Selection for synchronous breeding in the European starling

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

A more informative method for analyzing reproductive success

ABSTRACT.   Few investigators have examined how a female's prior history (i.e., number of clutches laid previously and whether those nesting attempts were successful or not) might influence the success of a particular clutch. Our objective was to determine the seasonal change in the mean number of fledglings successfully leaving nests in a population of Prairie Warblers ( Dendroica discolor ). To do so, we calculated the success of each clutch in the laying sequence (i.e., first, second, third, …, n ^th) and sorted them further by whether they were laid before (first-brood clutches) or after (second-brood clutches) the first brood fledged. In our sample population of 70 females we knew the fate of all clutches laid during a breeding season. Although the number of fledglings per successful clutch varied slightly, the probability that a clutch would be successful (i.e., produce at least one fledgling) and the number of fledglings per nest attempt increased during the breeding season. Thus, later clutches were more productive than earlier clutches. The low probability of producing a successful first clutch early in the breeding season would seem to favor selection for females that wait until later in the season to begin breeding. However, the seasonal reproductive success of females may be more dependent on breeding early (increasing the number of clutches laid in a season) than on the higher probability of success with later clutches. Our results indicate that further study of seasonal changes in reproductive success and their causes is needed.     

Differential maternal investment counteracts for late breeding in magpies Pica pica: an experimental study

Reproductive success in many avian populations declines throughout the breeding season. Two hypotheses have gained attention to explain such a decline: the "timing" hypothesis proposes that deteriorating food availability causes the decline in reproductive success (causal effect of breeding time), whereas the "quality" hypothesis proposes that individuals of lower phenotypic quality reproduce at the end of the breeding season, causing the correlation between breeding time and breeding success. We tested both of these hypotheses in a monogamous single breeder, the magpie Pica pica , by experimentally inducing some pairs to lay a replacement clutch, after removal of the first clutch. The first clutch was left in the nest of another magpie pair (matched by laying date and clutch size), and incubated and raised by these foster parents. In this way we obtained two clutches from the same magpie pair with full siblings raised in conditions of the first and second reproductive attempts. High quality pairs (with laying dates in the first half of the breeding season) reached similar breeding success in replacement clutches as compared to first clutches of the same female. In addition, experimental pairs reared significantly more offspring of similar quality in their replacement clutches as compared to late-season first clutches, thereby suggesting that late season first clutches were produced by pairs of lower phenotypic quality. Results indicate that high quality pairs trade-off clutch size for larger eggs in replacement clutches, which could help magpie pairs to partly compensate for poorer environmental conditions associated with a delayed breeding attempt.     

Breeding success and the question of clutch size of Northwestern Crows Corvus caurinus

The breeding biology of two island populations of the Northwestern Crow was studied in British Columbia over a period of five years. Both populations laid an average of four eggs per clutch, of which approximately 75% hatched. More young survived to fledge on Mandate than on Mitlenatch Island. The young from nests close (< 100 m) to the beach on Mitlenatch had a higher survival rate than those from further inland. The number of eggs lost or failing to hatch was lowest in clutches of four eggs. There was a non-significant trend for chick survival to be inversely related to clutch size. The net result was that clutches of five eggs produced no more young than did clutches of four eggs. It is suggested that the major limiting factor on clutch size is food availability but the decreased hatching success of any remaining eggs once hatching has started is also a contributing factor.     

Increased survival and breeding performance of double breeders in little penguins Eudyptula minor,New Zealand: evidence for individual bird quality?

The little penguin Eudyptula minor is unique among penguin species in being able to fledge chicks from two clutches in one breeding season. Pairs laying two clutches in a given season make a higher reproductive investment, and may be rewarded by a higher reproductive success as they may raise twice as many chicks as pairs laying one clutch. The higher effort made by pairs laying two clutches could correlate negatively with survival, future reproductive performance or offspring survival, indicating a cost of reproduction. Conversely, a positive relationship between the number of clutches produced in a given breeding season and survival, future reproductive performance or offspring survival would indicate that birds laying two clutches belonged to a category of birds with higher fitness, compared to birds laying only one clutch in the season. In this study we used a long‐term data set taken from an increasing population of little penguins in Otago, SE New Zealand. We modelled the relationship between the number of clutches laid in a breeding season and survival probability, reproductive performance in the next breeding season and first year survival of offspring using capture‐recapture modelling.
Birds laying two clutches produced 1.7 times more fledglings during a breeding season than pairs laying one clutch. We found that birds laying two clutches had a higher probability of breeding in the following breeding season, a higher probability of laying two clutches in the following breeding season and a higher survival probability. There was no overall difference in post‐fledging survival between the young of birds producing one clutch and the young of birds producing two clutches. However, the survival of young of single clutch breeders declined with laying date, whereas the young of double clutch breeders had the same survival rate irrespective of laying date. For a subset of data with birds of known age, we found evidence that the probability of laying two clutches increased with age. However, there were also indications for differences among birds in the tendency to lay two clutches that could not be attributed to age. We tentatively interpret our results as evidence of quality difference among little penguin breeders.     

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.     

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.     

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.     

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.     

Maternal investment in reproduction and its consequences in leatherback turtles

Maternal investment in reproduction by oviparous non-avian reptiles is usually limited to pre-ovipositional allocations to the number and size of eggs and clutches, thus making these species good subjects for testing hypotheses of reproductive optimality models. Because leatherback turtles (Dermochelys coriacea) stand out among oviparous amniotes by having the highest clutch frequency and producing the largest mass of eggs per reproductive season, we quantified maternal investment of 146 female leatherbacks over four nesting seasons (2001–2004) and found high inter- and intra-female variation in several reproductive characteristics. Estimated clutch frequency [coefficient of variation (CV) = 31%] and clutch size (CV = 26%) varied more among females than did egg mass (CV = 9%) and hatchling mass (CV = 7%). Moreover, clutch size had an approximately threefold higher effect on clutch mass than did egg mass. These results generally support predictions of reproductive optimality models in which species that lay several, large clutches per reproductive season should exhibit low variation in egg size and instead maximize egg number (clutch frequency and/or size). The number of hatchlings emerging per nest was positively correlated with clutch size, but fraction of eggs in a clutch yielding hatchlings (emergence success) was not correlated with clutch size and varied highly among females. In addition, seasonal fecundity and seasonal hatchling production increased with the frequency and the size of clutches (in order of effect size). Our results demonstrate that female leatherbacks exhibit high phenotypic variation in reproductive traits, possibly in response to environmental variability and/or resulting from genotypic variability within the population. Furthermore, high seasonal and lifetime fecundity of leatherbacks probably reflect compensation for high and unpredictable mortality during early life history stages in this species.     

Relationship between clutch size,egg volume and hatching success in a Yellow-legged Gull Larus michahellis colony in south-eastern Tunisia

This study aimed to test the hypothesis that clutch size covaries with egg volume and hatching success in the Yellow-legged Gull Larus michahellis. We determined clutch size and egg volume in a sample of 131 nests, and we used the data to check whether egg volume varied among nests according to clutch size, while taking into account the effects of egg laying order. We also estimated hatching success rate and investigated the relationship between hatching success and clutch size. Egg volume varied among clutches according to clutch size, with eggs being larger in three-egg clutches than in two-egg clutches. Moreover, three-egg clutches showed higher daily survival rates, and hence hatching success, than two-egg clutches. Overall, our results suggest that in the Yellow-legged Gull clutch size covaries with egg volume and hatching success, which could possibly reflect an age effect through different mechanisms. Indeed, older females could be hypothesised to exhibit greater breeding performance than younger females because of their higher experience in tapping energy resources for egg formation and defending nests from dangers. Moreover, due to their age, older females are likely to have lower residual reproductive potential and should invest more heavily in current breeding attempts.     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.     

Timing of current reproduction directly affects future reproductive output in European coots

Life-history theory suggests that the variation in the seasonal timing of reproduction within populations may be explained on the basis of individual optimization. Optimal breeding times would vary between individuals as a result of trade-offs between fitness components. The existence of such trade-offs has seldom been tested empirically. We experimentally investigated the consequences of altered timing of current reproduction for future reproductive output in the European coot (Fulica atra). First clutches of different laying date were cross-fostered between nests, and parents thereby experienced a delay or an advance in the hatching date. The probability and success of a second brood, adult survival until and reproduction in the next season were then compared to the natural variation among control pairs. Among control pairs the probability of a second brood declined with the progress of season. Delayed pairs were less likely and advanced pairs were more likely to produce a second brood. These changes were quantitatively as predicted from the natural seasonal decline. The number of eggs in the second clutch was positively related to egg number in the first clutch and negatively related to laying date. Compared to the natural variation, delayed females had more and advanced females had fewer eggs in their second clutch. The size of the second brood declined with season, but there was no significant effect of delay or advance. Local adult survival was higher following a delay and reduced following an advance. The effect of the experiment on adult survival was independent of sex. Laying date and clutch size of females breeding in the next year were not affected by treatment. The study demonstrates the existence of a trade-off between increased probability of a second brood and decreased parental survival for early breeding. Timing-dependent effects of current reproduction on future reproductive output may thus play an important role in the evolution of the seasonal timing of reproduction.     

Seasonal shifts in clutch size and egg size in the side-blotched lizard,Uta stansburiana Baird and Girard

Summary There is evidence that the side-blotched lizard, Uta stansburiana, and some other organisms of temperate latitudes produce fewer and larger eggs as the reproductive season progresses. There are at least two models that could explain this phenomenon.Proponents of the parental investment model claim that females are selected to increase egg size, at the cost of clutch size, late in the season in order to produce larger and competitively superior hatchlings at a time when food for hatchlings is in low supply and when juvenile density is high. In this model the selective agent is relative scarcity of food available to hatchlings late in the reproductive season, and the adaptive response is production of larger offspring.The alternative explanation (bet-hedging model) proposed in this paper is based on the view that the amount of food available to females for the production of late-season clutches is unpredictable, and that selection has favored conservatively small clutches in the late season to insure that each egg is at least minimally provisioned. Smaller clutches, which occur most frequently late in the season, are more likely to consist of larger eggs, compared to larger clutches, for two reasons. Firstly, unlike birds, oviparous lizards cannot alter parental investment after their eggs are deposited, and therefore, in cases of fractional optimal clutch size, the next lower integral clutch size is selected with the remaining reproductive energy allocated to increased egg size. With other factors constant, eggs of smaller clutches will increase more in size than eggs of larger clutches when excess energy is divided among the eggs of a clutch. Secondly, unanticipated energy that may become available for reproduction during energy-rich years will similarly increase egg size a greater amount if divided among fewer eggs.     

安徽滁州雌性丽斑麻蜥繁殖特征

在安徽滁州地区丽斑麻蜥(Eremias argus)年产两窝卵。窝卵数及窝卵重与雌体体长呈正相关,相对窝卵重与雌体体长无关,卵重与窝卵数无关。窝卵数、窝卵重及卵重在窝序间无明显的差异。卵长径与卵短径呈正相关,卵长径与窝卵数呈负相关,而卵短径与窝卵数无关。雌体主要通过增加窝卵数增加繁殖输出。     

Variation in reproductive output of marine turtles

Most marine turtle species are non-annual breeders and show variation in both the number of eggs laid per clutch and the number of clutches laid in a season. Large levels of inter-annual variation in the number of nesting females have been well documented in green turtle nesting populations and may be linked to environmental conditions. Other species of marine turtle exhibit less variation in nesting numbers. This inter-specific difference is thought to be linked to trophic status. To examine whether individual reproductive output is more variable in the herbivorous green turtle (Chelonia mydas Linneaeus 1758) than the carnivorous loggerhead (Caretta caretta Linneaeus 1758), we examined the nesting of both species in Cyprus over nine seasons. Green turtles showed slower annual growth rates (0.11 cm year⁻¹ curved carapace length (CCL) and 0.27 cm year⁻¹ curved carapace width (CCW)) than loggerhead turtles (0.36 cm year⁻¹ CCL, 0.51 cm year⁻¹ CCW). CCL was highly correlated to mean clutch size in both green (R²=0.51) and loggerhead turtles (R²=0.61) and maximal clutch size of green turtles (R²=0.58). Larger females did not lay a greater number of clutches or have a shorter remigration interval than smaller females of either species. On average, the size of green turtle clutches increased and that of loggerhead turtles decreased as the season progressed. Individual green turtles, however, produced more eggs per clutch through the season to a maximum in the third or fourth clutch. In loggerhead turtles, clutches 1-4 were very similar in size but the fifth clutch was 38% smaller than the first. No individuals of either species were recorded laying more than five clutches. Green turtles may not be able to achieve their maximum reproductive output with respect to clutch size throughout the season, whereas only loggerhead turtles laying five clutches (n=5) appear to become resource depleted. Green turtles nesting in years when large numbers of nests were recorded laid a greater number of clutches than females nesting in years with lower levels of nesting.     

Factors affecting variation in the reproductive investment of female treefrogs,Hyla intermedia

In anurans, fecundity (clutch size) is the most important determinant of female reproductive success. We investigated three possible causes responsible for fecundity variation in female Italian treefrogs, Hyla intermedia, during four breeding seasons: (i) variation in morphological (body size and condition) and life-history (age) traits; (ii) variation in the tradeoff between the number and the size of eggs; (iii) seasonal effects and within-season differences in the timing of deposition. At the population level, we found no evidence for a tradeoff between the number and the size of eggs, because they both correlated positively with females’ body size. Conversely, neither age nor post-spawning body condition showed any effect on female reproductive investment. Independent of body size, we found no evidence for variation in reproductive effort among different breeding seasons, but strong evidence for a decrease of clutch size and an increase of egg size with the advancing of a breeding season. To test for the functional significance of the observed temporal variation in allocation strategy, we carried out a rearing experiment in semi-natural conditions on a random sample of ten clutches. The experiment showed a negative effect of clutch size and a positive effect of egg size on both tadpole growth and developmental rates, suggesting that reproductive investment, although constrained by body size, can be adjusted by females to the time of deposition to increase the chances of offspring survival.     

Differential investment and costs during avian incubation determined by individual quality: an experimental study of the common eider (Somateria mollissima)

Individuals of different quality may have different investment strategies, shaping responses to experimental manipulations, thereby rendering the detection of such patterns difficult. However, previous clutch-size manipulation studies have infrequently incorporated individual differences in quality. To examine costs of incubation and reproductive investment in relation to changes in clutch size, we enlarged and reduced natural clutch sizes of four and five eggs by one egg early in the incubation period in female common eiders (Somateria mollissima), a sea duck with an anorectic incubation period. Females that had produced four eggs (lower quality) responded to clutch reductions by deserting the nest more frequently but did not increase incubation effort in response to clutch enlargement, at the cost of reduced hatch success of eggs. Among birds with an original clutch size of five (higher quality), reducing and enlarging clutch size reduced and increased relative body mass loss respectively without affecting hatch success. In common eiders many females abandon their own ducklings to the care of other females. Enlarging five-egg clutches led to increased brood care rate despite the higher effort spent incubating these clutches, indicating that the higher fitness value of a large brood is increasing adult brood investment. This study shows that the ability to respond to clutch-size manipulations depends on original clutch size, reflecting differences in female quality. Females of low quality were reluctant to increase investment at the cost of lower hatch success, whereas females of higher quality apparently have a larger capacity both to increase incubation effort and brood care investment.     

Breeding biology of the Barn Owl Tyto alba in central Mali

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

Female reproductive characteristics of the Balkan wall lizard (<Emphasis Type="Italic">Podarcis taurica</Emphasis>) in the northwestern periphery of its range

We present data on the female reproductive traits of the Balkan wall lizard in the Deliblato Sand, a large continental sandland in the Pannonian area in the northwestern periphery of the species range. The clutch and egg characteristics of the population were investigated on the basis of clutches laid in laboratory conditions by gravid females captured in one locality. Balkan wall lizards produced at least two clutches in a breeding season. Individual females laid clutches of commonly two (range 1–4) eggs. The female body size had no effect on clutch and egg size. There was no trade-off between egg size and clutch size.