Intraclutch Egg-size Variation Research Articles

Condition dependent strategies of egg size variation in the Common Eider Somateria mollissima.

We analysed intraclutch egg-size variation over the laying sequence in relation to clutch size, and the relation between clutch size and female body condition, in the Common Eider Somateria mollissima during an 8-year period. The aim was to assess if eiders adjusted egg size within the laying sequence depending on clutch sizes in response to body condition, as such an adjustment could have adaptive implications on reproductive success through a size advantage for the hatchlings. The analyses were performed on a population level; and then at the individual level using data from recaptured females that changed clutch size between years. Based on 1,099 clutches from 812 individual females, population clutch size averaged 4.13 eggs (range: 1-6), with 4- and 5-egg clutchesconstituting c.70% of all clutches, taking turns in being the most represented clutch size. Clutch size was positively related to female pre-laying body condition at both the population and individual levels. Egg size varied significantly within and between clutch sizes and changes were significantly related to the laying sequence. First eggs were significantly larger in 4-egg clutches and second eggs marginally smaller than in 5-egg clutches, a pattern also found among individual females changing clutch size between years. The relationship between female pre-laying body condition and clutch size, and the intraclutch egg-size pattern indicate that both clutch size and egg size are actively adapted to the pre-breeding body condition of the female. As egg size potentially optimise reproductive success through a size advantage in hatchlings, the observed pattern of intraclutch egg-size variation suggests that female eiders possesses a finely tuned conditional dependent mechanism that may optimize reproductive output in years were females are in suboptimal body condition for breeding.

Maternal provisioning by foam-nesting frogs of the genus Physalaemus (Anura, Leptodactylidae) in contrasting environments

In some environments prevailing conditions are unpredictable, thus anuran species use bet-hedging strategies and produce eggs of varied sizes. We investigated whether four species of Physalaemus (two from open, two from forest habitats) exhibit bet-hedging strategies, and if intra-clutch variation in egg size is greater for species that breed in the more unpredictable ponds of open habitats. All species studied adopted the bet-hedging strategy, with intra-clutch variation in egg size regardless of the environment; however, we found greater intra-clutch variation in the two species from open areas. The lower variation in egg-size found within forest habitats may be explained by the more stable environments that forest ponds provide for anuran eggs/embryos. Future studies in a phylogenetic context are needed to confirm the patterns detected here.

Intraclutch egg size variation is independent of ecological context among years in the European Starling Sturnus vulgaris

Female birds can potentially utilise various mechanisms, such as intraclutch egg size variation (ICESV), to modulate offspring quality. If this represents an ‘adaptive’ strategy allowing females to adjust reproductive effort after laying of the first egg, as information on resources becomes more predictable later in the laying sequence, then we would predict that ICESV would vary with ecological context, e.g. in ‘good’ versus ‘bad’ years. Here, in European Starlings (Sturnus vulgaris), we tested specific predictions that (1) females would lay relatively small last laid eggs in ‘poor’ years (with early laying and below-average breeding productivity); and (2) that ICESV would be lower in good years (late laying, above-average productivity) so that late-hatched chicks would not be disadvantaged. At the individual level we predicted (3) that repeatability of ICESV would be low if females adjusted this in relation to the environment in different years; and (4) that within-individual variation in ICESV should correlate with the number, and timing, of loss of chicks. Mean ICESV averaged −1.9% (measured as last egg mass relative to mean egg mass), typical of other hole-nesting passerines. However, our results did not support any of our hypotheses: (1) although there was annual variation in ICESV, in all years egg mass decreased through the laying sequence; (2) annual variation in ICESV was not consistent with variation in phenology or breeding productivity among years; (3) marked individual variation in ICESV (last laid egg varied from −20 to +10% relative to mean egg mass within clutches) was repeatable, i.e. individual females did not adjust ICESV in subsequent breeding attempts; and (4) variation in the magnitude of ICESV, alone and combined with hatching asynchrony, did not predict early brood loss. Our study suggests that ICESV is a relatively inflexible component of an individual female’s phenotype that might contribute to variation in individual quality.

Variación en el tamaño de los huevos en garzas (Aves: Ardeidae) que anidan en la ciénaga de Birama, Cuba

Intraclutch egg size variation in birds depends on many ecological factors and on the evolutive history of each species. In wading birds, a trend to smaller eggs with laying order has been described, but comparative reports are scarce. In this study, egg size variation patterns were described for nine Egrets and Heron species nesting in Birama' Swamp, Cuba. The patterns were described using external dimensions of 3142 eggs from 1875 nests of Butorides virescens, Bubulcus ibis, Ardea alba, Nycticorax nycticorax, Nyctanassa violacea and four Egretta species, taken in the field between 1998 and 2006. Results showed that eggs were 4.9-10% of adult weight and had volume variation coefficients between 6-9%. There were no general and consistent interspecies relationship between clutch size and egg sizes. Average volumes tend to get smaller with laying order, but it is not statistically detectable in Butorides and Bubulcus. Last egg was between 0.2% and 15% smaller than the first, showing an inverse relationship with it. Intraclutch asymmetry is light in E. thula and fluctuating around null in Bubulcus. Size only predicted laying or hatching order for the last egg, in nests with more than two eggs, with 72.4% of confidence.

Effects of intraclutch variation in egg size and hatching asynchrony on nestling development and survival in semi-precocial Herring Gulls

Intraclutch egg size variation may non-adaptively result from nutritional/energetic constraints acting on laying females or may reflect adaptive differential investment in offspring in relation to laying/hatching order. This variation may contribute to size hierarchies among siblings already established due to hatching asynchrony, and resultant competitive asymmetries often lead to starvation of the weakest nestling within a brood. The costs in terms of chick mortality can be high. However, the extent to which this mortality is egg size-mediated remains unclear, especially in relation to hatching asynchrony which may operate concomitantly. I assessed effects of egg size and hatching asynchrony on nestling development and survival of Herring Gulls (Larus argentatus), where the smaller size and later hatching of c-eggs may represent a brood-reduction strategy. To analyze variation in egg size, I recorded the laying order and laying date of 870 eggs in 290 three-egg clutches over a 3-yr period (2010-2012). I measured hatchlings and monitored growth and survival of 130 chicks from enclosed nests in 2011 and 2012. The negative effect of laying date ( =− 0.18 ± SE 0.06, P = 0.002) on c-egg size possibly reflected the fact that late breeders were either low quality or inexperienced females. The mass, size, and condition of hatchling Herring Gulls were positively related to egg size (all P < 0.0001). C-chicks suffered from increased mortality risk during the first 12 d, identified as the brood-reduction period in my study population. Although intraclutch variation in egg size was not directly related to patterns of chick mortality, I found that smaller relative egg size interactively increased differences in relative body condition of nestlings, primarily brought about by the degree of hatching asynchrony during this brood-reduction period. Thus, the value of relatively small c-eggs in Herring Gulls may lie in reinforcing brood reduction through effects on nestling body condition. A reproductive strategy Herring Gulls might have adopted to maintain a three-egg clutch, but that also enables them to adjust the number of chicks they rear relative to the prevailing environmental conditions and to their own condition during the nestling stage.

Intra-clutch pattern of albumen δ13C and δ15N in yellow-legged gulls Larus michahellis: female dietary shift or resource allocation strategy?

Energy or nutritional constraints associated to female dietary shifts during the clutch production period may play a role in generating intra-clutch egg size variation in yellow-legged gulls Larus michahellis. To explore this possibility, we determined albumen d 13 C and d 15 N values in three-egg clutches (modal clutch size) from three different breeding episodes: Ebro Delta 2004 and 2006, and Columbretes Islands 2004. Rather than a shift in females’ diet, consistent intraclutch patterns of variation in egg size and albumen isotopic values (particularly in the case of albumen d 13 C, which values held constant throughout the laying sequence) pointed to an intrinsic mechanism as the most feasible cause for the relatively smaller size of third/last-laid eggs. However, diet ‘‘quality’’ for breeding females seemed to affect intra-clutch egg size variation. In particular, a deficit of specific nutrients for egg formation associated to refuse scraps exploitation (as suggested by depleted albumen isotopic values) likely resulted in the more apparent intra-clutch egg size profile for the Ebro Delta 2004. In the absence of dietary shifts, the observation of consistently higher d 15 N values for third-albumens suggested a greater contribution of endogenous resources to their synthesis, as conversion of stored reserves into egg proteins results in greater isotopic fractionation, thereby yielding enriched isotopic signatures (particularly for d 15 N that shows greater isotopic fractionation with respect to that commonly assumed for d 13 C). We point to reabsorbed material derived from the hormonally-mediated regression of the female reproductive system (which is likely the intrinsic mechanisms resulting in the intra-clutch pattern of egg size variation: the hormonal hypothesis) as the most feasible endogenous source of nutrients for the synthesis of last-laid eggs, as optimize reproductive investment and maximize female fitness.

Between and within clutch variation of egg size in Greater Rheas

Abstract We describe egg characteristics, and analyze between and within clutch variation in egg size and mass in a natural population of Greater Rheas (Rhea americana). We assess the effect of this variation on nesting success and egg success. Yolk represented 29.5% of egg mass whereas albumen was 63.9%. Yolk mass increased with egg width but not with egg length, while mass of albumen increased mainly with egg length. The largest and smallest eggs were 10.3% larger and 25.3% smaller, respectively than mean intra-clutch values. The widest egg was 11.9% wider while the narrowest egg was 20.5% narrower than mean intra-clutch values. There was a significant decrease in egg size between clutches during the breeding season as a result of a decrease in egg length. There was no effect of laying order on intra-clutch variation in egg size, but we detected an increase in the variation of egg length within clutches with clutch size. We did not detect a relationship between egg size and nesting success, and between ...

Intraclutch Egg-Size Variation in Magellanic Penguins

Abstract We investigated patterns and consequences of intraclutch egg-size variation in Magellanic Penguins (Spheniscus magellanicus). First-laid eggs were significantly larger than second-laid eggs, although the mean difference represented only 2% of an average egg's volume. The degree of intraclutch egg-size variation was similar among years and females of different ages. Intraclutch egg-size variation did not affect intraclutch differences in chick hatching weights or fledging success. We found no selective advantage for laying eggs of different sizes. Because both eggs have an equal probability of being lost, chance favors equal provisioning of eggs. Egg volume explained 35% of the variation in hatching weight but did not determine fledging success. Laying order, year, and female age were better predictors of fledging success than egg size. Factors such as laying and hatching order, parental quality, oceanographic conditions, fights, and predation are more important in determining chick survival than are differences in egg size.

Patterns and fitness consequences of intraclutch variation in egg provisioning in tropical Australian frogs

Intraclutch variation in offspring size should evolve when offspring encounter unpredictable environmental conditions. This form of bet-hedging should maximise the lifetime reproductive success of individuals that engage it. We documented the numbers of eggs and means and variances of yolk volume in 15 frog species that occur in tropical savanna woodland. We experimentally determined the effects of initial yolk volume on larval growth patterns in four species. Intraclutch variation in yolk volume occurred to some degree in all species surveyed. Some species had very low, others had very high, intraclutch variation in yolk volume, but all species in which some clutches were highly variable also produced clutches with low variability. Species that occur in areas where the offspring environment is likely to be unpredictable had elevated levels of intraclutch variation in egg provisioning. There was no trade-off between egg size and number in any species surveyed. Under benign laboratory conditions, tadpoles from eggs with larger yolk volumes hatched at larger sizes, and these size differences persisted through a substantial proportion of the larval stage. This indicates that intraclutch variation in egg size has major offspring and thus parental fitness consequences, and is therefore a functional selection variable. This study provides evidence in support of models which predict that intraclutch variation in offspring provisioning can evolve in organisms that reproduce in unpredictable habitats.

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.

Effects of Environmental Conditions and Parental Quality on Inter- and Intraclutch Egg-Size Variation in the Collared Flycatcher (Ficedula Albicollis)

AbstractEgg size is a particularly important life-history trait mediating maternal influences on offspring phenotype. Females can vary their egg-size investment in relation to environmental circumstances, their own breeding condition, and the quality of their mate. Here we analyzed inter- and intraclutch variation in egg size in the Collared Flycatcher (Ficedula albicollis) on the basis of eight years of data. According to our results, mean egg size increased with female condition, but did not differ among young, middle-aged, and old females. The male’s age, body size, and forehead patch size did not influence egg size; thus, we found no evidence for differential investment in egg size in relation to male quality. We found no effect of laying date on egg size when controlling for ambient temperature during the egg formation period, yet temperature had a significant effect on egg size. That result indicates proximate constraints on egg formation. Furthermore, we report on annual differences in intraclutch egg-size variation. Egg size increased within clutches in years with a warm prelaying period; whereas in years when the weather during that period was cold, there was no significant intraclutch trend. Proximate considerations seem to explain the observed patterns of intraclutch egg-size variation; however, we cannot reject the adaptive explanation. Mean egg size and intraclutch egg-size variation were unrelated to clutch size. Therefore, we found no evidence for a trade-off between size and number of eggs within a clutch.

EFFECTS OF ENVIRONMENTAL CONDITIONS AND PARENTAL QUALITY ON INTER- AND INTRACLUTCH EGG-SIZE VARIATION IN THE COLLARED FLYCATCHER (FICEDULA ALBICOLLIS)

Abstract Egg size is a particularly important life-history trait mediating maternal influences on offspring phenotype. Females can vary their egg-size investment in relation to environmental circumstances, their own breeding condition, and the quality of their mate. Here we analyzed inter- and intraclutch variation in egg size in the Collared Flycatcher (Ficedula albicollis) on the basis of eight years of data. According to our results, mean egg size increased with female condition, but did not differ among young, middle-aged, and old females. The male’s age, body size, and forehead patch size did not influence egg size; thus, we found no evidence for differential investment in egg size in relation to male quality. We found no effect of laying date on egg size when controlling for ambient temperature during the egg formation period, yet temperature had a significant effect on egg size. That result indicates proximate constraints on egg formation. Furthermore, we report on annual differences in intraclutch...

INTRACLUTCH EGG-SIZE VARIATION IN MAGELLANIC PENGUINS

Abstract We investigated patterns and consequences of intraclutch egg-size variation in Magellanic Penguins (Spheniscus magellanicus). First-laid eggs were significantly larger than second-laid eggs, although the mean difference represented only 2% of an average egg's volume. The degree of intraclutch egg-size variation was similar among years and females of different ages. Intraclutch egg-size variation did not affect intraclutch differences in chick hatching weights or fledging success. We found no selective advantage for laying eggs of different sizes. Because both eggs have an equal probability of being lost, chance favors equal provisioning of eggs. Egg volume explained 35% of the variation in hatching weight but did not determine fledging success. Laying order, year, and female age were better predictors of fledging success than egg size. Factors such as laying and hatching order, parental quality, oceanographic conditions, fights, and predation are more important in determining chick survival than ...

Adaptive aspects of intraclutch egg-size variation in the High Arctic barnacle goose (Branta leucopsis)

The pattern of intraclutch egg-size variation in barnacle goose (Branta leucopsis) clutches and its adaptive implications was studied in Svalbard, Norway, from 1989 to 1998. Egg size was measured in relation to laying sequence, egg predation and hatching order were recorded to determine hatching success of eggs in different laying sequences, and the time when incubation started was examined. Egg size showed a rather consistent pattern, with a large second-laid egg and declining egg size for the remainder of the clutch. The first-laid egg was usually smaller than the second one, except in clutches with two and three eggs. Predation was highest for the first-laid egg, and last-laid eggs hatched last in most cases, although only one last-laid egg was abandoned. Four of six females started incubation before clutch completion. Both the "nutrient-allocation hypothesis" as well as the "early incubation start hypothesis" may contribute to explaining the expressed pattern of intraclutch egg-size variation. The fitness gains due to allocating fewer nutrients to eggs in unfavourable positions in the laying sequence may explain the small size of the first egg, whereas the multiple benefits of an early incubation start may have led to the decline in egg size later in the laying sequence as a mechanism to counteract hatching asynchrony.

Avian egg size: variation within species and inflexibility within individuals

Egg size is a widely-studied trait and yet the causes and consequences of variation in this trait remain poorly understood. Egg size varies greatly within many avian species, with the largest egg in a population generally being at least 50% bigger, and sometimes twice as large, as the smallest. Generally, approximately 70% of the variation in egg mass is due to variation between rather than within clutches, although there are some cases of extreme intra-clutch egg-size variation. Despite the large amount of variation in egg size between females, this trait is highly consistent within individuals between breeding attempts; the repeatability of egg size is generally above 0.6 and tends to be higher than that of clutch size or laying date. Heritability estimates also tend to be much higher for egg size (> 0.5) than for clutch size or laying date (< 0.5). As expected, given the high repeatability and heritability of egg size, supplemental food had no statistically significant effect on this trait in 18 out of 28 (64%) studies. Where dietary supplements do increase egg size, the effect is never more than 13% of the control values and is generally much less. Similarly, ambient temperature during egg formation generally explains less than 15% of the variation in egg size. In short, egg size appears to be a characteristic of individual females, and yet the traits of a female that determine egg size are not clear. Although egg size often increases with female age (17 out of 37 studies), the change in egg size is generally less than 10%. Female mass and size rarely explain more than 20% of the variation in egg size within species. A female's egg size is not consistently related to other aspects of reproductive performance such as clutch size, laying date, or the pair's ability to rear young. Physiological characteristics of the female (e.g. endogenous protein stores, oviduct mass, rate of protein uptake by ovarian follicles) show more promise as potential determinants of egg size. With regards to the consequences of egg-size variation for offspring fitness, egg size is often correlated with offspring mass and size within the first week after hatching, but the evidence for more long-lasting effects on chick growth and survival is equivocal. In other oviparous vertebrates, the magnitude of egg-size variation within populations is often as great or greater than that observed within avian populations. Although there are much fewer estimates of the repeatability of egg size in other taxa, the available evidence suggests that egg size may be more flexible within individuals. Furthermore, in non-avian species (particularly fish and turtles), it is more common for female mass or size to explain a substantial proportion of the variation in egg size. Further research into the physiological basis of egg-size variation is needed to shed light on both the proximate and ultimate causes of intraspecific variation in this trait in birds.

Plasma Amino Acid Concentrations as an Indicator of Protein Availability to Breeding Herring Gulls (Larus Argentatus)

AbstractIn a captive-feeding study using Herring Gulls (Larus argentatus), plasma amino-acid concentrations increased in response to an increase in dietary protein. Plasma amino-acid concentrations were also measured in wild Herring Gulls captured during incubation at eight Laurentian Great Lakes colonies. Those concentrations were used as an indicator of protein availability at those locations. Significant differences in amino acid concentrations were observed among colonies. Lower amino acid levels, particularly of the essential amino acids, were measured in gulls nesting on Lake Superior, whereas values in gulls captured on Lake Ontario and Lake Erie were greater. Those geographic differences in protein availability likely reflected spatial differences in availability of high quality prey (e.g. fish). Geographic differences in prey availability probably affected diet composition. Comparison of amino-acid levels in wild birds to reference values obtained through the captive feeding study indicated that gulls nesting on Lake Superior may have been protein limited. Colony-wide estimates of adult female body condition, intraclutch variation in egg size, and productivity were correlated with an index of plasma amino-acid concentrations.

Intraclutch egg‐size variation in acanthosomatid bugs: adaptive allocation of maternal investment?

If there are differences in predation risk among the offspring within a clutch, parents may allocate less resources to the offspring facing higher risk. I examined parental investment in terms of egg size within clutches in five species of stink bugs (Heteroptera, Acanthosomatidae). In subsocial Elasmucha and Sastragala species, the female guards her eggs and first‐instar nymphs against invertebrate predators by covering her clutch with her body. Large differences in survival from predation between offspring at the centre and offspring at the periphery of the clutch have been reported in such subsocial insects. I found that Elasmucha and Sastragala females laid significantly smaller eggs in the peripheral (and thus more vulnerable) part of the clutch. Phenotypic trade‐offs between egg size and clutch size were detected in these subsocial species. Egg size was positively correlated with hatched first‐instar nymph size: smaller nymphs hatched from smaller peripheral eggs. In asocial Elasmostethus humeralis, however, no significant difference in size was detected between the eggs at the centre of and those at the periphery of the clutch. Thus, in subsocial acanthosomatid bugs, females seem to allocate their resources according to the different predation risks faced by the offspring within the clutch.

Experimental (tamoxifen-induced) manipulation of female reproduction in zebra finches (Taeniopygia guttata).

Experimental manipulation of reproductive phenotype is a potentially powerful approach for understanding the fitness relationships of traits such as egg size, egg composition, and egg number. In this study, I investigated the effect of the antiestrogen tamoxifen on multiple, estrogen-dependent reproductive traits in female zebra finches (Taeniopygia guttata). Short-term tamoxifen treatment of egg-laying females (two daily injections before laying) had no effect on the timing or the pattern of egg laying compared to sham controls. However, tamoxifen treatment caused (1) a marked, but transient, decrease in egg size; (2) increased within-clutch egg-size variation; (3) a reduction in plasma vitellogenin (VTG) levels; and (4) lower dry yolk and yolk protein content of tamoxifen-treated females. Plasma levels of the second yolk precursor, very low density lipoprotein (VLDL), were not affected by tamoxifen, and tamoxifen appeared to have no effect on oviduct function in egg-laying females. These results are consistent with tamoxifen blocking estrogen receptors in the liver, suppressing VTG production, and decreasing the plasma pool of yolk precursors below a level required to maintain yolk formation at the normal rate. Tamoxifen treatment can therefore be used successfully to manipulate several components of the female reproductive phenotype (egg composition, intraclutch egg-size variation) to further explore the fitness consequences of these traits.

Maternal effects of egg size in brown trout (Salmo trutta): norms of reaction to environmental quality

The magnitude of fitness variation caused by maternal effects and, thus, the adaptive significance of maternal traits may depend on environmental quality, generating crossing reaction norms among offspring phenotypes that shape life-history evolution. By manipulating intraclutch variation in egg size and comparing siblings we examined the maternal effects of egg size on offspring performance and tested for the existence of reaction norms to environmental quality using the brown trout Salmo trutta. When sibling groups of small and large eggs were reared separately in a hatchery environment initial size differences disappeared rapidly. However, in semi-natural environments and under direct competition, juveniles from large eggs experienced growth and survival advantages over siblings from small eggs. Moreover, distinct reaction norms existed, with the differences in performance of juveniles from small and large eggs being most pronounced in the poorer growth environments. Our results provide the first direct evidence, to our knowledge, for a causal relationship between egg size and fitness-related traits in fishes, independent of potentially confounding genetic effects. Moreover, they indicate that previous studies have been biased by experimental conditions that excluded competitive asymmetries and environmental variability. The existence of reaction norms indicates a shift in optimal egg size across gradients of environmental quality that probably shapes the evolution of this trait.

Intraclutch Egg-Size Variation in the Eurasian Kestrel: Advantages and Disadvantages of Hatching from Large Eggs

Egg size in birds may be positively correlated with hatching success (Jarvinen and Vaisanen 1983, Magrath 1992a), hatchling mass (Magrath 1992a, Nilsson and Svensson 1993), and nestling growth and survival (Schifferli 1973, Moss et al. 1981, Williams 1990). Hence, egg size could influence offspring fitness (Williams 1994). Egg size varies among and within clutches, the former accounting for most of the variation (Ojanen et al. 1979, Jarvinen and Viisanen 1983, Wiggins 1990). However, variation in egg size within clutches may have greater fitness consequences than that among clutches because young that hatch from smaller eggs compete with larger siblings and have a higher risk of starvation (Nilsson and Svensson 1993). Generally, egg size varies with laying order following patterns that are typical for a species or population; e.g. increasing (Howe 1976, Mead and Morton 1985), decreasing (Parsons 1972, Nisbet and Cohen 1975), or increasing to the middle egg and decreasing thereafter (Arnold 1991, Williams et al. 1993). These patterns may change relative to food supply (Simmons 1994), year (Jover et al. 1993), or clutch size (Vifnuela 1997) and may result from physiological or nutritional constraints on the female and/or be adaptive strategies to improve breeding success. Several constraints could cause intraclutch variation in egg size. For example, egg size may increase with laying sequence because females are gearing up physiologically for starting egg production (Parsons 1976). Last-laid eggs may be smaller because prolactin blood levels, which increase with the onset of incubation, could reduce the size of developing follicles (Leblanc 1987). Egg size also could decline with laying sequence because energy reserves for egg formation are depleted throughout laying (Ryden 1978, Pierotti and Bellrose 1986). Finally, fluctuating environmental conditions at the time of egg laying associated with low temperatures (Ojanen et al. 1981, Jarvinen and Ylimaunu 1986, Magrath 1992b) or with changes in food availability (Jarvinen and Vaisanen 1983) may cause variation in egg size independent of laying order. At least three hypotheses have been proposed to explain the function of intraclutch variation in egg