Is there a cost of reproduction for Marsh Tits Parus palustris in a primeval forest?

We looked for evidence of a cost of reproduction in the Marsh Tit Parus palustris living in the last fragments of primeval temperate forest (Białowieża National Park, eastern Poland). Potential nest-holes were superabundant but the birds had to cope with a diverse set of predators, dangerous both to broods and to parents. Taking advantage of the natural variation in realized reproductive investment that this caused in terms of the loss of nests or mates, we expected to find differences in survival and future fecundity between birds which had lost broods (reduced effort), had reared young (controls) or were either provisioning young single-handed or had laid replacement clutches (increased effort). Despite 13 years of observation, even during seasons with very strenuous conditions, we have failed to demonstrate that the observed range of variation in parental investment caused any demographic cost of reproduction. Incubating females were regularly killed on the nest, which could indicate the existence of a cost operating in the earlier stages of the breeding cycle. Overall, these results suggest that the reproductive rate in Marsh Tits is not controlled proximately by reproductive cost.     

Serum immunoglobulin G levels are positively related to reproductive performance in a long-lived seabird, the common tern (Sterna hirundo)

The evolution of longevity requires a low risk of mortality from extrinsic factors, relative to intrinsic factors, so that individuals that differentially invest in physiological self-maintenance and minimize their annual reproductive costs will maximize lifetime fitness through a prolonged reproductive lifespan. The trade-off between reproductive effort and self-maintenance, as measured by immune function, has been well documented in short-lived birds, but is difficult to demonstrate in long-lived birds. To assess self-maintenance in a long-lived seabird, we measured serum protein levels, including immunoglobulin G (IgG = IgY), in 30 breeding pairs of common terns (Sterna hirundo) and their first-hatched (A) chicks. Most parents were of known age from banding as hatchlings; our sample was selected to contrast young breeders (6–9 years) with very old birds (17–23 years). Body-mass of the parents declined by 5% during the chick-rearing period, while serum protein levels were stable. Serum IgG levels were higher in parents of offspring with faster growth rates, while IgG levels were lower in parents whose broods were reduced by starvation. A-chicks in broods of two had higher IgG levels than singleton chicks. Albumin levels were not related to reproductive performance. Thus, despite adequate statistical power, we could find no evidence for a trade-off between reproduction and self-maintenance in common terns, even in old age. The results are consistent with life-history predictions for long-lived vertebrates, in which selection favors sustained self-maintenance across the reproductive lifespan. The positive relationships between IgG levels and reproductive performance indicate that IgG can be used as an index of parental “quality.”     

Age-dependent reproductive costs and the role of breeding skills in the Collared flycatcher

This study addressed whether there are any age‐related differences in reproductive costs. Of especial interest was whether young individuals increased their reproductive effort, and thereby their reproductive cost, as much as older birds when brood size was enlarged. To address these questions, a brood‐size manipulation experiment with reciprocal cross‐fostering of nestlings of young and middle‐aged female Collared flycatchers, Ficedula albicollis, was performed on the Swedish island of Gotland. Nestlings’ body mass, tarsus length and survival were recorded to estimate the parental ability and parental effort of the experimental female birds. Female survival and clutch size were recorded in the following years to estimate reproductive costs. We found that middle‐aged female flycatchers coped better with enlarged broods than younger females or invested more in reproduction. In the following year, young female birds that had raised enlarged broods laid smaller clutches than the females from all the other experimental groups. This result shows that the young female birds pay higher reproductive costs than the middle‐aged females. Both young and middle‐aged female flycatchers seemed to increase their reproductive effort when brood size was increased. However, such an increase resulted in higher reproductive costs for the young females. The difference in reproductive costs between birds of different ages is most likely a result of insufficient breeding skills of the young individuals.     

Brood size and immunity costs in zebra finches Taeniopygia guttata

Birds rearing experimentally enlarged broods have lower antibody responses to a novel antigen, and we tested three hypotheses that could explain this result. We used zebra finches Taeniopygia guttata inoculated with sheep red blood cells (SRBC) as a study system, for which this trade-off was previously demonstrated. 1. Compensatory cellular immunity: The humoral immune response is slow, and removal of SRBC through up-regulated cellular immunity could pre-empt an antibody response. However, cellular immune response to PHA decreased with increasing brood size, allowing rejection of this hypothesis. 2. Costs of antibody-production: Chicks in large broods grow less well, and birds with large broods may allocate resources to chicks instead of antibodies when these are costly. Compared to saline controls, SRBC suppressed metabolic rate in the hours following immunisation, but there was no effect in the following night, or at any time 4 and 8 days later. Fitness costs were measured by repeatedly immunising parents with SRBC while rearing young. Chick growth, parental condition, and subsequent reproduction of the parents were not affected by SRBC. We conclude that the costs of antibody formation cannot explain the trade-off between brood size and antibody responsiveness. 3. Costs of immune system maintenance: Maintaining a system enabling antibody-formation may be very costly, and birds rearing large broods may have down-regulated this system. Based on this hypothesis we predicted that antibody formation would still be reduced in parents rearing large broods when immunised after rearing the chicks. Our results confirmed this prediction, and we suggest that birds rearing large broods have lower antibody responses because they economised on the maintenance costs of the immune system.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.     

Food provisioning lowers inter-clutch interval in Moorhens Gallinula chloropus

Food supplements were provided to Moorhens Gallinula chloropus during production of their first brood, to test whether food availability limited the number of broods they could rear in a season. Supplemented territories had shorter inter-clutch intervals when individual differences were held constant, although they did not produce more broods. Because unmanipulated birds with short inter-clutch intervals produced more broods, food availability may be one of several factors limiting the production of multiple broods and therefore reproductive success.     

Experimentally increased reproductive effort alters telomere length in the blue tit (Cyanistes caeruleus)

Telomeres have recently been suggested to play important role in ageing and are considered to be a reliable ageing biomarkers. The life history theory predicts that costs of reproduction should be expressed in terms of accelerated senescence, and some empirical studies do confirm such presumption. Thus, a link between reproductive effort and telomere dynamics should be anticipated. Recent studies have indeed demonstrated that reproduction may trigger telomere loss, but actual impact of reproductive effort has not received adequate attention in experimental studies. Here, we experimentally manipulated reproductive effort by increasing the brood size in the wild blue tit (Cyanistes caeruleus). We show that parents attending enlarged broods experienced larger yearly telomere decay in comparison to control birds attending unaltered broods. In addition, we demonstrate that the change in telomere length differs between sexes, but this effect was independent from our treatment. To our knowledge, this is the first experimental study in the wild revealing that telomere dynamics may be linked to reproductive effort. Thus, telomere shortening may constitute one of the potential proximate mechanisms mediating the costs of reproduction.     

GROWTH RATES OF BIRDS IN THE HUMID NEW WORLD TROPICS

The growth curves of 40 species of lowland neotropical birds were fitted by logistic equation. The birds were mostly from Panama, Trinidad and Surinam. The growth constants of the fitted equations (asymptote A and growth rate K) were compared within and among species, and with previously published data on temperate species. Growth parameters of tropical passerines are about as variable within species as they are within temperate species. In both cases, variation in A and K between broods is greater than it is within broods. Panamanian birds breed during the dry-wet transition and conditions for growth apparently improve as rainfall increases. Asymptotes of growth curves are higher, and mortality within broods lower, as the breeding season progresses. Asynchronous hatching and the reduction of brood-size by selective starvation of young is a prominent phenomenon during the early part of the breeding season. Several instances are reported, however, of young persisting in nests with inadequate feeding and greatly subnormal weights. Slowed development under conditions of poor nutrition may be adaptive in the tropics if periods of low food availability are short and allow the possibility of recovery from undernourishment. As a group, neotropical lowland passerines (30 species) grow 23% more slowly than a sample of 51 temperate passerines. Variation of growth rates among these tropical species is similar to variation among temperate species, and it is related to adult body-size the length of the nestling period. Young of tropical and temperate species attain similar asymptotes, relative to adult body-weight, by the end of the nestling period. Hypotheses are advanced which might explain the slower growth rate of tropical species, and tested to the extent available data permits. (1) Because brood-size can be changed only by adding or removing whole young, changes in growth rate could provide finer adjustment of the energy requirements of the young to the feeding capacity of the parents. This model predicts different means and variances for growth rate within groups of species with different clutch-sizes, predictions not supported by available data. (2) Growth rate is shown to increase the maximum energy requirement of a nestling only if K exceeds some value determined by the energy requirement of the young, growth rate should vary in proportion to the level of basal maintenance metabolism. In a small sample of tropical species, rates of basal metabolism were 25% lower than in a comparable sample of temperate species. These data therefore support the hypothesis, although the cause of the lower metabolic rate of the tropical nestlings is not known. (3) Daily periods of hypothermia could reduce the energy requirement of the young and at the same time reduce their growth rate; but observations of body temperatures of tropical nestlings are contrary to this hypothesis. (4) The short day-length of tropical climates reduces the time during which young can assimilate energy relative to their energy expenditures. This model predicts that tropical nestlings would have less productive energy available, (consistent with their reduced growth rates), but it also predicts that arctic birds should grow faster than temperate species, which is not confirmed by available data. (5) The low nitrogen content of fruits may cause the slow growth of a few strictly frugivorous species (Oilbird and Bearded Bellbird), but among other tropical species growth rate is not correlated with the estimated proportion of fruit in the diet.     

Broodcare helpers in the cichlid fish Lamprologus brichardi: Their costs and benefits

‘Helping’ in birds and mammals involves seemingly altruistic behaviour. In the cichlid fish Lamprologus birchardi helpers are usually young of former broods staying in their parents' territories and participating in all kinds of parental duties (Broodcare, territory maintenance and defence). The discovery of helpers in fish offered the chance of attempting an extensive analysis of potential costs and benefits influencing the evolution of helpers in a vertebrate. Three factors proved to be of major importance in the cost-benefit analysis of helping as opposed to leaving for family-independent nonreproductive aggregations. Due to investment and to their rank within a family's hierarchy, helpers grow at a slower rate than non-helpers. This cost is compensated for by (i) a lower mortality risk to helpers caused by their access to a defended shelter and by protection afforded by bigger family members, and (ii) a positive contribution by helpers to the future reproductive success of their parents: females with helpers produce bigger clutches and consequently more free-swimming fry (=siblings). Other variables, such as the helpers' influence on the relative breeding success of their parents, broodcare experience through helping, the chances of territory take-over, parasitism of parents' reproduction and cannibalism are of minor importance. Similar social organizations in other fish are discussed with respect to their ecology and are compared with cooperatively breeding birds and mammals.     

Unpredictable food supply modifies costs of reproduction and hampers individual optimization

Investment into the current reproductive attempt is thought to be at the expense of survival and/or future reproduction. Individuals are therefore expected to adjust their decisions to their physiological state and predictable aspects of environmental quality. The main predictions of the individual optimization hypothesis for bird clutch sizes are: (1) an increase in the number of recruits with an increasing number of eggs in natural broods, with no corresponding impairment of parental survival or future reproduction, and (2) a decrease in the fitness of parents in response to both negative and positive brood size manipulation, as a result of a low number of recruits, poor future reproduction of parents, or both. We analysed environmental influences on costs and optimization of reproduction on 6 years of natural and experimentally manipulated broods in a Central European population of the collared flycatcher. Based on dramatic differences in caterpillar availability, we classified breeding seasons as average and rich food years. The categorization was substantiated by the majority of present and future fitness components of adults and offspring. Neither observational nor experimental data supported the individual optimization hypothesis, in contrast to a Scandinavian population of the species. The quality of fledglings deteriorated, and the number of recruits did not increase with natural clutch size. Manipulation revealed significant costs of reproduction to female parents in terms of future reproductive potential. However, the influence of manipulation on recruitment was linear, with no significant polynomial effect. The number of recruits increased with manipulation in rich food years and tended to decrease in average years, so control broods did not recruit more young than manipulated broods in any of the year types. This indicates that females did not optimize their clutch size, and that they generally laid fewer eggs than optimal in rich food years. Mean yearly clutch size did not follow food availability, which suggests that females cannot predict food supply of the brood-rearing period at the beginning of the season. This lack of information on future food conditions seems to prevent them from accurately estimating their optimal clutch size for each season. Our results suggest that individual optimization may not be a general pattern even within a species, and alternative mechanisms are needed to explain clutch size variation.     

Cooperation, conflict, and crèching behavior in goldeneye ducks

Crèching behavior, or brood amalgamation, results in offspring being reared by adults other than their genetic parents. Although a variety of hypotheses have been proposed to explain this behavior, most assume either that brood amalgamation is accidental (i.e., nonselected) or that adoption of young is selected for because of social benefits to the young and/or adopting parents. We propose, instead, that brood amalgamation is a function of two separate processes: brood desertion and brood adoption. To examine brood desertion, we develop a graphic model to predict when parents should abandon their young and we test this model experimentally for the Barrow's goldeneye (Bucephala islandica). As predicted, females deserted their offspring when the size of the brood was experimentally reduced. Brood adoption occurred when deserted ducklings joined other broods. However, the success of ducklings in doing so was strongly dependent on the availability of potential host broods and on the age of the recipient broods. Foreign ducklings were readily accepted into young broods (<10 d old) but invariably were rejected from old broods. We could detect no benefits or costs of brood adoption to the host females, contrary to the expectations of a social benefit hypothesis. Our experiments indicate that Crèching behavior is driven by selection on adults to abandon their brood when the benefits of continued investment are outweighed by the reduction in future reproduction and selection on deserted ducklings to join other broods to obtain parental care. Rather than a form of cooperative brood care, Crèching in goldeneyes is perhaps best considered as a form of reproductive parasitism, entailing parent-offspring conflict over brood desertion and intergenerational conflict over adoption of abandoned young.     

Brood size and the economy of brood defence: examining Lack's hypothesis in a biparental cichlid fish

Synopsis We tested the explanatory value of two hypotheses reviewed by Lack (1954) in the maintenance of brood size in free-ranging convict cichlidsCichlasoma nigrofasciatum: (1) physiological constraints on egg production, and (2) behavioural constraints imposed by brood defence. Number of free-swimming young in 13 experimental (E) broods was augmented to the upper limit of the size distribution of natural broods (150 young); 18 control (C) broods were handled in the same way but brood size was not changed (mean ± SE = 69.5 ± 11.0). E and C brood sizes were measured at 5 day intervals. At day 20 (just before independence from parental care), 50.3 ± 9.4 (n = 9) young remained in E broods and 30.8 ± 7.8 (n = 8) young remained in C broods (p> 0.05). Offspring number did not differ significantly (p> 0.05) between C and E broods after day 10. Mean growth rate of offspring was significantly lower in E broods than in C broods, perhaps in response to increased density of young in the former. Both the convergence of offspring number in E and C broods and suppression of growth in E broods support a behavioural constraint; that during the first 10 days in which the young are free swimming, two parents are unable to defend large broods as successfully as small broods. A trade-off exists in parental investment between current and future reproduction. Extra-parental investment in current reproduction (eggs) does not result in an increased number of young at independence, therefore a behavioural constraint during brood defence should stabilize the evolution of clutch size.     

High blowfly parasitic loads affect breeding success in a Mediterranean population of blue tits

The detrimental effects of ectoparasites on the breeding success of birds have been especially well demonstrated in the case of ectoparasites that affect both chicks and their parents. Since blowfly larvae of the genus Protocalliphora attack only nestling birds, they represent a good model for testing the consequences of parasitism on nestlings. A Corsican population of blue tit suffers extremely high rates of infestation by blowflies, which are suspected to negatively affect young birds. Comparing experimentally deparasitized (treated) and naturally infested (control) broods, we showed that the attack by Protocalliphora causes anaemia and an important disturbance to the chicks. Therefore, we expected that these effects would have a negative impact on body condition and survival in the infested broods. Although we did not find any effect of treatment on fledging success, our predictions were confirmed by lower growth rate, body mass at fledging and tarsus length at fledging in the control compared with the treated group. This suggests that in this population, blowflies decrease the probability of recruitment of young blue tits. Received: 6 May 1997 / Accepted: 14 July 1997     

Paternal influence on growth and survival of dark-eyed junco young: do parental males benefit?

Recent studies of typically monogamous passerine birds have suggested that the fitness benefits males derive by caring for their young may not be as great as was previously thought. This study was conducted to determine whether parental care by male dark-eyed juncos, Junco hyemalis, serves to increase either the quantity or quality of young that they produce. Over a 4-year period, males were caught at the time their eggs hatched, and the subsequent growth and survival of the young of unaided females and control pairs were compared. Broods raised by unaided females gained body mass more slowly and fledged at slightly lower mass than those raised by two parents. However, fledging mass was not correlated with survival to independence. There were no differences in tarsus growth between the two treatment groups. Entire brood loss to predators occurred as often among females without male help as it did among those with male help. However, partial brood loss was more common among female-only broods than among controls; this difference was largely attributable to higher rates of starvation and exposure in female-only broods. There appeared to be an interaction between growth and predation. Female-only broods that were below the median mass of combined treatment groups at 5 days of age were more likely than all other broods to experience partial or complete predation. Male impact on offspring survival varied with age of the offspring. When years were combined, males tended to increase survival during the first half of the nestling period, but their impact at the time of nest-leaving was minimal. In all years, from nest-leaving to independence (ca. 2 weeks), broods without male help survived only about half as well as did those with male help. Independent young raised by one parent were as likely to return the following spring as were young raised by two parents. Thus, paternal care benefits males by improving the survivorship of their fledglings, and may also act as a buffer against poor female parental quality and inclement weather. However, the magnitude of these benefits is such that bigamous males might achieve greater reproductive success than monogamous males. Various possible male strategies are discussed.     

Intersexual niche differentiation in the blue tit (Parus caeruleus)

The intersexual niche differentiation hypothesis, according to which sexual size dimorphism (SSD) has evolved to facilitate niche separation between the sexes, has received equivocal support in studies of birds. One explanation for the inconsistency of the results is that other factors crucial for reproductive success, such as variation in food abundance among territories, may obscure food niche differentiation between parents. We studied the effects of SSD, food availability and food demands of broods on niche differentiation in the blue tit ( Parus caeruleus ). Parents attending experimentally enlarged broods delivered the same amount, but more diverse food items, to their offspring than parents attending reduced broods, although this effect was significant in only one of the two study years. The time-lag (deviation from the peak food abundance in habitat in days) and squared time-lag were the most important determinants of the diversity of food items delivered to young and their condition (fitness surrogate). SSD, as measured by principal component scores of morphological traits, did not explain any variation neither in the diversity or amount of food provided, nor in offspring condition. These results suggest that synchronization of breeding with peak food availability is the most important determinant of diversity of food delivered to young and, more importantly, Their condition. No evidence for intersexual food niche differentiation was found; at best, it is likely to constitute only a weak selection pressure on the maintenance or enhancement of SSD in this system.     

Offspring pay sooner,parents pay later: experimental manipulation of body mass reveals trade-offs between immune function,reproduction and survival

Introduction

Life-history theory predicts that organisms trade off survival against reproduction. However, the time scales on which various consequences become evident and the physiology mediating the cost of reproduction remain poorly understood. Yet, explaining not only which mechanisms mediate this trade-off, but also how fast or slow the mechanisms act, is crucial for an improved understanding of life-history evolution. We investigated three time scales on which an experimental increase in body mass could affect this trade-off: within broods, within season and between years. We handicapped adult skylarks (Alauda arvensis) by attaching extra weight during first broods to both adults of a pair. We measured body mass, immune function and return rates in these birds. We also measured nest success, feeding rates, diet composition, nestling size, nestling immune function and recruitment rates.

Results

When nestlings of first broods fledged, parent body condition had not changed, but experimental birds experienced higher nest failure. Depending on the year, immune parameters of nestlings from experimental parents were either higher or lower than of control nestlings. Later, when parents were feeding their second brood, the balance between self-maintenance and nest success had shifted. Control and experimental adults differed in immune function, while mass and immune function of their nestlings did not differ. Although weights were removed after breeding, immune measurements during the second brood had the capacity to predict return rates to the next breeding season. Among birds that returned the next year, body condition and reproductive performance a year after the experiment did not differ between treatment groups.

Conclusions

We conclude that the balance between current reproduction and survival shifts from affecting nestlings to affecting parents as the reproductive season progresses. Furthermore, immune function is apparently one physiological mechanism involved in this trade-off. By unravelling a physiological mechanism underlying the trade-offs between current and future reproduction and by demonstrating the different time scales on which it acts, our study represents an important step in understanding a central theory of life-history evolution.

     

Brood-directed parental aggression and early brood loss in the coral reef fish, Acanthochromis polyacanthus (Pomacentridae)

Brood-directed parental aggression, in which parents attack and even kill their own young, is described for the coral reef fish Acanthochromis polyacanthus. This behaviour involves both parents, occurs on days 7–16 post-hatching, and is characteristic of all broods produced early in the spawning season. Such parental aggression appears to underlie widespread early brood loss in the species, i.e. disappearance of young before they reach an age at which they are viable in the absence of parental defence. Simultaneously, there is also a high frequency of sudden increases in the sizes of some tended broods. The data suggest that some brood-tending parents are forcibly expelling their broods from parental territories, and fostering them on to adults still tending broods. By expelling their young and re-spawning, pairs may realize a 7% increase in annual fecundity over pairs that do not expel their young, despite a low survival rate of expelled young.     

FACTORS AFFECTING EGG-WEIGHT, BODY-WEIGHT AND MOULT OF THE WOODPIGEON COLUMBA PALUMBUS

The heaviest clutches (2 eggs) laid by Woodpigeons Columba palumbus in a Cambridgeshire study area weighed 30% more than the lightest. Yet the variation in egg-weight within clutches was less than 1 %. Irrespective of initial weight, eggs lost weight at the same constant rate during incubation. Heavy eggs hatched more successfully than light eggs and none weighing less than 16 g hatched. There was no correlation between chicks' weight at hatching and their weight at day 6 during the July-September part of the breeding season. The ability to feed crop milk at this stage could compensate for low chick-weight, but this might not be true early in the season. Weight at day 6 was correlated with the weight at day 16 or 17. The growth pattern is discussed. Chicks in broods of one achieved a higher weight at day 17 than those in broods of two. The survival rate both in and after leaving the nest was the same in both brood-sizes. Chick-weight in artificially created broods of three was almost as high as in broods of two, but again data refer to the July-September period when abundant cereal food is available. Survival before and after fledging was lower in broods of three. Clutch- and egg-weight declined from April until September. It is suggested that this is adaptive, in that the adults produce heavier eggs when food supplies are most difficult to collect. The critical period probably occurs during the few days when the adult must produce crop milk and the young cannot be left unattended. Thus egg-weight depends on the female's capacity to acquire nutrients, and is related to the needs of embryonic development and the amount of compensation in nutrient supply which can be provided immediately after hatching. But clutch-size is more related to the bird's ability to feed and rear young to the point of fledging, thereby influencing the number of offspring which survive to leave progeny. Egg-weight and female body-weight were positively correlated in females weighing less than 480 g but not in heavier females. First-year birds did not acquire adult weight until midsummer and they would probably produce light eggs if they could breed before this month. However, their gonads do not recrudesce until July and this prevents them breeding in the spring. Seasonal changes in body-weight and fat content of adults and first-year birds are described and discussed; differences were noted between adult males and females which were considered to be adaptive. The moult is described. It begins in April and continues until November, approximately one pair of primaries being replaced per month. The moult ceases during the winter months, when it is known that food supplies become limiting. Woodpigeons lay light eggs relative to their body-weight but can achieve the extra parental care needed for the altricial chicks by producing crop milk. Because the moult is extended, the energy demands of moulting and breeding combined are relatively low and this enables the Woodpigeon to have a long breeding season and to moult coincidentally.     

Evolution of single-chick broods in the Swallow-tailed Gull Creagrus furcatus

Swallow-tailed Gulls lay single-egg clutches, and so raise single-chick broods. As they are pelagic seabirds, this small brood size is expected to relate to proximate food limitation owing to infrequent food deliveries. However, a previous brood doubling experiment detected an 82% increase in fledging success from experimentally doubled broods compared to controls. We repeated the brood doubling experiment, and found that none of 50 enlarged broods produced more than one independent offspring. Control and experimental parents produced fledglings of similar body size, which also had indistinguishable rates of fledging and subsequent survival and reproduction. A variety of parameters estimating survival and breeding costs of reproduction showed no treatment effect. Since two-chick broods yield dramatically higher fledging rates at some times, apparently without excess costs of reproduction, selection on brood size appears to favour a two-chick brood. However, selection may not favour a two-egg clutch if egg production is very costly. Additionally, our estimates of reproductive success do not incorporate the performance of experimental and control offspring as adults, which could differ, since growth of chicks differed slightly by treatment.     

Why don't birds lay more eggs?

Fifty years ago David Lack put forward a key hypothesis in life-history theory: that avian clutch is ultimately determined by the number of young that parents can provide with food. Since then, a plethora of brood manipulations has shown that birds can rear more young than the number of eggs they lay, and prompted a search for negative effects of increased effort on future reproduction. However, recent studies have shown that the demands of laying and incubating eggs generally omitted from experiments, could affect parental fitness. Lack's hypothesis, and the tests of its validity, need to be extended to encompass the full demands of producing and rearing the brood.