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.     

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.     

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.     

Fitness cost of incubation in great tits (Parus major) is related to clutch size

Life-history theory predicts that parents produce the number of offspring that maximizes their fitness. In birds, natural selection on parental decisions regarding clutch size may act during egg laying, incubation or nestling phase. To study the fitness consequences of clutch size during the incubation phase, we manipulated the clutch sizes during this phase only in three breeding seasons and measured the fitness consequences on the short and the long term. Clutch enlargement did not affect the offspring fitness of the manipulated first clutches, but fledging probability of the subsequent clutch in the same season was reduced. Parents incubating enlarged first clutches provided adequate care for the offspring of their first clutches during the nestling phase, but paid the price when caring for the offspring of their second clutch. Parents that incubated enlarged first clutches had lower local survival in the 2 years when the population had a relatively high production of second clutches, but not in the third year when there was a very low production of second clutches. During these 2 years, the costs of incubation were strong enough to change positive selection, as established by brood size manipulations in this study population, into stabilizing selection through the negative effect of incubation on parental fitness.     

Nest-building activity and laying date influence female reproductive investment in magpies: an experimental study

Nest size or nest-building activity has recently been hypothesized to be a postmating sexually selected signal in monogamous birds: females may assess a male's parental quality and willingness to invest in reproduction by his participation in nest building. Females may thus adjust their reproductive effort (i.e. clutch size) not only to their own abilities but also to those of their mates. We investigated whether female magpies, Pica pica, use nest-building activity rather than nest size to adjust their reproductive effort during replacement breeding attempts. After we removed their first clutch, high-quality pairs that built a large nest for the first clutch were more capable of building a replacement nest and females adjusted their clutch size in relation to the time it took to build the nest rather than nest size. We also found support for the hypothesized trade-off between clutch size and egg size in magpies. In replacement clutches females decreased clutch size and increased egg volume, thereby probably improving the survival probability of their offspring in less favourable conditions.Copyright 2002 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour     

Food availability affects the maternal transfer of androgens and antibodies into eggs of a colonial seabird

Mothers can improve the quality of their offspring by increasing the level of certain components in their eggs. To examine whether or not mothers increase deposition of such components in eggs as a function of food availability, we food-supplemented black-legged kittiwake females (Rissa tridactyla) before and during egg laying and compared deposition of androgens and antibodies into eggs of first and experimentally induced replacement clutches. Food-supplemented females transferred lower amounts of androgens and antibodies into eggs of induced replacement clutches than did non-food-supplemented mothers, whereas first clutches presented no differences between treatments. Our results suggest that when females are in lower condition, they transfer more androgens and antibodies into eggs to facilitate chick development despite potential long-term costs for juveniles. Females in prime condition may avoid these potential long-term costs because they can provide their chicks with more and higher quality resources.     

Reduced immunocompetence of nestlings in replacement clutches of the European magpie (Pica pica)

Laying date is one of the most important determinants of reproductive success and recruitment probability in birds. Late breeders usually fledge fewer chicks than individuals with earlier breeding dates, and fledglings produced late in the season have high mortality rates. Food availability and nestling mass have been evoked as the principal mechanistic links between laying date and offspring survival. Here we suggest that another factor may actually account for the difference in survival rate between early and late offspring: immunocompetence. We predicted that nestlings produced later in the season or in replacement clutches should have lower immune responses when challenged with an antigen, than early nestlings or nestlings produced in first clutches. This hypothesis was tested in a population of magpies (Pica pica), in which we experimentally induced breeding failure in a group of nests and compared the immune response of nestlings in replacement clutches with the immune response of first clutch nestlings. Cellular immune response, as measured by wing web swelling (a correlate of T-lymphocyte production after injection of phytohaemagglutinin-P), significantly decreased with hatching date and was significantly lower in nestlings of replacement clutches. Furthermore, coefficients of intraclutch variation in immune response were higher in nestlings of replacement clutches. This experiment demonstrates an inverse relationship between immune responsiveness and breeding date, and reduced recruitment probability of late nestlings may be a direct consequence of their inability to cope with parasites.     

The quality and the timing hypotheses evaluated using data on great reed warblers

The seasonal decline in reproductive success observed in many animal species may be caused by timing per se (timing hypothesis) or by variation in phenotypic quality between early and late breeding females (quality hypothesis). To distinguish between these two hypotheses, several studies of birds have used clutch removal experiments to manipulate breeding date. However, removal experiments also increase the females' previous reproductive effort due to the production of an extra clutch and a longer incubation period. According to life-history theory an increase in reproductive effort lowers future reproduction. Hence, life-history theory predicts lowered success of replacement broods for other reasons than expected from the timing hypothesis. Female great reed warblers, Acrocephalus arundinaceus , studied in Sweden are frequently exposed to nest predation, after which many lay replacement clutches. In order to examine possible effects of previous reproductive effort on different fitness components, we analysed the re-laying frequency and the reproductive success of replacement broods in relation to time of the season and previous reproductive effort (measured as the length of the previous breeding attempt, LPB). In clutches of re-laying females both the number of fledglings and the proportion of recruits were negatively correlated with LPB, whereas re-laying frequency and clutch size were not related to LPB. We expect such relationships to be present also among other species. Consequently, the use of replacement clutches, as for example in clutch removal experiments, in evaluations of the cause of the often observed seasonal decline in various fitness components, might exaggerate the importance of the timing hypothesis over the quality hypothesis.     

Time-dependent reproductive decisions in the blue tit

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

Reproductive investment in moa: a K-selected life-history strategy?

The exaggerated K-selected life-history strategy of moa has been suggested as an important factor causing their rapid extinction. Classically, this strategy is characterized by few, large offspring and low fecundity rates. Assuming clutches with one or two eggs as derived from the fossil record, we tested if eggs of moa were larger than the average of similar-sized birds, and estimated their unknown annual breeding frequencies. Therefore, we established allometries on body mass and different reproductive traits (i.e. egg mass, clutch mass and annual clutch mass). These were derived for r-selected (r-model) and for K-selected (K-model) bird species. In agreement with our initial expectations, moa had egg to body mass relations seen in “average” extant K-selected birds. While the K-model pointed to a clutch size of one or two eggs for moa corroborating fossil data and a K-selected life-history, the r-model predicted two to three times larger sizes. Under clutch sizes between one and two eggs and an annual clutch mass as observed in other similar-sized flightless island birds (e.g. rails, ratites), the annual clutch mass allometry suggested one to three clutches per year for moa. Even when assuming less than one brood per year (K-model predicts 0.5 clutches per year); annual clutch masses were still consistent with the K-model. Further studies are needed to clarify whether or not the reproductive strategy of flightless island birds and/or of the birds underlying the K-model fits better to the moa strategy. The approach presented herein, illustrates that combining biological and paleontological data can assist in the reconstruction of species traits, which are insufficiently or not preserved in fossils, but are necessary to understand the evolution of traits.     

Effects of a food supplementation experiment on reproductive investment and a post-mating sexually selected trait in magpies Pica pica

Food availability is an important factor affecting breeding success in birds. Food supplementation experiments in birds have in general focused on the effects on reproductive success in terms of female investment (laying date, clutch size, egg size), however, it is also known that the estimation of mate quality based on sexually selected signals influences female reproductive investment. In the particular case of magpies, females use nest size, a post-mating sexually selected signal, to assess male's likelihood to invest in reproduction, and accordingly adjust reproductive investment (clutch size). Then, the possible effects of food supplementation on female reproductive investment could be mediated by other variables related to parental quality, such as nest size in magpies. In the present study, we explore if higher food availability in a magpie territory affected both male sexually selected traits (i.e. nest size) and female reproductive investment (laying date, egg size, clutch size). We performed a food supplementation experiment in which we experimentally increased food availability in several magpie territories, keeping others as controls. In food-supplemented territories, males built significantly larger nests and females significantly increased egg size by 4.1% compared to control females. Results suggest that the continuous provisioning of protein rich food allowed magpie females to increase egg size. However, laying date and clutch size did not differ between control and food-supplemented magpie pairs. Food availability also affected the relationship between female reproductive investment and nest size. In control territories, females decreased their egg size in response to a larger nest, whereas a tendency for the opposite relationship was revealed in food-supplemented territories. We discuss the possibility that magpie females adopt different strategies for reproductive investment according to food availability.     

The ‘Widow Effect’ and its Consequences for Reproduction in Burying Beetles,Nicrophorus vespilloides (Coleoptera: Silphidae)

Burying beetles tend their young on small vertebrate carcasses, which serve as the sole source of food for the developing larvae. Single females are as proficient at rearing offspring as male-female pairs, yet males opt to remain with their broods throughout most of the larval development. One potential benefit of a male's extended residency is that it affords him the opportunity of additional copulations with the female, which could ensure his paternity in a replacement brood should the female's first egg clutch fail to hatch. We tested this hypothesis by manipulating males' access to their mates during the production of replacement clutches, using genetic colour markers to determine the paternity of offspring. Females were induced to produce a replacement brood by removing their first clutch of eggs. In one experimental treatment, we removed the female's mate upon the removal of her first egg clutch (‘widowed’ females); in a second treatment, the female was permitted to retain her mate up until she produced a replacement clutch. There was no significant difference in paternity between males removed from females before the initiation of replacement clutches and those permitted to remain with their mates. However, widowed females produced fewer offspring in replacement broods than did females permitted to retain their mates. This difference occurred primarily because a significantly greater proportion of widowed females opted not to produce a replacement clutch, a result we refer to as the ‘widow effect’. This widow effect was further shown in those replicates in which females of both treatments produced replacement clutches: widowed females took significantly longer to produce a replacement clutch than did females permitted to retain their mates. The loss of her mate could be a signal to a female that a take-over of the carcass is imminent. Her reluctance to produce a replacement clutch under these circumstances might constitute a strategy by which she conserves carrion for a subsequent reproductive attempt with an intruding male successful at ousting her previous mate. Regardless of its functional significance, the widow effect favours the extended residency of males and therefore contributes to the selective maintenance of male parental care.     

Incubation capacity limits maximum clutch size in black-tailed gulls Larus crassirostris

Factors determining clutch size of birds have long been the central issue in studies in life histories. It is assumed that the configuration of brood patches could limit the maximum clutch size. To test this hypothesis we manipulated clutch sizes and measured egg temperature as well as reproductive consequences in black-tailed gulls Larus crassirostris , which usually lay two egg clutches and have three brood patches. Mean egg temperature in 4-egg clutches (32.6±1.0°C) was significantly lower than in 2-egg (34.6±0.4°C) and 3-egg clutches (34.1±0.4°C), because egg temperature of the coolest egg within a 4-egg clutch was often substantially lower than the other three eggs. The proportion of eggs hatching from 4-egg clutches (11.6%) was lower than those of 2-egg (49.1%) and 3-egg clutches (52.0%). Four-egg clutches had longer incubation periods (29.6±1.3 day) than 2-egg (28.1±1.7 day) and 3-egg clutches (28.0 ±1.3 day). The results indicate that incubation capacity, which may be determined by the configuration of brood patches, limits the maximum clutch size in black tailed gulls, but not the actual clutch size typically laid.     

A comparative study of egg mass and clutch size in the Anseriformes

The factors explaining interspecific differences in clutch investment in precocial birds are poorly understood. We investigated how variations in clutch characteristics are related to environmental factors in a comparative study of 151 extant species of ducks, geese and swans (Anseriformes). Egg mass was negatively related to clutch size in a phylogenetic regression, a relationship that was much stronger when controlling for female mass. Nest placement was related to both egg size and clutch size, with cavity-nesting species laying more but smaller eggs. Egg size was positively correlated with incubation period and with female mass, and also with sexual size dimorphism (i.e. male mass relative to that of the female). Clutch size was not related to female mass. Species with long term pair bonds laid smaller clutches and larger eggs. The size of the breeding range was strongly positively correlated with clutch size and clutch mass, and its inclusion in multivariate models made other biogeographical variables (hemisphere, breeding latitude or insularity) non-significant. The small clutches in insular species appear to be a product of small range size rather than insularity per se. Our results suggest there is an evolutionary trade-off between clutch and egg size, and lend support to Lack’s resource-limitation hypothesis for the waterfowl.     

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.     

INTERACTIVE EFFECTS OF OFFSPRING SIZE AND TIMING OF REPRODUCTION ON OFFSPRING REPRODUCTION: EXPERIMENTAL,MATERNAL, AND QUANTITATIVE GENETIC ASPECTS

We demonstrate that egg size in side-blotched lizards is heritable (parent-offspring regressions) and thus will respond to natural selection. Because our estimate of heritability is derived from free-ranging lizards, it is useful for predicting evolutionary response to selection in wild populations. Moreover, our estimate for the heritability of egg size is not likely to be confounded by nongenetic maternal effects that might arise from egg size per se because we estimate a significant parent-offspring correlation for egg size in the face of dramatic experimental manipulation of yolk volume of the egg. Furthermore, we also demonstrate a significant correlation between egg size of the female parent and clutch size of her offspring. Because this correlation is not related to experimentally induced maternal effects, we suggest that it is indicative of a genetic correlation between egg size and clutch size. We synthesize our results from genetic analyses of the trade-off between egg size and clutch size with previously published experiments that document the mechanistic basis of this trade-off. Experimental manipulation of yolk volume has no effect on offspring reproductive traits such as egg size, clutch size, size at maturity, or oviposition date. However, egg size was related to offspring survival during adult phases of the life history. We partitioned survival of offspring during the adult phase of the life history into (1) survival of offspring from winter emergence to the production of the first clutch (i.e., the vitellogenic phase of the first clutch), and (2) survival of the offspring from the production of the first clutch to the end of the reproductive season. Offspring from the first clutch of the reproductive season in the previous year had higher survival during vitellogenesis of their first clutch if these offspring came from small eggs. We did not observe selection during these prelaying phases of adulthood for offspring from later clutches. However, we did find that later clutch offspring from large eggs had the highest survival over the first season of reproduction. The differences in selection on adult survival arising from maternal effects would reinforce previously documented selection that favors the production of small offspring early in the season and large offspring later in the season—a seasonal shift in maternal provisioning. We also report on a significant parent-offspring correlation in lay date and thus significant heritable variation in lay date. We can rule out the possibility of yolk volume as a confounding maternal effect—experimental manipulation of yolk volume has no effect on lay date of offspring. However, we cannot distinguish between genetic effects (i.e., heritable) and nongenetic maternal effects acting on lay date that arise from the maternal trait lay date per se (or other unidentified maternal traits). Nevertheless, we demonstrate how the timing of female reproduction (e.g., date of oviposition and date of hatching) affect reproductive attributes of offspring. Notably, we find that date of hatching has effects on body size at maturity and fecundity of offspring from later clutches. We did not detect comparable effects of lay date on offspring from the first clutch.     

6. THE PROBLEM OF NOMENCLATURE

Hockey, P. A. R. 1983. Aspects of the breeding biology of the African Black Oystercatcher. Ostrich 54:26-35. Fifty-five pairs of African Black Oystercatchers Haematopus moquini bred at Marcus Island in 1979–1980. Sixteen pairs laid replacement clutches: the mean interval between loss of the fist clutch and laying of a replacement clutch was 22,2 days. Mean inter-nest distance was 19,4 m. The modal clutch size was 2, with a mean of 1.74. Mean dimensions of 105 eggs were 60,7 × 40,l mm and mean fresh egg mass was 55,s g. There were differences in egg mass and dimensions between eggs in one- and two-egg clutches. Rate of egg loss was high, due mainly to depredation by Kelp Gulls Larus dominicanus promoted by human disturbance. Fledging success was lower at a disturbed site than at undisturbed sites, with highest chick mortality occurring in the first week of life. Mortality of first-year birds of eight days and older was estimated at 48% and 69% in two successive seasons. All juvenile birds dispersed from the natal sites, and were resighted up to 168 km away. Dispersed juveniles were concentrated at the edge of an area of high oystercatcher density.     

Measuring temporal variation in reproductive output reveals optimal resource allocation to reproduction in the northern grass lizard, Takydromus septentrionalis

We measured the reproductive output of Takydromus septentrionalis collected over 5 years between 1997 and 2005 to test the hypothesis that reproductive females should allocate an optimal fraction of accessible resources in a particular clutch and to individual eggs. Females laid 1–7 clutches per breeding season, with large females producing more, as well as larger clutches, than did small females. Clutch size, clutch mass, annual fecundity, and annual reproductive output were all positively related to female size (snout–vent length). Females switched from producing more, but smaller eggs in the first clutch to fewer, but larger eggs in the subsequent clutches. The mass-specific clutch mass was greater in the first clutch than in the subsequent clutches, but it did not differ among the subsequent clutches. Post-oviposition body mass, clutch size, and egg size showed differing degrees of annual variation, but clutch mass of either the first or the second clutch remained unchanged across the sampling years. The regression line describing the size–number trade-off was higher in the subsequent clutch than in the first clutch, but neither the line for first clutch, nor the line for the second clutch varied among years. Reproduction retarded growth more markedly in small females than in large ones. Our data show that: (1) trade-offs between size and number of eggs and between reproduction and growth (and thus, future reproduction) are evident in T. septentrionalis ; (2) females allocate an optimal fraction of accessible resources in current reproduction and to individual eggs; and (3) seasonal shifts in reproductive output and egg size are determined ultimately by natural selection.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 315–324.     

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.     

Different Temperature and Cooling Patterns at the Blunt and Sharp Egg Poles Reflect the Arrangement of Eggs in an Avian Clutch

Incubation is an energetically demanding process during which birds apply heat to their eggs to ensure embryonic development. Parent behaviours such as egg turning and exchanging the outer and central eggs in the nest cup affect the amount of heat lost to the environment from individual eggs. Little is known, however, about whether and how egg surface temperature and cooling rates vary among the different areas of an egg and how the arrangement of eggs within the clutch influences heat loss. We performed laboratory (using Japanese quail eggs) and field (with northern lapwing eggs) experiments using infrared imaging to assess the temperature and cooling patterns of heated eggs and clutches. We found that (i) the sharp poles of individual quail eggs warmed to a higher egg surface temperature than did the blunt poles, resulting in faster cooling at the sharp poles compared to the blunt poles; (ii) both quail and lapwing clutches with the sharp poles oriented towards the clutch centre (arranged clutches) maintained higher temperatures over the central part of the clutch than occurred in those clutches where most of the sharp egg poles were oriented towards the exterior (scattered clutches); and (iii) the arranged clutches of both quail and lapwing showed slower cooling rates at both the inner and outer clutch positions than did the respective parts of scattered clutches. Our results demonstrate that egg surface temperature and cooling rates differ between the sharp and blunt poles of the egg and that the orientation of individual eggs within the nest cup can significantly affect cooling of the clutch as a whole. We suggest that birds can arrange their eggs within the nest cup to optimise thermoregulation of the clutch.