Display rate by male fairy-wrens (Malurus cyaneus) during the fertile period of females has little influence on extra-pair mate choice

Empirical and theoretical studies have only recently begun to examine how females use complex multi-component displays when selecting mates. Superb fairy-wrens are well suited to the study of female choice because females have control over extra-group paternity and cuckold their mates at high rates, while males possess a variety of sexually selected traits. Available evidence suggests that females base their extra-group mate choice on the timing of male moult into breeding plumage or the onset of display. However, males continue to perform elaborate displays throughout the season, and direct most displays to females during their fertile period. We therefore conducted focal observations on fertile females to quantify the frequency of male display and used microsatellite genotyping to compare the role of display rate during the breeding season and the timing of male moult on female mate choice. We show that the addition of data on male display rate does not improve our ability to predict which males obtain extra-group paternity. The timing of male moult into breeding plumage remains the only predictor of male extra-group reproductive success. Nevertheless, we found that males displayed more to females that were unable to select extra-group mates on the basis of the timing of moult or the onset of display. This raises the possibility that there are circumstances when females use display rate to discriminate between potential extra-group sires. Overall this study supports the theoretical prediction that females are more likely to base their mate choice on reliable indicators of male quality such as fixed morphological traits and displays of endurance, in this case an early moult into breeding plumage and the performance of an elaborate display during the winter, than a flexible behavioural trait such as display rate during the breeding season. Received: 26 January 2000 / Revised: 1 August 2000 / Accepted: 26 August 2000     

Male-male competition ensures honest signaling of male parental ability in the three-spined stickleback (Gasterosteus aculeatus)

The importance of socially imposed costs for the evolution and maintenance of honest sexual signals has received less attention than other costs. Here I show that male-male competition can increase the honesty of sexual signaling in relation to male parental ability in a species with flexible signaling. When four three-spined stickleback (Gasterosteus aculeatus) males were allowed to court a female first separately and then in competition, red nuptial coloration under competition reflected male parental ability more accurately than red coloration when separated. Parental ability was determined as the ability of the male to raise a clutch of eggs to the hatching stage under interaction with other breeding and non-breeding males. The increased honesty under competition was probably due to subordinate males of poor parental ability decreasing their color expression under interaction to reduce the risk of fights with superior males. However, socially imposed costs of signaling were probably not the main factors maintaining honest signaling, as red coloration reflected male parental ability also in the absence of competition, although less accurately. Nevertheless, the small-scale differences that male-male competition induced can significantly facilitate adaptive female choice and have large impacts on sexual selection. Received: 7 July 2000 / Revised: 31 August 2000 / Accepted: 4 September 2000     

Female distribution affects mate searching and sexual selection in male northern water snakes (Nerodia sipedon)

Mating systems and sexual selection are assumed to be affected by the distribution of critical resources. We use observations of 312 mating aggregations to compare mate-searching success of male northern water snakes (Nerodia sipedon) in two marshes in which differences in mating substrate availability resulted in more than fourfold differences in female dispersion. Reproductive males had significantly larger home ranges where females were dispersed than where females were clumped. The number of females encountered by males increased significantly with male home range size where females were dispersed, and decreased significantly where females were clumped. Where females were clumped, males were more likely to encounter other males when they located females. We found no evidence in either population that mate searching was energetically expensive or that males with relatively more energy had larger home ranges. However, males with greater fat reserves at the start of the season participated in more mating aggregations when females were dispersed, suggesting that fat reserves could affect a male’s willingness to attempt mating or to persist in aggregations. When females were dispersed there was weak stabilizing selection acting to maintain male body size (β=–0.14), but strong directional selection favoring larger (β=0.50) and fatter (β=0.37) males. Over 7 years, the intensity of selection favoring larger males varied substantially (β=0.14–1.15), but that variation was not related to variation in the operational sex ratio. We found no evidence of directional selection on either body size (β=0.05) or fat reserves (β=0.10) of males when females were spatially clumped. Overall, the distribution of females had a pronounced effect on male behavior, on the factors that affected male success in locating females, and probably on the extent of sperm competition once females had been located. Received: 23 November 1998 / Received in revised form: 9 August 1999 / Accepted: 18 August 1999     

Ectoparasites of house sparrows (Passer domesticus): an experimental test of the immunocompetence handicap hypothesis and a new model

Conspicuous secondary sexual traits may have evolved as handicap-revealing signals or as badges of status. We present results of an experiment using males of the sexually dimorphic house sparrow (Passer domesticus), that support the idea that the male-specific bib can be both a handicap-revealing signal and a reliable badge indicating the physical condition of the bird. In a test of the immunocompetence handicap hypothesis, wild-caught adult male house sparrows were studied in captivity. Birds implanted with elevated doses of testosterone were more dominant, had higher circulating levels of both testosterone and corticosterone and they also harboured relatively larger ectoparasite loads. Higher parasite loads were also associated with individuals showing lower immunocompetence and larger changes in bib size. A new model for immunocompetence effects in sexual selection is introduced, integrating actions that the hypothalamopituitary axis exerts on gonads, adrenals and the thyroid gland. The ”integrated immunocompetence model” synthesizes both the ”handicap” (i.e. survival-decreasing) and ”badge of status” (i.e. survival- enhancing) models for evolution of secondary sexual traits. Received: 15 May 1999 / Received in revised form: 2 November 1999 / Accepted: 6 November 1999     

Testosterone is involved in acquisition and maintenance of sexually selected male plumage in superb fairy-wrens, Malurus cyaneus

Testosterone has been proposed as a physiological link between the level of sexual signalling and male condition. Bright plumage is one of the most noticeable sexual signals and is often used by females as a basis for mate choice. Yet bright male plumage is not necessarily testosterone dependent. We investigated the role of testosterone in the moult into seasonal nuptial plumage in male superb fairy-wrens. Early pre-nuptial moult is under intense intersexual selection and males can acquire the bright plumage any time between autumn and the next spring. Testosterone was always undetectable or very low in males in dull eclipse plumage. During the pre-nuptial moult, both the number of males with detectable testosterone and average testosterone levels increased sharply. High testosterone was more correlated with nuptial plumage than with presence of the cloacal protuberance (indicative of sperm storage). Subcutaneous testosterone implants always induced the pre-nuptial moult within 2–3 weeks after implantation, even well outside the natural time range of moulting. Moreover, removal of the implants before the nuptial plumage was completed, arrested the moult process. The evidence suggests that development of the nuptial plumage is testosterone dependent, although we cannot exclude that testosterone exerts its action after conversion to a metabolite such as oestrogen. Once the nuptial plumage was completed, all males maintained substantially elevated testosterone, sometimes months before the onset of breeding. These high levels could be necessary to maintain the plumage, and/or are involved in courtship displays. The results are discussed with respect to potential costs involved in acquiring and maintaining the nuptial plumage. Received: 17 January 2000 / Received in revised form: 24 February 2000 / Accepted: 25 February 2000     

Assessing the aerodynamic effects of tail elongations in the house martin (Delichon urbica): implications for the initial selection pressures in hirundines

Of the three species of hirundine that breed sympatrically across the U.K., one, the barn swallow, has outer tail feathers elongated into streamers, whereas the other two species, the house martin and the sand martin, do not. The tail streamer of the barn swallow is regarded as a classic example of a sexually selected trait. Recent evidence, however, has suggested that streamers may have evolved largely through natural selection for enhanced flight performance and increased maneuverability. We tested the hypotheses that small streamers (1) increase performance in turning flight, but (2) decrease performance in flight variables related to velocity. We manipulated the lengths of house martin outer tail feathers and measured changes in their free-flight performance, using stereo-video to reconstruct the birds" three-dimensional flight paths. Five flight variables were found to best describe individual variation in flight performance. Of these five, the three variables determining maneuverability predicted that flight performance would be optimized by a 6- to 10-mm increase in the length of the outer tail feathers. In contrast, for mean velocity and mean acceleration, extension of the outer tail feathers appears to have a detrimental effect on flight performance. We suggest that the initial selection pressure for streamers in ancestral short-tailed "barn swallows" was via natural selection for increased maneuverability. In addition, we propose that the benefits of increased maneuverability have differed between hirundines in the past, such that the cost of increasing the length of the outer tail feather has, to date, outweighed the benefits of doing so in streamerless hirundines. Received: 14 February 2000 / Revised: 2 July 2000 / Accepted 18 July 2000     

Racketed tail of the male and female turquoise-browed motmot: male but not female tail length correlates with pairing success,performance, and reproductive success

Both males and females of many avian species maintain elaborate plumage traits, and elaborate monomorphic plumage may convey adaptive benefits to one or both sexes as inter- or intraspecific signals. Both sexes of the turquoise-browed motmot (Eumomota superciliosa) are elaborately plumed with long racket-tipped tail. I investigated whether the racketed tail functions as a sexually selected signal in one or both sexes by testing the predictions that males and/or females with the largest tails have: (1) greater pairing success, (2) greater reproductive performance (clutch-initiation date, clutch size, and hatching success), and (3) greater reproductive success. Yearling males with longer denuded rachises (wires) on the central tail feathers had greater pairing success. In addition, adult males with longer wires paired with females who laid larger clutches, had greater hatching success independent of clutch size, and fledged more young. There was no relationship between female tail plumage and pairing success, reproductive performance, or fledgling success. These results are consistent with the hypothesis that male tail plumage functions as a mate choice or status signal, but that the tail of the female does not function in a sexually selected context. I discuss alternative hypotheses for the evolutionary maintenance of the elaborate female tail plumage.     

Testosterone and the allocation of reproductive effort in male house finches (Carpodacus mexicanus)

Testosterone has been proposed to serve as the mediator that controls the relative effort that an individual male bird will devote to mating effort versus parental effort. Here, we demonstrate a testosterone-influenced trade-off between parental and mating efforts in male house finches. Male house finches with experimentally elevated testosterone fed nestlings at a significantly lower rate, but sang at a higher rate than males without manipulated testosterone levels. Females mated to testosterone-implanted males fed nestlings at a significantly higher rate than females mated to males without testosterone implants, resulting in similar feeding rates for both treated and untreated pairs. The effects of testosterone on male house finches, however, were not as dramatic as the effects of testosterone observed in some other socially monogamous species of birds. Because extra-pair copulations are uncommon in house finches and males provide substantial amounts of parental care, these more modest effects may be due to differences in how the allocation of reproductive effort affects the costs and benefits of different reproductive behaviors. Received: 6 June 2000 / Accepted: 17 July 2000     

Post-mating odor in females of the solitary bee, Andrena nigroaenea (Apoidea, Andrenidae), inhibits male mating behavior

We investigated odor changes and their behavioral significance in the solitary, ground-nesting bee Andrena nigroaenea. We used gas chromatography with electroantennographic detection and performed behavioral tests with males in the field using natural odor samples and synthetic compounds. We found that only cuticle extracts of young females elicited copulation attempts in the males. We demonstrated that among the 17 compounds which triggered electroantennographic responses, all-trans-farnesyl hexanoate and all-trans-farnesol were significantly more abundant in unattractive cuticle extracts of A. nigroaenea females. Dufour’s gland extracts of these females also contained greater amounts of both compounds. In bioassays using synthetic farnesyl hexanoate and farnesol we found that these compounds inhibit copulation behavior in the males. Farnesyl hexanoate is probably synthesized in Dufour’s gland and used by females for lining brood cells. We interpret the semiochemical function of farnesyl hexanoate and its precursor farnesol to have evolved secondarily. As an outcome of sexual selection, it facilitates the discrimination by males of receptive females from nesting and thus already mated individuals. The dual function of these compounds represents an elegant parsimony in the chemical communication system of this insect. Received: 19 January 2000 / Revised: 29 May 2000 / Accepted: 24 June 2000     

Female resistance and male preference in a stream-dwelling isopod: effects of female molt characteristics

In the stream-dwelling isopod, Lirceus fontinalis, mating contests between males and females occur prior to pair formation. We examined the relative contribution of male preference and female resistance to contest outcomes. We first quantified male and female behavior during typical mating interactions and examined the relationship between time until molt (TTM) and mating outcomes. We then examined the role of male preference and female resistance in determining mating outcomes when females differed in molt type (growth, egg deposition) and appeared to differ in TTM (due to hormone applications). Both male preference and female resistance contributed to different components of the mating sequence but female resistance ultimately determined whether or not pair formation occurred. Males expressed a preference for females that appeared to be close to molt, using variation in levels of molt hormone as a cue. However, males did not discriminate between females based on molt type. Received: 5 March 1999 / Received in revised form: 10 August 1999 / Accepted: 16 October 1999     

Enhancement of reproductive success through mate choice in a social rock-wallaby, Petrogale assimilis (Macropodidae) as revealed by microsatellite markers

The reason why a female who is socially paired to one particular male seeks extra-pair copulations (EPCs) with others has important implications in life history models and to the study of behaviour. The Allied rock-wallaby, Petrogale assimilis, lives in spatially isolated colonies in tropical north Queensland, Australia. Extensive observations of a colony at Black Rock showed that intense behavioural bonding occurs between pairs of adult males and females; about two-thirds of males paired with one female, the remainder paired with two females simultaneously. Single-locus microsatellite profiling determined the paternity of 63 offspring from 21 females for which long-term behavioural data were available. One-third of the young were fathered by males which were not paired socially with the mother. The mating system was heterogeneous: (1) all offspring of 11 females were fathered by the mother's partner, (2) all young of 5 females were fathered by extra-pair males, and (3) only some of the young of 5 females were fathered by their regular consort. Analysis of individual longitudinal demographic records showed that females whose young were always fathered by their consort had higher reproductive success than those whose young were always fathered as a result of (EPCs). However, females with some offspring fathered by their regular consort and others via EPCs had the highest probability of raising young to independence. These females were significantly more likely to have an offspring fathered as a result of an EPC if their previous young had failed to survive to pouch emergence. These results are consistent with the hypothesis that females choose mates for their genetic quality. Comparison of the males with which these females sought EPCs and the regular consorts suggested that arm length rather than body weight or testes size was used as the index of genetic quality. Results from a second colony of rock-wallabies in which the reproductive rate was accelerated were also consistent with the genetic-quality hypothesis. These results imply that by choosing better-quality fathers irrespective of social pairing, females are able to maximise their overall lifetime reproductive success, and presumably, those of their offspring. Received: 8 June 1997 / Accepted after revision: 28 February 1998     

The effect of male coloration on female mate choice in closely related Lake Victoria cichlids (Haplochromis nyererei complex)

We studied the effect of male coloration on interspecific female mate choice in two closely related species of haplochromine cichlids from Lake Victoria. The species differ primarily in male coloration. Males of one species are red, those of the other are blue. We recorded the behavioral responses of females to males of both species in paired male trials under white light and under monochromatic light, under which the interspecific differences in coloration were masked. Females of both species exhibited species-assortative mate choice when colour differences were visible, but chose non-assortatively when colour differences were masked by light conditions. Neither male behaviour nor overall female response frequencies differed between light treatments. That female preferences could be altered by manipulating the perceived colour pattern implies that the colour itself is used in interspecific mate choice, rather than other characters. Hence, male coloration in haplochromine cichlids does underlie sexual selection by direct mate choice, involving the capacity for individual assessment of potential mates by the female. Females of both species responded more frequently to blue males under monochromatic light. Blue males were larger and displayed more than red males. This implies a hierarchy of choice criteria. Females may use male display rates, size, or both when colour is unavailable. Where available, colour has gained dominance over other criteria. This may explain rapid speciation by sexual selection on male coloration, as proposed in a recent mathematical model. Received: 11 April 1997 / Accepted after revision: 27 July 1997     

Evolution of worker sterility in honey-bees (Apis mellifera): how anarchistic workers evade policing by laying eggs that have low removal rates

In anarchistic honey-bee colonies, many workers’ sons are reared despite the presence of the queen. Worker-laid eggs are normally eaten by other workers in queenright colonies. Workers are thought to discriminate between queen-laid and worker-laid eggs by the presence or absence of a queen-produced egg-marking pheromone. This study compared the survival of three classes of eggs (worker-laid eggs from anarchistic colonies, worker-laid eggs from non-anarchistic queenless colonies, and queen-laid eggs) in both queenright normal colonies and queenright anarchistic colonies, in order to test the hypothesis that anarchistic workers evade policing by laying more acceptable eggs. As expected, few worker-laid eggs from non-anarchistic colonies survived more than 2 h. In contrast, worker-laid eggs from anarchistic colonies had much greater acceptability, which in some trials equalled the acceptability of queen-laid eggs. Anarchistic colonies were generally less discriminatory than normal queenright colonies towards worker-laid eggs, whether these originated from anarchistic colonies or normal queenless colonies. This indicates that the egg-removal aspect of the anarchistic syndrome involves both worker laying of eggs with greater acceptability and reduced discriminatory behaviour of policing workers. Received: 19 July 1999 / Received in revised form: 3 November 1999 / Accepted: 20 November 1999     

The influence of potential reproductive rate and variation in mate quality on male and female choosiness in the sand goby, Pomatoschistus minutus

Recent models of choosiness in mate choice have identified two particularly important factors: the potential reproductive rate (PRR) of the choosing sex relative to that of the chosen sex, and the variation in quality of potential mates. This experimental study tested how these factors affected choosiness in male and female sand gobies, Pomatoschistus minutus. We manipulated relative PRR by means of water temperature, and mate quality by means of body length. The choosing male or female was offered a choice between two mates with either a small or a large difference in body length representing a small or a large variation in mate quality. Choosiness was measured as (1) preference for the larger mate, and (2) as whether or not spawning occurred with the smaller mate, while the larger mate was visible but screened off. We found that females preferred large males, and that their level of choosiness was affected by variation in male quality, but not by their own relative PRR. Males, on the other hand, seemed unselective in all treatments and were in general more likely than females to spawn with their provided partner. This suggests that in the sand goby, variation in male mate quality has a greater influence than relative PRR on facultative changes in female choosiness. However, a general difference in PRR between males and females may be one important factor explaining the observed sex difference in choosiness. Received: 17 April 2000 / Revised: 24 June 2000 / Accepted: 17 July 2000     

Maternal investment in rhesus macaques (Macaca mulatta): reproductive costs and consequences of raising sons

Maternal investment in offspring is expected to vary according to offspring sex when the reproductive success of the progeny is a function of differential levels of parental expenditure. We conducted a longitudinal investigation of rhesus macaques to determine whether variation in male progeny production, measured with both DNA fingerprinting and short tandem repeat marker typing, could be traced back to patterns of maternal investment. Males weigh significantly more than females at birth, despite an absence of sex differences in gestation length. Size dimorphism increases during infancy, with maternal rank associated with son’s, but not daughter’s, weight at the end of the period of maternal investment. Son’s, but not daughter’s, weight at 1 year of age is significantly correlated with adult weight, and male, but not female, weight accounts for a portion of the variance in reproductive success. Variance in annual offspring output was three- to fourfold higher in males than in females. We suggest that energetic costs of rearing sons could be buffered by fetal delivery of testosterone to the mother, which is aromatized to estrogen and fosters fat accumulation during gestation. We conclude that maternal investment is only slightly greater in sons than in daughters, with mothers endowing sons with extra resources because son, but not daughter, mass has ramifications for offspring sirehood. However, male reproductive tactics supersede maternal investment patterns as fundamental regulators of male fitness. Received: 23 July 1999 / Received in revised form: 23 February 2000 / Accepted: 13 March 2000     

Genetic evidence for mixed parentage in nests of the emu (Dromaius novaehollandiae)

Parentage in emus (Dromaius novaehollandiae) was examined by microsatellite analysis using four independent loci. Of 106 chicks sampled in one breeding season from 18 nests, 54 (51%) were not fathered by the nesting male, 12 (11%) were not from the observed mate of the sitting male, and 9 (8%) represented intra-specific brood parasitism, having no alleles in common with either nest parent. Some males (11%) fathered all chicks in their nests, but the majority showed high levels of cuckoldry. Those males commencing incubation earliest in the season tended to have the highest levels of paternity in their own nests. These results reveal a high frequency of extra-pair fertilisations and resultant cuckoldry in a predominantly socially monogamous bird and support recent reports which have described the emu mating system as a complexity of polyandrous, promiscuous and monogamous behaviour. Parentage assignment of chicks resulting from extra-pair fertilisations revealed an evenly scattered pattern of paternity that did not show any particular male dominance in reproductive success. These results lead to a reassessment of behavioural observations of emus, the consequences of parentage distribution, and theories about mating systems and sexual selection. The frequency of extra-pair copulations and intra-specific brood parasitism suggests patterns of descent that differ greatly from those implied by social monogamy. Received: 27 November 1999 / Received in revised form: 8 January 2000 / Accepted: 8 February 2000     

Isopropyl (Z9)-hexadecenoate as a male attractant pheromone from the sternal gland of the rove beetle Aleochara curtala (Coleoptera: Staphylinidae)

Summary. In the rove beetle Aleochara curtula, a male specific sternal gland is described. Isopropyl (Z9)-hexadecenoate has been identified by GC/MS of surface and sternal gland extracts as a male specific compound. Its emission in the air was demonstrated by closed-loopstripping-analyses. In field experiments, conspecific males and females were attracted by the odor of carrion, caged males from laboratory cultures, and by the synthetic ester. Isopropyl (Z9)-hexadecenoate or live males, combined with fresh carrion attract more beetles than a fresh carcass alone. Isopropyl hexadecanoate as a minor compound was not attractive. The term “aggregation” pheromone and the ecological significance of attracting females to a fresh carcass allowing early copulation and egg-deposition are discussed. Received 20 August 1998; accepted 15 January 1999.     

Lifetime mating success, sexual selection and life history of fallow bucks (Dama dama)

We used data from a long-term study (15 years) of fallow deer to report for the first time the lifetime mating success, overall variance in lifetime mating success, and age-specific mortality levels of males. Fallow bucks that gain matings have higher social dominance rank, higher rates of fighting, and invest more in vocal display during the breeding season than unsuccessful males. Therefore, we examined if mating was associated with trade-offs in terms of survival, lifespan, and mating potential. We found that the variance in lifetime mating success was very high: 34 (10.7%) males mated, and of those, the 10 most successful males gained 73% of all matings (n=934). Mortality rates were generally high and only 22.3% (71/318) of males reached social maturity, i.e., 4 years. The oldest male was 13 years old. We found that fallow bucks that mated were not more likely to die during the following year, did not suffer from a reduction in lifespan, and did not incur lower mating success later in life as a result of mating during the early years of social maturity. Our results show that mating males at age 5 years (and possibly 9 years) may be more likely to survive than non-mating males. Additionally, the number of matings gained by males during the first years of social maturity was positively correlated with lifespan. We suggest that mating males are of higher quality than non-mating males because they are not more likely to incur trade-offs as a result of their increased reproductive efforts. Received: 9 November 1999 / Revised: 30 April 2000 / Accepted: 27 May 2000     

Electrosensory prey detection in the African sharptooth catfish, Clarias gariepinus (Clariidae), of a weakly electric mormyrid fish, the bulldog (Marcusenius macrolepidotus)

During their seasonal runs in the Okavango and other freshwater bodies in southern Africa, the sharptooth catfish, Clarias gariepinus, specialises on the bulldog, Marcusenius macrolepidotus, as its main prey. We examined whether the catfish can locate bulldogs by detecting their pulse-type electric organ discharges (EODs). The electrosensory threshold for single-cycle, monopolar square-wave pulses was exceedingly low (down to 13 μV_p–p/cm for 4-ms pulses) in trained, food-rewarded sharptooth catfish (n=4), confirming the results of Lissmann and Machin who were, however, unable to identify a biological function. Other stimulus pulse waveforms (single-cycle, monopolar as well as bipolar sine-wave pulses) were also effective stimuli according to their spectral low-frequency energy contents (0– 30 Hz). Male bulldogs display an EOD pulse approximately 10× the duration of female EODs (≈0.5 ms). The C. gariepinus threshold for field-recorded playbacks of a male bulldog EOD (of long duration) was 103 μV_p–p/cm, whereas the brief female and juvenile EODs were not detected (using intensities of natural EODs). EODs of other mormyrids were detected when either monopolar or of long duration. Signal source amplitude increased linearly with standard length (SL) in bulldogs. Signal reach, as calculated from signal source amplitude and receiver sensitivity, is up to 150 cm for a large male bulldog (SL 27.5 cm), and 83 cm for a male that has just turned sexually mature (SL 12.6 cm). Therefore, most bulldogs eaten by catfish are probably male, in agreement with the size distribution of bulldogs found in catfish stomachs. These results suggest that sharptooth catfish rely heavily on their acute electrical sense during hunting, and an important function for electroreception in an African catfish has been identified. Received: 13 December 1999 / Revised: 20 April 2000 / Accepted: 20 May 2000     

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