Does time of the season influence filial cannibalism in the sand goby, <Emphasis Type="Italic">Pomatoschistus minutus</Emphasis>?

According to life-history theory, filial cannibalism by fish that breed over one season only should be more beneficial early than late in the season if they eat eggs to invest energy into later clutches. Also, filial cannibalism may be more costly late in the season if finding ripe females for replacing eaten eggs is harder then. On the other hand, offspring hatching early may have a competitive advantage over fry hatching late and hence provide higher fitness to the parent. Using data collected over three successive years, I tested if sand goby males are more prone to eat of their eggs early than late in the reproductive season. I found no difference in the amount of eggs eaten or in the frequency of males eating the whole clutch between early and late in the season. Furthermore, there was no difference in the frequency of males who ate parts of their clutches, early compared to late. This might reflect a trade-off between quality (early hatching offspring) and quantity (producing as many offspring as possible over a long reproductive season). If so, the lack of seasonal pattern of filial cannibalism found in sand gobies might be the result of opposing selection pressures.     

Asynchronous hatching and brood reduction by filial cannibalism in the burying beetle Nicrophorus quadripunctatus

Despite decades of intensive research, there is still much debate about the adaptive significance of asynchronous hatching. A major obstacle in understanding the significance of this process is the difficulty involved in separating the hypotheses that explain asynchronous hatching as an adaptive trait from those that explain it as a by-product of physiological constraints on hatching or egg-laying patterns. We investigated the burying beetle Nicrophorus quadripunctatus, a species in which the parent can eliminate less-adaptive offspring (e.g., slower-growing offspring) by filial cannibalism and adjust the age structure of offspring to an adaptive pattern. The main aim of this study was to determine the age composition of offspring that survived and to determine the effect of larval growth on filial cannibalism. We investigated how the point in time at which each group of larvae hatched affects the timing of filial cannibalism by the female parent. We found that N. quadripunctatus exhibited asynchronous hatching, and reared larvae of different ages. We also found that later-hatching larvae had lower survival and growth rates; therefore, filial cannibalism plays a role in eliminating later-arriving, slower-growing, and hence less-adaptive offspring.     

When to care for, abandon, or eat your offspring: the evolution of parental care and filial cannibalism

Parental care and filial cannibalism (the consumption of one's own offspring) co-occur in many animals. While parental care typically increases offspring survival, filial cannibalism involves the killing of one's young. Using an evolutionary ecology approach, we evaluate the importance of a range of factors on the evolution of parental care and filial cannibalism. Parental care, no care/total abandonment, and filial cannibalism evolved and often coexisted over a range of parameter space. While no single benefit was essential for the evolution of filial cannibalism, benefits associated with adult or offspring survival and/or reproduction facilitated the evolution of cannibalism. Our model highlights the plausibility of a range of alternative hypotheses. Specifically, the evolution of filial cannibalism was enhanced if (1) parents could selectively cannibalize lower-quality offspring, (2) filial cannibalism increased egg maturation rate, (3) energetic benefits of eggs existed, or (4) cannibalism increased a parent's reproductive rate (e.g., through mate attractiveness). Density-dependent egg survivorship alone did not favor the evolution of cannibalism. However, when egg survival was density dependent, filial cannibalism invaded more often when the density dependence was relatively more intense. Our results suggest that population-level resource competition potentially plays an important role in the evolution of both parental care and filial cannibalism.     

You eat what you are: personality‐dependent filial cannibalism in a fish with paternal care

Many animal parents invest heavily to ensure offspring survival, yet some eventually consume some or all of their very own young. This so‐called filial cannibalism is known from a wide range of taxa, but its adaptive benefit remains largely unclear. The extent to which parents cannibalize their broods varies substantially not only between species, but also between individuals, indicating that intrinsic behavioral differences, or animal personalities, might constitute a relevant proximate trigger for filial cannibalism. Using a marine fish with extensive paternal care, the common goby (Pomatoschistus microps), we investigated the influence of animal personality on filial cannibalism by assessing (1) behavioral consistency across a breeding and a nonbreeding context; (2) correlations between different breeding (egg fanning; filial cannibalism) and nonbreeding (activity) behaviors, and, in a separate experiment; (3) whether previously established personality scores affect filial cannibalism levels. We found consistent individual differences in activity across contexts. Partial filial cannibalism was independent of egg fanning but correlated strongly with activity, where active males cannibalized more eggs than less active males. This pattern was strong initially but vanished as the breeding season progressed. The incidence of whole clutch filial cannibalism increased with activity and clutch size. Our findings indicate that filial cannibalism cannot generally be adjusted independently of male personality and is thus phenotypically less plastic than typically assumed. The present work stresses the multidimensional interaction between animal personality, individual plasticity and the environment in shaping filial cannibalism.     

Experimental demonstration of the insurance value of extra eggs in an obligately siblicidal seabird

A variety of organisms regularly produce more offspring thanthey raise. Despite the apparent energetic waste of such areproductive tactic, overproduction may be favored by naturalselection in some cases. One such case is when surplus offspringcan serve as replacements, or insurance, for failed siblings.We tested the Insurance Egg Hypothesis (IEH) as an explanationfor the overproduction of offspring in an obligately siblicidal seabird, the Nazca booby (Sula grant)i, which fledges a maximumof one nestling regardless of its clutch size. We manipulatedclutch sizes within the range of natural variation encounteredin this species (one-two eggs). The IEH predicts that parentswith two-egg clutches should have higher reproductive successthan those with one-egg clutches because the second egg canprovide a nestling when the first egg fails to hatch, or when the first chick dies young. Consistent with the IEH, naturalone-egg clutches that were enlarged to two eggs produced morehatchlings and fledglings than control one-egg clutches did,and natural two-egg clutches that were reduced to one egg producedfewer hatchlings and fledglings than control two-egg clutchesdid. We also evaluated aspects of the Individual Optimization Hypothesis, which proposes that individual optimal clutch sizesdiffer, as an explanation for clutch size variation in thisspecies. In Nazca boobies, selection driven by replacementvalue appears to favor clutches larger than one even thoughfinal brood size is invariably one. One-egg clutches may be produced by parents experiencing some proximate limitation,such as a lack of food.     

Parents benefit from eating offspring: density-dependent egg survivorship compensates for filial cannibalism

Abstract Why should animals knowingly consume their own young? It is difficult to imagine many circumstances in which eating one's own young (i.e., filial cannibalism) actually increases an individual's fitness; however, filial cannibalism commonly co‐occurs with parental care in fishes. The evolutionary significance of filial cannibalism remains unclear. The most commonly accepted explanation is that filial cannibalism is a mechanism by which caring males gain energy or nutrients that they reinvest into future reproduction, thereby increasing net reproductive success. There is mixed support for this hypothesis and, at best, it can only explain filial cannibalism in some species. A recent alternative hypothesis suggests that filial cannibalism improves the survivorship of remaining eggs by increasing oxygen availability, and thus increases current reproductive success. This theory has received little attention as of yet. We evaluated the hypothesis of oxygen‐mediated filial cannibalism in the sand goby by examining the effect of oxygen and egg density on the occurrence of filial cannibalism, evaluating the effects of partial clutch cannibalism on the survivorship of remaining eggs, and comparing potential costs and benefits of filial cannibalism related to the net number of eggs surviving. Indeed, we found that oxygen level and egg density affected the occurrence of cannibalism and that simulated partial clutch cannibalism improved survivorship of the remaining eggs. Additionally, because increased egg survivorship, stemming from partial egg removal, compensated for the cost of cannibalism (i.e., number of eggs removed) at a range of cannibalism levels, filial cannibalism potentially results in no net losses in reproductive success. However, oxygen did not affect egg survivorship. Thus, we suggest a more general hypothesis of filial cannibalism mediated by density‐dependent egg survivorship.     

Maternal manipulation of hatching asynchrony limits sibling cannibalism in the predatory mite Phytoseiulus persimilis

1. Sibling cannibalism is a common phenomenon in the animal kingdom but entails a high risk of direct and inclusive fitness loss for the mother and her offspring. Therefore, mechanisms limiting sibling cannibalism are expected to be selected for. One way of maternal manipulation of sibling cannibalism is to influence hatching asynchrony between nearby laid eggs. This has rarely been tested experimentally. 2. We examined the ability of ovipositing females of the cannibalistic predatory mite Phytoseiulus persimilis to influence the occurrence of sibling cannibalism among offspring by manipulating hatching asynchrony of nearby laid eggs. 3. In the first experiment, we assessed the occurrence of sibling cannibalism in relation to the hatching interval (24 h and 48 h) between nearby laid eggs. In the second experiment, we tested whether ovipositing females discriminate sites containing young (24-h old) and old (48-h old) eggs, fresh and old traces (metabolic waste products and possibly pheromones) left by the same female (24 h and 48 h ago), or young eggs plus fresh female traces and old eggs plus old female traces. Both experiments were conducted with and without prey. 4. Without prey, siblings were more likely to cannibalize each other if the hatching interval between nearby laid eggs was short (24 h). Cannibalism occurred less often when senior siblings (protonymphs) experienced a delay in the opportunity to cannibalize junior siblings (larvae). 5. Independent of prey availability, females preferentially added new eggs to sites containing old eggs plus old female traces but did neither distinguish between young and old eggs presented without own traces nor between fresh and old traces presented without eggs. 6. We discuss cue perception and use by P. persimilis females and contrast the outcome of our experiments and theoretical predictions of sibling cannibalism. We conclude that P. persimilis mothers increase hatching asynchrony of nearby laid eggs to prevent sibling cannibalism on the last produced offspring. Such a behaviour may be considered a simple form of maternal care increasing the survival prospects of offspring.     

Maternal investment increases with altitude in a frog on the Tibetan Plateau

Reproducing females can allocate energy between the production of eggs or offspring of different size or number, both of which can strongly influence fitness. The physical capacity to store developing offspring imposes constraints on maximum clutch volume, but individual females and populations can trade off whether more or fewer eggs or offspring are produced, and their relative sizes. Harsh environments are likely to select for larger egg or offspring size, and many vertebrate populations compensate for this reproductive investment through an increase in female body size. We report a different trade‐off in a frog endemic to the Tibetan Plateau, Rana kukunoris. Females living at higher altitudes (n = 11 populations, 2000–3500 m) produce larger eggs, but without a concomitant increase in female body size or clutch size. The reduced diel and seasonal activity at high altitudes may impose constraints on the maximum body size of adult frogs, by limiting the opportunity for energy accumulation. Simultaneously, producing larger eggs likely helps to increase the rate of embryonic development, causing tadpoles to hatch earlier. The gelatinous matrix surrounding eggs, more of which is produced by large females, may help buffer developing embryos from temperature fluctuations or offer protection from ultraviolet radiation. High‐altitude frogs on the Tibetan Plateau employ a reproductive strategy that favours large egg size independent of body size, which is unusual in amphibians. The harsh and unpredictable environmental conditions at high altitudes can thus impose strong and opposing selection pressures on adult and embryonic life stages, both of which can simultaneously influence fitness.     

Spawning behaviour and acoustic communication in Atlantic cod

Cannibalism of small numbers of offspring by a parent has been proposed as an adaptive parental strategy, by providing energy to support parental care. There are few empirical studies, however, to support this hypothesis. The beaugregory damselfish, Stegastes leucostictus , is a marine teleost that does not actively ventilate its eggs by fanning them. Partial cannibalism is common in this species, but in field studies was found to be unrelated to ration level. Filial cannibalism differed from predation in the pattern of egg eating; filial cannibalism was characterised by a random pattern of egg loss from a clutch rather than an aggregated distribution. Embryos developed quicker and had higher survival rates when they were at low densities and in nest sites where oxygen levels were high, and experimental reduction of oxygen levels increased rates of filial cannibalism. Here I present a hypothesis for filial cannibalism in the beaugregory damselfish; males cannibalise egg clutches in order to reduce clutch density and improve oxygen supply to the remaining embryos. I use a model of filial cannibalism to demonstrate how oxygen mediated cannibalism may be adaptive, and discuss the evolution of filial cannibalism in the beaugregory damselfish and other teleosts.     

Hatching order and sex ratio in Southern Grey Shrike Lanius meridionalis in relation to clutch size

Despite major advances in sex ratio theory, how offspring sex should vary with hatching order remains unclear. We examine nestling sex ratio in the Southern Grey Shrike Lanius meridionalis according to hatching order and clutch size. Southern Grey Shrike nestlings present a different sex ratio with body‐mass rank order depending on clutch size. When the clutch size was five eggs (with a very low risk of brood reduction; 13%) the less costly sex (male) was found at the end of the body mass hierarchy. However, when clutch size was six eggs (with a high risk of brood reduction; 42%) the larger sex (female) was found at intermediate positions in the hatching order, possibly to decrease competitive asymmetries.     

Adaptive filial cannibalism in the beaugregory damselfish

Cannibalism of small numbers of offspring by a parent has been proposed as an adaptive parental strategy, by providing energy to support parental care. There are few empirical studies, however, to support this hypothesis. The beaugregory damselfish, Stegastes leucostictus, is a marine teleost that does not actively ventilate its eggs by fanning them. Partial cannibalism is common in this species, but in field studies was found to be unrelated to ration level. Filial cannibalism differed from predation in the pattern of egg eating; filial cannibalism was characterised by a random pattern of egg loss from a clutch rather than an aggregated distribution. Embryos developed quicker and had higher survival rates when they were at low densities and in nest sites where oxygen levels were high, and experimental reduction of oxygen levels increased rates of filial cannibalism. Here I present a hypothesis for filial cannibalism in the beaugregory damselfish; males cannibalise egg clutches in order to reduce clutch density and improve oxygen supply to the remaining embryos. I use a model of filial cannibalism to demonstrate how oxygen mediated cannibalism may be adaptive, and discuss the evolution of filial cannibalism in the beaugregory damselfish and other teleosts.     

Filial cannibalism improves survival and development of beaugregory damselfish embryos

Cannibalism of small numbers of offspring by a parent has been proposed as an adaptive parental strategy, by providing energy to support parental care. However, there are few empirical studies to support this hypothesis. We conducted field and laboratory experiments to investigate partial filial cannibalism in Stegastes leucostictus, a coral reef fish with paternal care. Partial cannibalism was shown to be common, and males were found to remove developing embryos from throughout a clutch in a random pattern, rather than in the more aggregated pattern seen during embryo predation. Males that received a diet supplement grew faster than control males, but did not engage in less cannibalism. Also, males did not concentrate cannibalism on early embryonic stages with the highest energetic value. Experimental reduction of embryo densities was found to significantly increase embryo development rate and survival from egg deposition to hatching, and experimental reduction of oxygen levels significantly increased rates of partial filial cannibalism by males. Artificial spawning sites with low oxygen levels were avoided by spawning females, and cannibalism rates by males were higher. We propose that partial filial cannibalism serves as an adaptive parental strategy to low oxygen levels in S. leucostictus by increasing the hatching success of embryos.     

Maternal hatching synchronization in a subsocial burrower bug mitigates the risk of future sibling cannibalism

Sibling cannibalism—the killing and consumption of conspecifics within broods—carries a high risk of direct and inclusive fitness loss for parents and offspring. We reported previously that a unique vibrational behavior shown by the mother of the subsocial burrower bug, Adomerus rotundus (Heteroptera: Cydnidae), induced synchronous hatching. Maternal regulation may be one of the most effective mechanisms for preventing or limiting sibling cannibalism. Here, we tested the hypothesis that synchronous hatching induced by maternal vibration in A. rotundus prevents sibling cannibalism. Mothers and their mature egg masses were allocated to three groups: synchronous hatching by maternal vibration (SHmv), synchronous hatching by artificial vibration (SHav), and asynchronous hatching (AH). We then investigated the influence of each hatching strategy on the occurrence of sibling cannibalism of eggs and early‐instar nymphs in the laboratory. No difference in the proportion of eggs cannibalized was observed among the three groups. However, the proportion of nymphs cannibalized was higher in the AH group than in the SHmv group. The difference in the number of days to first molting within clutch was significantly higher in the AH group than in the SHmv group. Junior nymphs were sometimes eaten by senior nymphs. However, immediately after molting, senior nymphs were at a high risk of being eaten by junior nymphs. Our results indicate that synchronous hatching of A. rotundus is necessary to mitigate the risk of sibling cannibalism.     

A limit on the extent to which increased egg size can compensate for a poor postnatal environment revealed experimentally in the burying beetle,Nicrophorus vespilloides

It is often assumed that there is a positive relationship between egg size and offspring fitness. However, recent studies have suggested that egg size has a greater effect on offspring fitness in low‐quality environments than in high‐quality environments. Such observations suggest that mothers may compensate for poor posthatching environments by increasing egg size. In this paper we test whether there is a limit on the extent to which increased egg size can compensate for the removal of posthatching parental care in the burying beetle, Nicrophorus vespilloides. Previous experiments with N. vespilloides suggest that an increased egg size can compensate for a relatively poor environment after hatching. Here, we phenotypically engineered female N. vespilloides to produce large or small eggs by varying the amount of time they were allowed to feed on the carcass as larvae. We then tested whether differences between these groups in egg size translated into differences in larval performance in a harsh postnatal environment that excluded parental care. We found that females engineered to produce large eggs did not have higher breeding success, and nor did they produce larger larvae than females engineered to produce small eggs. These results suggest that there is a limit on the extent to which increased maternal investment in egg size can compensate for a poor posthatching environment. We discuss the implication of our results for a recent study showing that experimental N. vespilloides populations can adapt rapidly to the absence of posthatching parental care.     

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.     

The evolution of filial cannibalism and female mate choice strategies as resolutions to sexual conflict in fishes

Filial cannibalism (the consumption of one's own viable offspring) is common among fish with paternal care. In this study, I use a computer simulation to study simultaneous evolution of male filial cannibalism and female mate choice. Under certain conditions, selection on parental males favors filial cannibalism. When filial cannibalism increases a male's probability to raise the current brood successfully, filial cannibalism also benefits the female. However, when egg eating is a male investment into future reproduction, a conflict between female and male interests emerges. Here I investigate how female discrimination against filial cannibals affects evolution of filial cannibalism and how different female choice criteria perform against filial cannibalism. The introduction of discriminating females makes the fixation of filial cannibalism less likely. I introduced three different female choice criteria: (1) females who could discern a male's genotype, that is, whether the male was going to eat eggs as an investment in future reproductive events; (2) energy-choosing females that preferred to mate with males who had enough energy reserves to live through the current brood cycle without consuming eggs; and (3) females that preferred to mate with already mated males, that is, males with eggs in their nest. Genotype choice never coexisted with filial cannibals at fixation and filial cannibals were unable to invade a population with genotype-choosing females. Energy choice was successful only when males had high energy reserves and were less dependent on filial cannibalism as an alternative energy source. The egg choosers frequently coexisted with the cannibals at fixation. When the female strategies were entered simultaneously, the most frequent outcome for low mate sampling costs was that both the cannibals and the egg choice was fixed and all other strategies went extinct. These results suggest that sexual conflicts may not always evolve toward a resolution of the conflict, but sometimes the stable state retains the conflict. In the present case, this was because the egg-preference strategy had a higher fitness than the other female strategies. The outcome of this simulation is similar to empirical findings. In fish with paternal care, male filial cannibalism and female preference for mates with eggs commonly co-occur.     

Population density, body size, and phenotypic plasticity of brood size in a burying beetle

Theory predicts that organisms living in heterogeneous environmentswill exhibit phenotypic plasticity. One trait that may be particularlyimportant in this context is the clutch or brood size becauseit is simultaneously a maternal and offspring characteristic.In this paper, I test the hypothesis that the burying beetle,Nicrophorus orbicollis, adjusts brood size, in part, in anticipationof the reproductive environment of its adult offspring. N. orbicollisuse a small vertebrate carcass as a food resource for theiryoung. Both parents provide parental care and actively regulatebrood size through filial cannibalism. The result is a positivecorrelation between brood size and carcass size. Adult bodysize is an important determinant of reproductive success forboth sexes, but only at higher population densities. I testthree predictions generated by the hypothesis that beetles adjustbrood size in response to population density. First, averageadult body size should vary positively with population density.Second, brood size on a given-sized carcass should be larger(producing more but smaller young) in low-density populationsthan in high-density populations. Third, females should respondadaptively to changes in local population density by producinglarger broods when population density is low and small broodswhen population density is high. All three predictions weresupported using a combination of field and laboratory experiments.These results (1) show that brood size is a phenotypically plastictrait and (2) support the idea that brood size decisions arean intergenerational phenomenon that varies with the anticipatedcompetitive environment of the offspring.     

Larger clutch size increases fledging success and offspring quality in a precocial species

1. We tested the hypothesis that the ability of parents to raise viable offspring limits clutch size in the greater snow goose ( Anser caerulescens atlanticus L.), a precocial bird.
2. We manipulated clutch size by exchanging complete clutches between pairs of nests to increase or decrease the clutch size by zero (control), one, two or three eggs in 314 nests over 2 years.
3. Pre-fledging survival of goslings increased in enlarged broods and decreased in reduced broods compared to control. Consequently, enlarged broods fledged more offspring and the reverse was true for reduced broods.
4. Size and mass of goslings near fledging was also higher in enlarged broods than in control, which suggests that offspring quality was also enhanced by the manipulation. This is contrary to the common trade-off between offspring numbers and quality.
5. Large families were dominant over smaller ones in feeding sites, which could explain the increased survival and growth of enlarged broods.
6. Our results suggest that the ability to raise young does not limit clutch size in this species and that parents could be more successful (i.e. increase both the number and quality of their offspring) by laying more eggs. However, the time required to lay additional eggs reduces the viability of all offspring and may explain why females do not lay more eggs.     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

Reproductive performance of a lacertid lizard at the core and the periphery of the species' range

The range boundaries of organisms are frequently interpreted in terms of a decline in the extent to which the life histories of outer populations are able to adapt to local environmental conditions. To test this hypothesis, we compared the reproductive characteristics of two Iberian populations of the lizard Psammodromus algirus (Linnaeus, 1758). One of them (Lerma) is close to the northern edge of the species' range, whereas the other one (El Pardo) occupies a typical core habitat 200 km further south. Gravid females were captured in the field and transported to the lab for egg laying. Second clutches were less frequent at Lerma (where clutch size and clutch mass were larger for first than for second clutches) than at El Pardo. The total mass of both clutches combined was similar at both sites. Thus, the higher frequency of second clutches at El Pardo appeared to balance the between-sites difference in energy allocation to the first clutch. Females from Lerma laid more but smaller eggs than those from El Pardo. When incubated at the same temperature, eggs from Lerma hatched sooner even when controlling for between-sites differences in mean egg size. These differences are interpreted in the light of the advantages of early hatching and high fecundity in the northern population, as opposed to large offspring size in the core population. We conclude that the life-history traits studied show enough variation, presumably of an adaptive nature, to cope with environmental challenges at the edge of the species' range.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 87–96.