Causes and consequences of egg mass variation between and within blue tit clutches

Causes of egg-size variation between and within clutches were studied in clutches of the blue tit ( Parus caeruleus L.). We measured the mass of each egg in the laying sequence in unmanipulated clutches, in clutches of parents experimentally supplied with extra food before egg-laying, and in clutches of parents supplemented with extra food after the start of egg-laying. Hatchlings were weighed at an age of two days and their mass was found to be positively related to egg mass. No general trend of decreasing or increasing egg mass was found within the laying sequence. Females provided with extra food before egg-laying laid clutches with significantly less variation in egg mass than did control females. The reason for this was that the first-laid egg of unmanipulated females was lighter than the rest of the eggs in the clutch. This pattern disappeared in clutches of females receiving extra food. Thus, the reduction in egg mass variation among clutches of foodsupplemented females depended on an ability of these females, in contrast to control females, to lay a first egg of the same mass as the rest of the clutch. Eggs laid after the initiation of incubation were significantly heavier than equivalent eggs in those clutches where incubation started after clutch completion. The difference was small, however, and the adaptive significance of the finding is questionable. We argue that intra-clutch variation in egg mass is connected with greater fitness consequences than in inter-clutch variation. Furthermore, our results indicate that energetical constraints on the laying female are more important as a cause of the observed intra-clutch variation in egg mass than are adaptive responses to the environment.     

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.     

Within‐ and between‐season repeatability of eggshell colouration in the great reed warbler Acrocephalus arundinaceus

While avian eggshell colouration has attracted biologists for decades, little is known about its variation within individuals. The main goal of this study was to explore within‐ and between‐season repeatability of eggshell appearance in the great reed warbler Acrocephalus arundinaceus. To achieve this, we measured eggshell reflectance of first and replacement clutches of individual females within one breeding season, and the reflectance of their first clutches in two following breeding seasons. As environmental conditions may affect egg colouration, repeatability was estimated from linear mixed‐effects models where we initially included temperature and rainfall during egg laying, as well as year, clutch order and egg‐laying date as fixed effects, and clutch identity nested in females as random effects. Eggshell appearance within clutches showed moderate repeatability, while both within‐ and between‐season repeatability in clutch colouration for individual females was low. Our findings indicate that low intra‐clutch variation may have a function within the context of brood parasitism, i.e. in facilitating host recognition of alien eggs. With variable eggs between successive clutches, however, host females may need to relearn the appearance of their eggs with every clutch they lay. This could represent a significant constraint for host egg‐discrimination abilities. Yet, whether environmental or intrinsic physiological factors are responsible for the variation in eggshell colouration between successive clutches of the same females still remain to be discovered.     

Intraclutch egg-size variation in the Barnacle Goose Branta leucopsis: an egg-removal experiment

We carried out an egg-removal experiment on Barnacle Geese Branta leucopsis to test the hypothesis that tactile contact between the brood patch and eggs accumulating in the nest during laying was causally related to intraclutch egg-size variation, acting through the development of incubation behaviour. Egg removal had no effect on mean egg mass, relaying interval or clutch size: mean (±s.e.) clutch sizes for first clutches were 4.9 ± 0.9 and 4.7 ± 0.9 eggs for control and experimental females, respectively. We therefore conclude that Barnacle Geese have a determinate pattern of egg laying. Daily removal of all eggs from the first-laid egg did not affect the within-clutch pattern of egg-size variation: in both control and experimental clutches the first egg was relatively small, the second or third egg was largest and there was a linear decline in egg size to the smallest, last-laid egg. There was no significant difference in either the absolute or relative size of the last-laid eggs in control and experimental clutches. Tactile contact with eggs in the nest, therefore, is not required for the expression of intraclutch egg-size variation in Barnacle Geese. In this respect, determinate-laying Barnacle Geese differ from species with indeterminate laying patterns (e.g. gulls) where egg removal does affect the normal pattern of within-clutch egg-size variation. If hormonal changes (e.g. prolactin) associated with onset of incubation are causally related to the intraclutch decrease in egg size, then the stimuli involved appear to differ between determinate and indeterminate layers.     

Causes and consequences of egg-size variation in the European Starling Sturnus vulgaris

Variations in egg-size (weight, or an index of volume derived from the length and breadth of eggs) were analysed for four years' data from a colony of Starlings in southern England. Variation was much greater among clutches than within clutches. There were no consistent trends for egg-size to increase or decrease through the laying sequence, but the first- and last-laid eggs tended to be smaller than the clutch mean. Loss of single nestlings (perhaps by starvation) was more frequent if the smallest, or last-laid egg was particularly small. Effects of egg-size on the growth of surviving chicks were slight-most variation in development was attributable to factors such as clutch or brood size, and nestlings' relative weights fluctuated so that they continually changed ranks during the nestling period. In Starlings, egg-size was inversely correlated with clutch size, but three open-nesting species (Bullfinch, Song Thrush and Blackbird) showed the opposite pattern. These results are discussed in relation to their implications for understanding the adaptive aspects of egg-size variation.     

Maternal androgens in the pied flycatcher: timing of breeding and within-female consistency

Maternal hormones can have substantial phenotypic effects in the progeny of many vertebrates. It has been proposed that mothers adaptively adjust hormone levels experienced by particular young to optimize their reproductive output. In birds, systematic variation in egg hormone levels has been related to different female reproductive strategies. Because in many bird species prospects of the offspring change seasonally and with brood number, strategic adjustment of yolk androgen levels would be expected. To test this idea, we induced pied flycatcher (Ficedula hypoleuca) females to nest twice during the same season by removing their first clutches shortly after clutch completion. We collected eggs of first and replacement clutches to measure yolk concentrations of androstenedione (A4) and testosterone (T) and captured the females that laid these clutches for phenotypic measurements. Although average egg androgen levels were remarkably consistent within females, hormone patterns differed considerably between first and replacement clutches. Eggs of replacement clutches were heavier with larger yolks compared to first clutches, but they contained on average lower levels of androgens. Within clutches, androgen concentration increased over the laying sequence in the first clutch, but decreased or remained more constant over the laying sequence in the replacement clutch. Mean yolk T, but not A4 levels, were negatively associated with laying date for both breeding attempts. Moreover, females in good body condition produced eggs containing lower levels of androgens than females in poor condition. Our results are consistent with the idea that differences in yolk androgen levels may be one mechanism underlying seasonal variation in reproductive success and it is possible that changes in egg androgen patterns may reflect a change in female reproductive strategy. High within-female consistency also highlights the possibility that there may be some underlying genetic variation in yolk androgen levels.     

Variation and trade-off in reproductive output of the Andean frog<Emphasis Type="Italic"> Hyla labialis</Emphasis>

I analysed variation in number and size of eggs in association with variation in body size of Hyla labialis females from three altitudinally widely separated populations in the Colombian Andes. Females at 3,500 m were significantly larger and heavier than those at 2,700 m, which in turn were larger and heavier than those at 2,000 m. After accounting for interpopulational differences in body size, clutch size (i.e. number of eggs) and egg size differed significantly between populations, but clutch mass did not. At 3,500 m, large relative clutch mass tended to be associated with larger eggs, whereas at lower altitudes it was associated with relatively larger clutch size. Within each population, females showed a significant trade-off between clutch size and egg size, even though relative clutch mass varied substantially and a clutch rarely used up the carrying capacity of their body cavity. At 3,500 m, comparisons between the early and late breeding season within a year suggested that climate fluctuations may play a role in reproductive-output variation within a population. Laboratory-kept females in a high somatic condition after spawning laid larger clutches in the next breeding event, but the trade-off persisted despite ad libitum food supply. Comparisons between field and laboratory suggest that limits to reproductive output were due to physiological, but not morphological restrictions. Therefore, the trade-off is better explained by energy limitation for egg growth depending on clutch size, than by the body cavity limitation hypothesis.     

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.     

Egg-size variation in the bluethroat (Luscinia s. svecica): constraints and adaptation

We examined inter- and intra-clutch egg-size variation in the bluethroat (Luscinia s. svecica), an open-nesting passerine breeding in the sub-alpine region in southern Norway. By removing first clutches shortly after egg-laying, we induced laying of a repeat clutch. Females significantly reduced the number of eggs from the first to the second nesting attempt, but increased mean egg size. Females in good condition laid significantly larger eggs than those in poor condition. Consistent with predictions of the brood survival hypothesis, assuming an adaptive investment in last eggs to ensure survival of all eggs in the clutch, we found that the size of the last eggs in first clutches was generally larger than the mean egg size of the clutch, and that the relative size of the last egg increased with clutch size. However, a large last egg reflected a general increase in egg size throughout the laying sequence rather than a specific investment in the last egg only. Egg size was not significantly influenced by sex or paternity (within-pair versus extra-pair) of the embryo. In repeat clutches the last egg was not consistently larger than the mean for the clutch. We conclude that female bluethroats face resource limitations during egg formation early in the season, and that the patterns of increase in egg size with laying order for first clutches, and from first to repeat clutches, can largely be explained by proximate constraints on egg formation.     

Egg size and clutch size in the reproductive investment of American Kestrels

We studied causes and consequences of egg-size variation among clutches of American kestrels ( Falco sparverius ). Egg. from 275 clutches were measured 1990 to 1992. To test the hypothesis that the size of eggs was contrained by food availability in the pre-laying period, we censused small mammal populations in the three years and performed a food supplementation experiment in 1990 and 1991. Kestrels did not advance the date they laid their first egg but did lay significantly larger eggs in response to extra food. The size of eggs was correlated with small mammal abundance on the territoty, and females in good body condition tended to lay large eggs. Body size did not affect egg size, and there were no relationship between agg size and laying date except in 1900, the poorest food year. Clutches with a large mean egg volume had better hatching success than clutches containing small eggs. We argue that there is a phonetypic component to egg size in kestrels, and that kestrels use egg size to fine-tune reproductive investiment to available resources.     

Effects of experimental calcium availability,egg parameters and laying order on Great Tit Parus major eggshell pigmentation patterns

Many bird species lay eggs speckled with protoporphyrin‐based spots, however, for most of them the function of eggshell spotting is unknown. A plausible hypothesis is that protoporphyrin might have a structural function in strengthening the eggshell and is therefore deposited when calcium is scarce. In this study, we experimentally provided Great Tit Parus major females with supplemental calcium to examine its effect on the protoporphyrin‐based maculation of their eggs. In addition, we studied variation in eggshell pigmentation patterns in relation to other egg parameters and laying order. Calcium‐supplemented females laid larger eggs but shell thickness was not significantly affected by the treatment. Calcium supplementation may reduce the time and energy females devote to searching for calcium‐rich material, so that they can collect more nutrients and so lay larger eggs. Furthermore, pigment darkness was associated with egg volume and shape, which suggests that female quality and environmental food availability may also influence the shell pigmentation pattern. Within clutches, later‐laid eggs had larger and darker spots that were distributed more unevenly on the shell surface. This within‐clutch pattern could be explained by the increase in egg volume and egg shape and a decline in shell thickness with egg‐laying order, which characteristics were all related to shell‐spotting pattern. Eggs with a coronal ring had thinner shells, but pigment intensity and spot size were not related to shell thickness. Thus, our results suggest that concentrated spotting distribution may have a mechanical function, supporting the structural‐function hypothesis.     

Maternal resource variation across the laying sequence in Canada geese Branta canadensis maxima

Variation in maternal investments to offspring presumably reflects an optimization of resource allocation such that a female's fitness is maximized. In birds, both egg size and yolk constituents are examples of resources that can vary among offspring within a clutch. Egg size and maternally-derived steroid hormone concentrations present in yolk have been characterized for many species that lay small clutches or have altricial young, but little information is available for species that lay moderate to large clutches of precocial young. In this study, we recorded laying position, measured fresh egg mass and determined maternally-derived testosterone and estradiol concentrations present in yolks for whole clutches of free-living Canada geese Branta canadensis maxima to assess variation in maternal resources within clutches. We found that egg size varied non-linearly across the laying sequence such that first laid eggs were small, the largest eggs in the clutch occurred in the second and third positions, and size declined in eggs laid in subsequent positions. Concentration of testosterone in the yolk followed a pattern in which the first and second laid eggs have the highest concentrations within a clutch and declining concentrations in subsequently laid eggs. In contrast, maternally-derived yolk estradiol concentrations (measured in a subset of clutches) did not change across the laying sequence.     

The effects of low dietary calcium during egg-laying on eggshell formation and skeletal calcium reserves in the Zebra Finch Taeniopygia guttata

Many small passerines forage intensively for calcium-rich foods during laying. Increased incidences of shell defects in eggs of small passerines have been reported, particularly in western Europe, and these have been explained in terms of declining calcium availability in soils, resulting from prolonged anthropogenic acid deposition. Studies in the field have provided laying birds, nesting in areas of low calcium availability, with calcium supplements. An alternative approach was adopted in this study by allowing captive Zebra Finches Taeniopygia guttata to lay first clutches on ad libitum calcium, switching them to a low calcium diet for 72 hours for the formation of all but the first egg of the second clutch and reinstating ad libitum calcium for the final clutch. Control females had access to ad libitum calcium for all three clutches. Clutch sizes did not vary significantly between birds on low calcium and controls. The former took over three days longer to lay clutch 3 than did controls but the difference was not statistically significant. Birds on low calcium laid eggs that declined in shell ash mass with laying sequence, indicating that birds may have been calcium-limited. Although not statistically significant, eggshell thickness also declined with laying sequence in clutches laid by females on low calcium. The remaining egg measurements (shell mass, shell surface area and volume] of clutches laid by birds on low calcium did not differ significantly from those of controls. Furthermore, females on low calcium did not resort to skeletal reserves to provide sufficient calcium for egg formation. Dietary calcium appears to be of paramount importance in providing sufficient calcium for clutch formation.     

Indeterminate laying and flexible clutch size in a capital breeder, the common eider

Clutch size control in capital breeders such as large waterfowl has been much debated. Some studies have concluded that clutch size in ducks is determined before the start of laying and does not change in response to egg additions or removals. The response, however, may depend on the timing of tests, and experiments may have been too late for females to alter the number of eggs. We here study clutch size responses to predation of first and second eggs in the common eider, using protein fingerprinting of egg albumen to verify that the same female continues laying in the nest after predation. Sixty of 79 females with early egg predation (one or both of the two first eggs) deserted the nest. Among the 19 females that stayed and continued laying, the mean number of eggs produced was 4.4, significantly higher than the 3.7 in non-predated nests. The staying females had similar egg size and clutch initiation date as females that deserted, and their body mass and clutch initiation date was similar to that of females whose clutches were not predated. Even capital-breeding common eiders may therefore be indeterminate layers, as many females in which early eggs are removed lay more eggs than others. A previous study has shown that they can reduce their laying if eggs are added. Our results add to increasing evidence that ducks have more flexible egg production than previously thought.     

Intra-Clutch Ratio of Yolk Progesterone Level Changes with Laying Date in Rockhopper Penguins: A Strategy to Influence Brood Reduction?

Hatching asynchrony in avian species generally leads to a size hierarchy among siblings, favouring the first-hatched chicks. Maternally deposited hormones affect the embryo and chick's physiology and behaviour. It has been observed that progesterone, a hormone present at higher levels than other steroid hormones in egg yolks, is negatively related to body mass in embryos, chicks and adults. A differential within-clutch progesterone deposition could therefore be linked to the size hierarchy between siblings and to the resulting brood reduction. We tested whether yolk progesterone levels differed between eggs according to future parental ability to feed the entire clutch in wild rockhopper penguins Eudyptes chrysocome. This species presents a unique reversed egg-size dimorphism and hatching asynchrony, with the larger second-laid egg (B-egg) hatching before the smaller first-laid egg (A-egg). Yolk progesterone levels increased only slightly with female body mass at laying. However, intra-clutch ratios were not related to female body mass. On the other hand, yolk progesterone levels increased significantly with the date of laying onset for A-eggs while they decreased for B-eggs. Early clutches therefore had proportionally more progesterone in the B-egg compared to the A-egg while late clutches had proportionally less progesterone in the B-egg. We propose that females could strategically regulate yolk progesterone deposition within clutches according to the expected food availability during chick growth, an adaptive strategy to adjust brood reduction to conditions. We also discuss these results, relating to yolk progesterone, in the broader context of other yolk steroids.     

Producing extra eggs does not deplete macronutrient reserves in European Starlings Sturnus vulgaris

Recent studies have suggested that the size of avian clutches may be determined in part by the costs of egg production, which are thought to involve the depletion of endogenous nutrient reserves. I tested the hypothesis that producing extra eggs would deplete protein and lipid reserves in a passerine species, the European Starling Sturnus vulgaris . The first three eggs of clutches were removed on the days they were laid to induce females to lay extra eggs. Females responded to this manipulation by laying, on average, 1.3 eggs more than controls. However, based on the distribution of clutch sizes, it appears that some females responded to the removal of three eggs by producing more than one extra egg, whereas others responded by laying no extra eggs or by deserting. In unmanipulated females, the lean dry mass of the flight muscles and total body lipid were found to decline throughout breeding, an observation which is often interpreted as evidence that egg production depletes endogenous reserves. However, females experimentally induced to produce an extra egg did not have reduced stores of protein or lipid at clutch completion. No effect was observed in starlings even though this study had substantial (80%) power to detect a change in protein reserves of the same magnitude as that observed in a previous study of gulls. The decrease in flight muscle mass that has frequently been observed during breeding in passerines may not be due simply to mobilization of endogenous protein reserves.     

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.     

Maternal androgens in black-headed gull (Larus ridibundus) eggs: consequences for chick development

We tested the hypothesis that mother birds counterbalance the negative effects of hatching asynchrony for later-hatched chicks by increasing the yolk androgen concentrations in consecutive eggs of their clutch. In doing so, they may adaptively tune each offspring's competitive ability and, thus, growth and survival. However, evidence in support of this hypothesis is contradictory. The yolk concentrations of maternal androgens in the eggs of black-headed gulls increase significantly with the laying order of the eggs in a clutch. We experimentally tested the functional consequences of this increase on chick development under natural conditions by injecting eggs with either an oil or androgen solution. We created experimental clutches in which androgen levels either stayed constant or increased with laying order while controlling for differences in egg quality by using only first-laid eggs. We then compared development, growth and survival between these broods. Androgen treatment enhanced embryonic development because androgen-treated eggs hatched half a day earlier than controls, while their size at hatching was similar to oil-treated controls. Androgen treatment did not increase chick survival, but it enhanced growth. Androgen-treated, third-hatched chicks had a higher body mass and longer legs than third-hatched chicks that hatched from oil-treated eggs. At the same time, growth of first chicks (which were all oil treated) was reduced by the presence of two androgen-treated siblings, suggesting that yolk androgens enhance the competitive ability of later-hatched chicks. Our results support the hypothesis that transfer of different amounts of androgens to the eggs of a clutch is a mechanism by which mothers maximize their reproductive output.     

Clutch size limitation in waders: experimental test in redshank Tringa totanus

Several hypotheses have been raised to explain the upper limit of clutch size at four eggs in waders (suborder Charadrii), which may play an important role in the evolution of the variety of mating and parental care systems in this group. Experimental tests of the hypotheses have produced conflicting results. It was recently suggested that the combined effects of several incubation costs of a larger clutch suffice to limit its size to four eggs in this group. Here we test the incubation-limitation hypothesis in a field experiment, in redshank Tringa totanus. We created five-egg clutches by adding one egg from another nest to a just completed four-egg clutch. Four-egg control clutches were created by replacing one of the eggs by an egg from another nest. All egg removals, additions and replacements were done before incubation started. Incubation time in five-egg clutches increased by 1 day to 24.3ǂ.23 days, compared to 23.3ǂ.32 days in four-egg clutches. Egg hatchability and nest predation rates did not differ significantly between treatments. On average five-egg clutches produced one extra chick at hatching (4.5ǂ.26 chicks) compared to four-egg clutches (3.5ǂ.27 chicks). Also when several additional costs from incubating enlarged clutches are added, redshanks by laying a fifth egg would on average increase their reproductive success at hatching by an estimated 22%. The incubation-limitation hypothesis therefore is clearly rejected in this species. Possible mechanisms behind the four-egg clutch limit in waders and ways of testing the alternatives are discussed.     

Effects of egg size and composition on the size, quality and survival of lapwing Vanellus vanellus chicks

The influence of egg size and composition on the size, quality and survival of lapwing chicks was examined on two farmland study sites in the Midland Valley of Scotland. Eggs comprised 33.1% yolk, 61.3% albumen and 5.6% shell. Whereas the yolk and shell proportions decreased with increasing egg size, the albumen proportion increased. Most variation in egg size was attributable to differences between females but was also influenced by clutch number (eggs in replacement clutches on the rough grazing, but not the arable, site were smaller), clutch size (eggs were smaller in smaller clutches), maternal body condition (females in good condition produced larger eggs) and habitat (since females on the arable site fed more successfully, they were in better condition and laid larger eggs). Chick size, weight and survival were all influenced by egg size. The incubation period varied between 21 and 28 days (mean = 25.2) and was shorter in clutches laid later in the season.