The function of hatching asynchrony in the blue-footed booby

The blue-footed booby (Sula nebouxii) commonly hatches two eggs 4 days apart; then the senior (first-hatched) chick aggressively dominates the other and sometimes kills it. Two hypotheses explaining the function of the hatching interval were tested by creating broods with synchronous hatching: the facultative brood reduction hypothesis of Lack (1954) and the sibling rivalry reduction hypothesis of Hahn (1981). The results contradicted most predictions of both hypotheses: synchronous broods formed an aggressive hierarchy similarly to asynchronous broods (controls), and subordinate chicks grew poorly (Fig. 1) and died frequently, similarly to junior chicks in control broods. However, compared with synchronous broods, asynchronous broods showed less aggression (Fig. 2), diminished food allocation to subordinate chicks (Fig. 3) and less total food consumption (30% fewer feeds at age 0–10 days). These behavioral comparisons took into account the different ages of chicks in different treatments. The results suggest that natural asynchrony makes brood reduction more efficient and decreases the costs of sibling aggression to parents, in terms of their future survival or fecundity, as proposed by Mock and Ploger (1987). Further, in exaggeratedly asynchronous broods (8-day hatch interval) junior chicks suffered more aggression (Fig. 4) and grew more slowly than junior chicks in control broods. This result supports the hypothesis of optimal hatch asynchrony of Mock and Ploger (1987).     

Evidence of kin-selected tolerance by nestlings in a siblicidal bird

Behaviorally dominant members of blue-footed booby (Sula nebouxii) broods can effect siblicide by restricting access of subordinate siblings to parents providing food. In spite of their capacity for siblicide, dominant chicks permit subordinates to feed during short-term food shortage; in fact, the proportion of the food that the dominant takes is independent of the total amount delivered in older chicks. A model of optimal food distribution suggests that dominant chicks maximize their inclusive fitness with this pattern, rather than by satisfying their own food requirements and leaving what remains for the subordinate sibling. The indirect reproductive potential represented by a chick's sibling appears to have influenced the evolution of siblicidal brood reduction in this species.     

Food-dependent benefits of hatching asynchrony in American kestrels Falco sparverius

Food supply and hatching asynchrony were manipulated for 90 broods of American kestrels (Falco sparverius) during 1989–1991. We measured the growth and mortality of nestlings within four treatment groups (asynchronous, synchronous, food-supplemented, unsupplemented) to test the brood reduction hypothesis of Lack (1947, 1954). Fledging success did not differ between synchronous and asynchronous broods when food was poor but consistent with the brood reduction hypothesis, nestlings died at a younger age in asynchronous broods. When food was supplemented, mortality did not occur in the synchronous broods but youngest nestlings still died in asynchronous nests despite apparently adequate food for the brood. Oldest nestlings in asynchronous broods fledged with a greater mass than their younger siblings, also consistent with Lack's hypothesis. Average nestling quality in synchronous broods was very dependent on food levels. Synchronous young that were supplemented were, on average, the heaviest of any treatment group but young from unsupplemented synchronous broods were the lightest. Overall, patterns of mortality and growth for kestrels support the brood reduction hypothesis when food is limited, but not when it is abundant. This food-dependent benefit of asynchrony in the nestling period is a prerequisite for facultatively adjusted hatching spans during laying.     

Opposing selective pressures on hatching asynchrony: egg viability, brood reduction, and nestling growth

At least 19 hypotheses have been proposed to explain the evolutionary significance of avian hatching asynchrony, and hatching patterns have been suggested to be the result of several simultaneous selective pressures. Hatching asynchrony was experimentally modified in the black kite Milvus migrans by manipulating the onset of incubation during the laying period. Delayed onset of incubation reduced egg viability of first-laid eggs, especially when ambient temperature during the laying period was high. Brood reduction (nestling mortality by starvation or siblicide) was more commonly observed in asynchronous nests. The growth rate was slower in synchronous broods, probably due to stronger sibling rivalry in broods with high size symmetry. Last-hatched chicks in synchronous broods fledged at a small size/mass, while in control broods, hatching order affected growth rates, but not final size. Brood reduction, variable growth rates, and the ability to face long periods of food scarcity are probably mechanisms to adjust productivity to stochastic food availability in a highly opportunistic predator. The natural pattern of hatching asynchrony may be the consequence of opposing selective forces. Extreme hatching synchrony is associated with slow growth rates, small final size of last-hatched chicks, and low viability of first-laid eggs, while extreme hatching asynchrony is associated with high mortality rates. Females seem to facultatively manipulate the degree of hatching asynchrony according to those pressures, because hatching asynchrony of control clutches was positively correlated with temperature during laying, and negatively correlated with the rate of rabbit consumption. Received: 25 October 1999 / Revised: 30 May 2000 / Accepted: 25 June 2000     

Sibling aggression, hatching asynchrony, and nestling mortality in the black kite (Milvus migrans)

In siblicidal species, hatching asynchrony could act to reduce sibling rivalry or promote the death of last-hatched chicks. The pattern of hatching asynchrony was experimentally altered in the black kite Milvus migrans. Hatching asynchrony in control broods was intermediate between those of experimentally synchronised and asynchronised broods. Sibling aggression and wounds on the chicks were more commonly observed early in the nestling period and in synchronous nests. Serious injuries were observed on last-hatched chicks in asynchronous nests, as were observations of intimidated or crushed chicks. Sibling aggression was related to food abundance, but some chicks died at an early age in nests with abundant food (cainism). Cainism was more commonly found in asynchronous nests. For species with facultative siblicide, moderate hatching asynchrony could be a compromise between reducing sibling rivalry and avoiding large size differences between sibs that would result in cainism. Female black kites preferentially fed the smallest chicks and exhibited behaviours to reduce sibling aggression, contrary to observations in other siblicidal species. In a highly opportunistic forager such as the black kite, a strategy may exist to protract the life of all the chicks in the brood, waiting for unpredictable situations of food overabundance. This would induce the appearance of a parent-offspring conflict over brood reduction, reflected in the existence of a possible anticipated response by some of the chicks (cainism) and in the appearance of special behaviours by the parents to selectively feed smaller chicks or reduce sibling aggression. In this facultatively siblicidal species, cainism does not seem to be the final stage of an evolutionary trend favouring the raising of high-quality chicks, but a manifestation of a parent-offspring conflict over brood size. Received: 9 March 1998 / Accepted after revision: 8 August 1998     

Is asynchronous hatching adaptive in herring gulls (Larus argentatus)?

Hatching asynchrony commonly induces a size hierarchy among siblings and the resultant competition for food between siblings can often lead to starvation of the smallest chicks within a brood. We created herring gull (Larus argentatus) broods with varying degrees of hatching synchrony by manipulating the timing of incubation while maintaining the originally laid eggs. The degree of hatching asynchrony affected sibling size hierarchy at the time of hatching of the last-hatched ”c-chick.” In unmanipulated broods, there was no disadvantage of being a c-chick. However, when asynchrony was experimentally increased, we found reduced survival of the c-chick only in the exaggerated asynchronous experimental group. The effects were observable only during the first 10 days of chick life. We recorded no cases of the chicks dying of starvation. Furthermore, behavioral observations indicated that there was no sibling competition, and no selective feeding of larger sibs in the study colony. We propose that the observed lower survival rates of c-chicks in exaggerated asynchronous broods resulted from their lesser motor abilities, affecting their chances of escaping predators. Fledging success for the whole colony was generally high and almost half of all pairs fledged all three chicks, which is indicative of a good feeding environment. We argue that normal hatching asynchrony is a favorable solution in a good feeding environment, but that increased asynchrony reduces breeding success. We do not view asynchrony in the herring gull as an adaptation for brood reduction and propose instead that it may come about because there has been selection for incubation to start before clutch completion. Received: 14 April 1999 / Received in revised form: 20 October 1999 / Accepted: 23 January 2000     

Begging and provisioning in broods of asynchronously-hatched yellow-headed blackbird nestlings

Studies of begging have found a positive relationship between begging level and provisioning level. Studies of unequal nestlings, however, have found that small nestlings generally beg more but are fed less than their larger siblings. We manipulated the begging levels of yellow-headed blackbird (Xanthocephalus xanthocephalus) chicks to investigate how begging benefits individuals in broods of unequal siblings. Food-deprived chicks begged more and were fed more; satiated chicks begged less and were fed less. When we deprived each chick of a brood in turn, large and small chicks generally increased begging and received more provisioning. Small chicks, however, rarely received more food than their larger siblings even when they behged relatively more. Parent yellow-headed blackbirds increase provisioning to hungry begging chicks, but also allocate food based on relative offspring size.     

Parent blue-footed boobies suppress siblicidal behavior of offspring

Behaviorally dominant nestlings routinely kill sibling nestmates in blue-footed booby (Sula nebouxii) broods during periods of food shortage. Previous work demonstrated that these dominant, first-hatching “A-chicks” regulate the lethality of their behavior towards subordinate, second-hatching “B-chicks,” showing tolerance towards B-chicks except during chronic food shortages. Siblicide by A-chicks usually occurs after the hatchling stage. Results of an interspecific cross-fostering experiment indicated that A-chicks also attempt siblicide shortly after hatching, but parents apparently exert control over these attempts, and thwart them, when chicks are young. Theory predicts selection for such regulation in siblicidal birds that are likely to experience genetic parent-offspring conflict over the value of subordinant nestlings; our evidence of post-hatching parental regulation is consistent with that prediction. Received: 6 June 1998 / Accepted after revision: 12 July 1998     

Do males and females differ in the feeding of large and small siblings? An experiment with the bluethroat

Males and females have been reported to differ in their feeding of large and small siblings in several species of birds. According to recent hypotheses, this phenomenon may be related to a sexual conflict over avian hatching patterns. We designed an experiment to test for the existence of such a sex difference by manipulating nestling size hierarchies of the bluethroat (Luscinia s. svecica) in two directions; half the broods were “asynchronized” to yield large size-differences within broods and the other half were “synchronized” to yield small size-differences. In all broods, nestlings were categorized as being either large or small according to body mass. We recorded male and female food distribution by video early (day 4 after hatching) and late (day 8) in the nestling period. Males and females did not differ in their distribution of food among different-sized nestlings. With large size-differences, both males and females fed large nestlings nearly twice as often as small ones. In contrast, when the size-differences were small, food was more evenly distributed among nestlings. Early in the nestling period, males fed more nestlings during each feeding visit than did females. Our finding that male and female bluethroats do not differ in the feeding of large and small siblings is in contrast to most previous studies. Variation in costs and benefits to males and females from feeding different-sized nestlings, and restrictions to parental choice due to nestling interactions, may explain interspecific variation. Received: 27 June 1997 / Accepted after revision: 26 January 1998     

Influences of sex,sex composition of brood and hatching order on mass growth in common terns<Emphasis Type="Italic"> Sterna hirundo</Emphasis>

We studied sex-dependent mass growth of chicks of the monomorphic common tern Sterna hirundo which fledged at a colony site in Wilhelmshaven, Germany, between 1997 and 1999. Brood size, brood mates' sex and hatching order (a-, b- and c-chicks) were known for many individuals, which were sexed by molecular techniques. Daily growth rates and age of fledging were independent of sex. However, in all years male chicks reached higher peak mass and fledged up to 5.2% heavier than female chicks. Broods with at least two fledglings showed that besides sex, brood size interacting with sex composition affected chick mass. In mixed broods, brothers had higher peak and pre-fledging mass than sisters they were reared with. Especially in the combination a-daughter and b-son the brothers were heavier. Lowest mass was found in broods with three nest mates of the same sex. A detailed study of 24 three-fledgling broods showed that male c-chicks were heavier than their siblings. The results reveal an advantage for chicks in mixed broods, especially for sons, and more especially if the son was a c-chick. Higher mass and possibly dominance of sons in c-position might be related to higher maternal androgen levels, which are known to increase with each egg laid. The results suggest that even in a monomorphic species, sons might be more expensive to rear, and are discussed with respect to sibling competition, parental effort, survival of sons, as well as to fitness benefits favoring parents producing sons.     

Brown pelican siblicide and the prey-size hypothesis

Summary We asked whether the brown pelican (Pelecanus occidentalis) practices facultative brood reduction and tested two predictions of Mock's (1985) prey-size hypothesis: (1) if chicks take food directly from the parental mouth, nestmates should compete aggressively; (2) aggression between nestmates should increase during the developmental transition from indirect feeding (parents deposit food on the substrate) to direct feeding (parents pass food from mouth to mouth). Eggs in two-egg and three-egg clutches were laid and hatched 2 days apart. Junior (second- and third-hatched) chicks, fed less than their nestmates, grew more slowly and died more frequently. Of non-hatchling victims 74% were underweight, 55% bore peck wounds and 43% were found outside the nest, implying starvation, aggression and expulsion from the nest. Aggression occurred in all observed sibships, and junior chicks were subordinate in eight of ten cases. A progressive transition from indirect to direct feeding occurred during the 30 days after hatching, but there was no evidence of an increase in pecking during the transition. In conclusion, the brown pelican practices facultative brood reduction, and chick loss is mediated by sibling aggression as predicted by the preysize hypothesis. Pecking during indirect feeds seems to contradict the prey-size hypothesis, but may serve partly to establish a relationship of dominance-subordinance.Correspondence to: H. Drummond     

Effects of territory quality, food availability and sibling competition on the fledging success of oystercatchers (Haematopus ostralegus)

We investigated the fledging probability of oystercatcher, Haematopus ostralegus, chicks as a function of hatching order, brood size, territory quality and food availability. Sibling dominance was related to the hatching order in both low- (’leapfrogs’) and high-quality (’residents’) territories. Differences in hatchling mass might have aided the establishment of a dominance hierarchy, since breeders produced small late eggs and hatchlings. These mass differences were most pronounced in leapfrogs, and in large broods in years with lower food availability (’poor’ years). Late hatchlings fledged less often and with lower body masses compared to early hatchlings in all situations. Leapfrogs produced smaller broods and hatched their broods more asynchronously in poor years than leapfrogs breeding in years with more available food (’good’ years) and residents breeding in both poor and good years. Large brood sizes resulted in lower survival of hatchlings in poor years. These results favour the ’brood reduction’ hypothesis. However, contrary to the expectations of this hypothesis, hatching order also affected fledging success in residents. Moreover, large brood size resulted in higher survival of hatchlings in good years, particularly in residents. Thus, although large broods experienced losses due to sibling competition in some years, they nevertheless consistently produced more fledglings per brood in all years, both as leapfrogs and residents. We believe this effect is due to parental quality correlating with initial brood size. Most leapfrogs, at best, fledged one chick successfully each year, losing chicks due to starvation. Nevertheless, leapfrog broods were reduced in size after hatching significantly less quickly than resident broods. These results suggest that breeders lay and hatch insurance eggs to compensate for unpredictable losses due to the high predation rates on both nests (ca 50%) and chicks (ca 90%), in accordance with the ’nest failure’ hypothesis. Received: 14 February 2000 / Revised: 27 September 2000 / Accepted: 10 June 2000     

Social hierarchy among siblings in broods of the oystercatcher Haematopus ostralegus

Summary On Skokholm Island, Wales, the young of the Oystercatcher (Haematopus ostralegus) are fully precocial, yet are totally dependent on parents for food. Though all young (commonly three) usually hatch within one day, small hatching intervals do exist, and first-hatched nestlings are usually slightly heavier at hatching than those hatched later. Firsthatched young and/or initially heavier young become socially dominant over their siblings; a stable, linear, non-aggressive social hierarchy among siblings develops, which determines the partition of food among them and results in unequal distribution when food is scarce. Food shortage impairs the growth rate of subordinate young. They are rather hungry and restless and consequently predation on them is higher than on dominant young.It is proposed that Oystercatchers achieve effective brood-reduction by the parents' control of differential egg weight, by the eggs' hatching order, and by the young's acceptance of a non-aggressive social hierarchy. The latter is an adaptation associated with the unusual phenomenon of fully precocial wader young being exclusively fed by parents.     

Brood reduction in the American white pelican (Pelecanus erythrohynchos)

Summary American white pelicans (Pelecanus erythrorhynchos) breeding in colonies at East Shoal Lake, Manitoba, Canada exhibited a mean hatching asynchrony of 2.5 days in 2-egg clutches. This resulted in a size difference between chicks which facilitated sibling dominance, harassment and lack of food for the subordinate chick. Only one young survived per nest. In marked broods, the secondhatched chick survived in 20% of successful nests. Manipulated clutch sizes (1, 2 and 3 eggs or chicks per nest) revealed that the presence of a second chick contributes significantly to the reproductive success of the parents. Results support the hypothesis that the second egg functions as a form of insurance against early loss of the first egg or chick. The parents, by establishing hatching asynchrony, by nonintervention in sibling aggression, and by selectively feeding the dominant chick, maximize their chance of rearing the most viable young.     

Sex-specific effects of albumen removal and nest environment manipulation on Barn Swallow nestlings

In avian species, maternal provisioning to the eggs is predicted to be more valuable for the offspring under adverse environmental conditions and intense sibling competition. However, studies manipulating both the amount of maternal pre-hatching resources and the harshness of post-hatching environment have seldom been performed to date. In this experimental study of Barn Swallow (Hirundo rustica) nestlings, we tested the consequences of a reduction in the albumen content of the eggs for fitness-related offspring traits, while performing an unbalanced partial cross-fostering soon after hatching, either increasing or decreasing brood size by one nestling. By molecular sexing of the chicks, we additionally tested for sex-specific sensitivity of individual nestlings to experimental treatments and to sex ratio variation in nestmates. We predicted that chicks hatching from albumen-deprived eggs should suffer more than control chicks from the harsher rearing conditions of enlarged broods. However, although albumen removal depressed chick body mass, chicks hatching from control eggs did not fare better than those hatching from eggs with reduced albumen content in enlarged vs. reduced broods. Albumen removal had sex-specific effects on immunity, with males, but not females, hatching from eggs with reduced albumen content showing a lower T-cell-mediated immune response than controls, suggesting that the two sexes were differentially susceptible to resource deprivation during early ontogeny. In addition, both immune response and chick body mass at age 7 days, when maximum growth rate is attained, declined with an increasing proportion of male nestmates. The effect of brood size manipulation on chick body mass at age 12 days, when peak body mass is attained, was also found to depend on brood sex composition, in that an increase in the proportion of male nestmates depressed offspring body mass in reduced broods, while the reverse was true in enlarged broods. On the whole, these findings suggest that sex differences may exist in environmental sensitivity and patterns of resource allocation among different body functions, and that brood size variation and sex composition may affect offspring fitness-related traits.     

Benefits of large broods by higher chick survival and better territories in a precocial shorebird

When reproductive success is constant in one breeding phase, different tactics that increase variation in reproductive success among individuals may evolve in other phases. For instance, in shorebirds, which usually have a limited clutch size of four eggs, variation in reproductive tactics among individuals is expected either before egg-laying (e.g. diverse mating systems) or after hatching of the young (e.g. diverse parental care). In this paper, I studied the pied avocet (Recurvirostra avosetta), a shorebird with a modal clutch size of four eggs, to test whether post-hatch chick adoption as an alternative tactic can be linked to increased variation in annual reproductive success. When predation was high, naturally adopting pairs produced more filial fledglings than did pairs not adopting chicks and not losing chicks to adoption. The number of filial fledglings increased with the number of adopted young, possibly through diluting the chances of predation on filial young. Experimental chick addition did not lead to more fledged young due to low brood integrity as shown by the frequent loss of chicks from some experimental broods. When predation was low, larger broods occupied feeding territories with higher prey abundance than smaller broods, possibly due to their dominance over smaller ones. Pairs that lost chicks to adoption (donors) fledged as many filial young in their broods as did non-adopters/non-donors, whereas the total number of donors’ filial fledglings, including those raised in adopting broods, approached that of adopters. These findings show, for the first time, that post-hatch alternative reproductive tactics can lead to variation in annual reproductive success and to higher success for some pairs even in species where past adaptations limit variation in reproductive success in a certain phase of reproduction.     

Proximate control of siblicide in cattle egrets: a test of the food-amount hypothesis

Avian sibling aggression plays an important role in controlling food distribution among nest mates. The food-amount hypothesis predicts that the amount of food obtained by siblings acts as a proximate cue controlling their level of aggression. The importance of this cue to cattle egrets (Bubulcus ibis) in Oklahoma was evaluated by studying parental feeding rates, food distribution among nestlings, and nestling aggression in three-chick broods [A-(oldest), B-, and C-chicks]. We compared nests in which siblicide occurred to nests where all three chicks survived (nonsiblicide nests). Relative to nonsiblicide nests, siblicide nests received decreasing amounts of food as the day of siblicide approached. Similarly, aggression levels of all three chicks in siblicide nests increased significantly compared with nonsiblicide nests, with most of the aggressive behavior involving B- and C-chicks. As a result, on a per-day basis, food amount and aggression exhibited a strong, negative correlation, thus providing support for the food-amount hypothesis. Though the total amount of food delivered to siblicide nests declined on the days prior to siblicide, the amount of food received by A-chicks in these nests did not. Relative to nonsiblicide nests, A-chicks in siblicide nests received an increased share (percent of volume) of the food delivered at the expense of C-chicks, with the percent remaining the same for B-chicks. The percentage of food B-chicks received the day after siblicide increased significantly from that received the day before siblicide, with no change occurring in either the amount or percentage of food received by A-chicks; clearly, B-chicks were the beneficiary of their aggressive actions that resulted in the death of C-chicks. We provide evidence supporting the hypothesis that aggression is proximately linked to food amount, possibly by a relatively simple cue such as hunger. Received: 25 November 1994/Accepted after revision: 10 February 1996     

Maternal effects on begging behaviour: an experimental demonstration of the effects of laying sequence, hatch order, nestling sex and brood size

Differential resource allocation by females across the laying sequence has been hypothesised as a mechanism through which females could either compensate nestlings that hatch last in asynchronous broods or promote brood reduction. In this study we artificially incubated eggs and cross-fostered offspring to manipulate nestlings’ position in the hatching order, to identify whether the competitive ability of nestlings is dependent on position in the laying sequence. In both control and experimentally reversed broods, first hatched chicks had a higher survival than last hatched siblings. Yet, nestlings that hatched from eggs laid in the second half of a clutch begged with a greater intensity than nestlings hatched from eggs laid in the first half of a clutch. In natural broods, the greater begging competitiveness of nestlings from later-laid eggs led to a moderation of sibling competition and these nestlings achieved the same body size and weight as nestlings from eggs laid in the first half of the clutch. The lack of a substantial difference in the size and condition of surviving nestlings in respect to laying order suggests that differential resource allocation across the egg-laying sequence partially compensates for hatching last in asynchronous broods and reduces the negative effects of the nestling size hierarchy. The effect of laying order, brood size and experimental treatment also differed for male and female nestlings. Our study highlights the need to be aware of the complex and subtle effects of nestling sex and laying sequence when investigating genetic and environmental influences on individual fitness.     

Is natural hatching asynchrony optimal? An experimental investigation of sibling competition patterns in a facultatively siblicidal seabird

In unpredictable environments, any tactic that enables avian parents to adjust brood size and, thus, energy expenditure to environmental conditions should be favoured. Hatching asynchrony (HA), which occurs whenever incubation commences before clutch completion, may comprise such a tactic. For instance, the sibling rivalry hypothesis states that the hierarchy among chicks, concomitant to HA, should both facilitate the adjustment of brood size to environmental conditions and reduce several components of sibling competition as compared to synchronous hatching, at both brood and individual levels. We thus predicted that brood aggression, begging and feeding rates should decrease and that older chick superiority should increase with HA increasing, leading to higher growth and survival rates. Accordingly, we investigated the effects of an experimental upward and downward manipulation of HA magnitude on behaviour, growth and survival of black-legged kittiwake (Rissa tridactyla) chicks. In line with the sibling rivalry hypothesis, synchronous hatching increased aggression and tended to increase feeding rates by parents at the brood level. Begging rates, however, increased with HA contrary to our expectations. At the individual level, as HA magnitude increased, the younger chick was attacked and begged proportionally more often, experienced a slower growth and a higher mortality than its sibling. Overall, the occurrence of energetic costs triggered by synchronous hatching both for parents and chicks, together with the lower growth rate and increased mortality of the younger chick in highly asynchronous broods suggest that natural HA magnitude may be optimal.     

Costs and benefits of surplus offspring in the lesser kestrel (Falco naumanni )

Lesser kestrels (Falco naumanni) lay clutches which appear excessive as only 3% of them yield as many young as eggs laid. Four hypotheses may explain the adaptive value of producing surplus eggs: (1) the bet-hedging hypothesis assumes that the environment varies unpredictably and surplus eggs serve to track uncertain resources; (2) the ice-box hypothesis suggests that surplus offspring serve as a reserve food during a period of shortage; (3) the progeny choice hypothesis says that parents produce surplus offspring in order to choose these with higher fitness; and (4) the insurance-egg hypothesis proposes that extra eggs are an insurance against the failure of any egg. To test the significance of this strategy in the lesser kestrel, an experiment manipu-lating brood size at hatching was carried out over 2 years, with good and bad feeding conditions. The experiment consisted of adding a chick to experimental broods where one egg failed to hatch or removing a randomly selected chick from experimental broods where all eggs had hatched. Independently of annual food availability, pairs with brood sizes reduced by one chick fledged more nestlings than pairs with brood size equalling their clutch sizes. Body condition of young was also better in the former group, but only in 1993 (a high-food year). Independently of year, mean local survival of parents with complete broods at hatching was lower than for parents raising reduced broods. These results supported only the insurance-egg hypothesis which says that surplus eggs may be an insurance against the failure of any egg, but parents may suffer reproductive costs when all eggs hatch. Received: 17 January 1997 / Accepted after revision: 27 April 1997