three (21.6 ± 0.7 days). Eggs of enlarged clutches also lost weight more slowly during incubation in both years compared with control clutches. No difference was found in the incubation behaviour or weight loss of parents between reduced, control and enlarged clutches. We found no evidence to support the nest predation hypothesis, since neither the proportion of nests predated nor the number of chicks hatched was different between reduced, control and enlarged clutches. The results of this study are most consistent with the incubation ability hypothesis, although the parental ability hypothesis has remained untested.     

THE FOOD OF SOME LARUS GULLS

The food of L. argentatus on Skomer and Skokholm Islands (Wales) was studied by the analysis of stomach contents, pellets and direct observation. L. argentatus is mainly a scavenger and exists for the most part on fish-waste and garbage discarded by man. At Ncwborough Warren argentatus feeds more on arable land and on the shore than do the Skokholm and Skomer birds.
Although argentatus frequently eats limpets Patella spp., there is little competition for food between them and Oystercatchers Haematopus ostralegus because the two species eat different sized limpets.
A summary of previous studies into the food of argentatus is given.
L. marinus is mainly a predator feeding on the young of other gulls, sea-birds and Rabbits but it also eats carrion, offal and fish-waste when available and during the winter. The numbers of Manx Shearwaters Procellaria puffinus killed on Skokholm and Skomer are discussed.
L. fuscus obtains most of its food on the shore and on arable land. There appears to be little competition, at least in the areas studied, between .fuscus and argentatus for food.     

Breeding biology of rooks (Corvus frugilegus L.) in Hawke's Bay,New Zealand

Abstract

Three rookeries in Hawke's Bay were studied during 1966–68. First or replacement clutches were started between 26 August and 23 October. First clutches averaged 4.3 eggs and replacements 3.7 eggs. The mean size of first clutches varied between years from 4.1 to 4.6 eggs. Incubation took 17–18 days. Most losses occurred around hatching, when about 40% of the eggs or young were lost. Incubated eggs and small nestlings incurred losses of 20% and 10% respectively, and all nestlings older than 10 days survived to at least 20 days. On average, 1.4 young were reared per nest in which eggs were laid; successful nests averaged 2.2 young. First clutches averaged 1.3 young (2.4 per successful first clutch). During the season, mean clutch size declined from 4.2 to 3.5, the mean number of young hatched declined from 2.0 to 0.6 per clutch, and the mean number of young fledged from all clutches declined from 1.3 to 0.4 per clutch. Mean nestling weight increased with age from 14 g on the first day after hatching to 360 g on the 19th day. The causes of egg and nestling mortality and the adaptiveness of clutch size are discussed.     

SURVIVAL OF MANX SHEARWATERS PUFFINUS PUFFINUS

The breeding success of Manx Shearwaters at Skokholm Island, Pembrokeshire, was followed in study burrows, and data on survival of the young were augmented by ringing large numbers as they were about to leave the island and recapturing them in later years.
The weight of the young at fledging and the date at which they leave affect their chances of survival; overall, as many as 30% of the young birds may survive to reach breeding age, which is thought to be normally about 5–6 years. Adult mortality varies between wide limits of about 5 and 20%, but juvenile survival appears to be of about the right order to balance adult losses.
The significance of the one-egg clutch is discussed. At the normal time of laying the female may not be able to obtain sufficient food to form a second egg. A two-egg clutch, laid at a later date, would not produce more surviving young since these would fledge too late in the season to have a good chance of survival.
The advantage of leaving the island at a heavy weight is discussed, and it is concluded that the fat stores laid down give heavy young a greater chance than light young of migrating a long distance without food. It is possible that the young may even reach their winter quarters (off Brazil) without having to feed on passage if they depart with enough stored fat.     

THE BREEDING BIOLOGY OF THE BLACK-FACED DIOCH QUELEA QUELEA IN NIGERIA

A study of the breeding biology of Quelea quelea in Nigeria, and particularly at a large breeding colony near Lake Chad, showed that losses of eggs and young were extremely small. 95% of eggs laid hatched successfully, and 87% give rise to fledglings. Nestling deaths were density-dependent and apparently due to starvation.
The incubation period was 10 days or less. By day, eggs were heated to 34°- 37° C. by the sun; at night the females incubated. The nestlings were initially fed mainly on insects, their diet gradually changing to one of seeds—mostly of the grass Echinochloa pyramidalis . The deep body temperatures of young birds were determined. It is suggested that the nestlings left the nest after, on average, 11¹/₂ days to escape intolerable temperature conditions in the nest.
Fat reserves were accumulated by nestlings and fledglings, and were utilized when the young became independent. The adults put on fat during the incubation period and lost it during the time spent feeding nestlings.
It is concluded that the most common clutch-size of Q. quelea , which is everywhere three, corresponds to the largest number of young the parents can normally nourish. This conforms to Lack's theory on the significance of clutch-size, and gives no support to Skutch's opinion that the theory does not apply to tropical birds.     

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 cost of incubation in relation to clutch-size in the Collared Flycatcher Ficedula albicollis

This study examines the importance of avian incubation costs as determinants of clutch-size variation by performing clutch-size and brood-size manipulations in the same population of Collared Flycatchers Ficedula albicollis during the same breeding season. In 2 5 cases when three or more clutches of the same size were completed on the same day, we moved two eggs on the day after the last egg had been laid from one randomly selected clutch (C) to another (C) and moved two other eggs from this to a third clutch (C⁺). In 20 other cases of simultaneously completed clutches of the same size, we moved two randomly selected young from one brood to a second and from that moved two other young to a third (B, B and B⁺groups). Most females were weighed the day after completion of the clutch and 1–4 days before hatching of the young, and some of them also 10–14 days after hatching of the young. We measured the daily energy expenditure of females incubating manipulated clutches of 4, 6 and 8 eggs by means of the doubly-labelled water (D₂¹⁸O) technique and also recorded their nest attendance. Hatching success of fertilized eggs was reduced in the enlarged clutches compared with control and reduced clutches. Females expired on average 3142.6 ml CO₂ and expended 78.6 kJ per day while incubating, which corresponds to a metabolic intensity of 3.3 times BMR. Daily energy expenditure increased with clutch-size due to higher costs while incubating, and not because of changed activity patterns. There were no significant differences in length of incubation, female mass or mass changes between phases for the C, C and C⁺groups. In both the C and B groups, enlarged broods produced significantly more fledged young than control broods, and those significantly more than reduced broods. Fledgling tarsus-length and mass did not differ significantly between treatments in either the C or B groups. There was no significant difference in breeding success between clutch and brood manipulations. In this season, incubation costs did not entail significant fitness losses, expressed either as fledgling production or female condition. Also, control females could have raised more young to fledging age than they did with no apparent costs.     

HERITABILITY AND SELECTION ON CLUTCH SIZE IN DARWIN'S MEDIUM GROUND FINCHES (GEOSPIZA FORTIS)

I studied the causes of variation and selection on clutch size in a population of Darwin's Medium Ground Finches (Geospiza fortis) on Isla Daphne Major, using data collected over a nine-year period (1976–1984). Quantitative-genetic analyses were carried out using the first clutch laid by a female in a given year. I used both unadjusted clutch-size values and values adjusted for between-year differences in mean clutch size for repeatability and regression analyses. Repeatability of clutch size was small (≤8%) and nonsignificant in all cases. Sib-sib analyses and parent-offspring regressions gave no evidence of a significant additive genetic component to clutch-size variation. Slopes of mother-daughter regressions were actually negative, suggesting possible maternal effects of mother's clutch size on daughter's clutch size. There was a small positive relationship between female age and clutch size but no effect of male or female body size or of large-scale differences in habitat quality on clutch size. Selection on clutch size was generally directional and positive: in almost all years in which successful breeding occurred, large clutches tended to fledge more chicks and produce more young surviving to the following year, possibly because there was no trade-off between clutch size and the weights of individual chicks at fledging. Thus, sustained directional selection for large clutch size may have reduced additive genetic variation in clutch size to low levels in this population. The size of a female's clutch may be primarily determined by unidentified proximate environmental factors which vary from year to year, rather than by any long-term optimization of clutch size with respect to adult survival.     

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.     

The breeding biology of the starling Sturnus vulgaris in northern Finland

The breeding biology of the starling Sturnus vulgaris was studied in three populations in northern Finland, with additional reference localities in nearby areas. Results from a total of 329 nests are presented from years 1962-1976.
The onset of laying in the Oulu area took place on 5 May, the median date being 10 May, about one week later than in southern Finland and 2–3 weeks later than in central Europe. The clutch-size was 5.29 ± 0.05 in the first nests and 5.06 ± 0.26 in later ones. Incubation time was 12.1 ± 0.6 d. An average of 3.31 ± 0.10 fledglings was produced per pair, 3.69 ± 0.08 in succesful clutches, which is significantly less than usually reported from central Europe. Most of the losses were due to starvation of the young.
No second broods were observed in the Oulu area, and repeated nesting was not common, probably because of the general late start of laying.
The reasons for the recent decline in several Finnish starling populations do not lie in the production capacity but must be sought for elsewhere.     

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.     

Intraclutch egg-size variation and offspring survival in the Kentish Plover Charadrius alexandrinus

We studied the consequences of intraclutch egg-size variation in Kentish Plovers Charadrius alexandrinus in southern Spain to test the hypothesis that females allocate resources preferentially to eggs with the greatest survival potential. Second eggs were larger and had a greater hatching success than the other eggs in a clutch. However, we did not find unequivocal support for the differential allocation hypothesis, because egg failures according to laying order were not due to the causes predicted by the hypothesis, and the change in egg-size with laying order was not consistent between successive clutches of renesting females. This suggests that nutrient availability during egg formation could act as a proximate factor affecting egg-size independently of laying order. Larger eggs produced heavier chicks and, within broods, heavier chicks were most likely to be recruited into the breeding population. By laying clutches with a larger mean egg volume in replacement clutches, females took longer to lay again after failure of the first clutch. Thus, opposing selective forces may act to increase and decrease egg volume. Although a trade-off between egg-size and clutch-size has not been found in precocial birds, our results suggest that there may be a trade-off between egg-size and fecundity, since an increase in the size of eggs within clutches may limit the time remaining within the season to lay second or replacement clutches.