Clutch desertion and re-nesting in pied flycatchers: an experiment with progressive clutch removal

Clutch desertion and re-nesting are important components of fitness when predation is frequent. In nestbox populations however, nest predation and desertion are rare but can be studied by experimental manipulations. We experimentally reduced clutches of pied flycatcher,Ficedula hypoleucaby removing one egg per day until desertion occurred. The size of the clutch at desertion and whether females re-nested or not were used as measures of the female response. Of the deserting females, 74% re-nested in our study area. Re-nesting frequency was correlated with date but not with the size of the clutch laid. The majority of the non-re-nesting females deserted empty nests, while the majority of re-nesting females deserted one egg. Clutch size at desertion was not correlated with the size of the clutch laid nor with laying date; it was smaller than the size predicted by an optimality analysis of the value of both the current (deserted) and the replacement clutch. For the re-nesting females, there was a negative correlation between fledging rate of the replacement clutch and the size of the clutch at desertion. Our predictions, made under the hypothesis that desertion and re-nesting are adaptive behaviours, were partly supported by the data; we explain the discrepancy by the constraint of searching for a new nest site or mate for re-nesting.     

Nest desertion by a cowbird host: an antiparasite behavior or a response to egg loss?

Natural selection can favor songbirds that desert nests containingeggs of the parasitic brown-headed cowbird (Molothrus ater).However, the high variability in desertion of parasitized nestswithin species is perplexing in light of the typically highcosts of parasitism. Because nest desertion can also be a responseto partial clutch predation, we first asked if Bell's vireos(Vireo bellii) deserted nests in response to the presence ofcowbird eggs (antiparasite response hypothesis) or to egg removalby predators and female cowbirds (egg predation hypothesis).Second, we asked whether variation in nest desertion was dueto intrinsic differences among individuals or to variation innest contents. We monitored a large number of nests (n = 494)and performed a clutch manipulation experiment to test thesehypotheses. The number of vireo eggs that remained in a nestwas a strong predictor of desertion both within and among pairs.Neither the presence of a single cowbird egg, which leads tonest failure for this host, nor the number of cowbird eggs receivedin a vireo nest influenced nest desertion. Furthermore, vireosdid not desert experimental nests when we immediately exchangedcowbird eggs for vireo eggs but deserted if we removed vireoeggs and replaced them with cowbird eggs the following morning.Desertion of parasitized nests by Bell's vireos can be almostentirely explained as a response to partial or complete clutchloss and does not appear to have been altered by selection frombrood parasitism.     

Indeterminate laying and flexible clutch size in a capital breeder, the common eider

Clutch size control in capital breeders such as large waterfowl has been much debated. Some studies have concluded that clutch size in ducks is determined before the start of laying and does not change in response to egg additions or removals. The response, however, may depend on the timing of tests, and experiments may have been too late for females to alter the number of eggs. We here study clutch size responses to predation of first and second eggs in the common eider, using protein fingerprinting of egg albumen to verify that the same female continues laying in the nest after predation. Sixty of 79 females with early egg predation (one or both of the two first eggs) deserted the nest. Among the 19 females that stayed and continued laying, the mean number of eggs produced was 4.4, significantly higher than the 3.7 in non-predated nests. The staying females had similar egg size and clutch initiation date as females that deserted, and their body mass and clutch initiation date was similar to that of females whose clutches were not predated. Even capital-breeding common eiders may therefore be indeterminate layers, as many females in which early eggs are removed lay more eggs than others. A previous study has shown that they can reduce their laying if eggs are added. Our results add to increasing evidence that ducks have more flexible egg production than previously thought.     

Effects of clutch sizes and incubation stage on nest desertion in the female Common Eider Somateria mollissima nesting in the high Arctic

In bird species, one of the trade-offs between reproduction and survival appears in the parental decision to desert the nest. Nest desertion is modulated by several factors including clutch size. However, the incubation stage at which predation occurs is also an important factor. In this study, we examined whether nest desertion was linked to initial clutch size, partial clutch predation (final clutch size) and the incubation stage at which it happened in a capital breeder: the female common eider (Somateria mollissima) nesting in the high Arctic. The study was performed in Kongsfjorden in 2002 on the western coast of the Svalbard Archipelago (78°55′N, 20°07′E). We observed that nest desertion was higher when the initial clutch size was small. Also, females deserted their nests more during the first third of incubation than later. Thus, as incubation proceeded, nest desertion was less likely to occur even after egg reduction. Our results pointed out that this parental decision in female eiders seemed to depend on initial clutch size and on the date into incubation of clutch reduction. Sophie Bourgeon and Francois Criscuolo contributed equally to the work     

A molecular genetic analysis of the communal nesting of the ostrich (Struthio camelus)

The ostrich breeding system is complex and unique; communal clutches are laid by several females, although only one female, the major female, and the resident territorial male provide parental care. More eggs are laid in the nest than can be incubated and the major female ejects surplus eggs from the incubated central clutch. Microsatellite markers were used to analyse the parentage of communal nests in Nairobi National Park. This revealed that major females contributed a disproportionate number of fertile eggs to the central, incubated clutch and that multiple paternity and maternity within a nest were common; 68.9% of all incubated eggs on a nest were not parented by both the resident territorial male and the major female of that nest. All the males fertilized eggs on the clutches of neighbouring males. Unexpectedly, every major female with her own nest was also simultaneously a minor female with incubated eggs on neighbouring clutches. The relatedness between females laying in the same nest was not significantly different from the population average and significantly less than that between chicks hatched from the same nest.     

Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.     

Offspring desertion and parental care in the Whiskered Tern Chlidonias hybrida

In species with biparental care, a conflict of interest can arise if one mate tries to maximize its own reproductive success at the expense of the other's. One of the mates can desert the brood to accrue a number of benefits to enhance its own fitness, leaving parental care to the remaining parent. This study is the first to describe the desertion pattern in a tern species (Sternidae). We investigated offspring desertion in the Whiskered Tern Chlidonias hybrida, a species with semi‐precocial chicks. Offspring desertion was recorded in 52% of nests prior to fledging (n = 131 nests). Females also deserted during the post‐fledging period. Of the deserters, 97% were females. Desertions started when chicks were 5 days old and no longer required intense brooding. Desertions before fledging did not affect fledging success. Provisioning rates between pair members differed, and females supplied much less food than males. Female provisioning rate affected the chances of nest desertion significantly: daily desertion rates were lower when females supplied more food. After females had deserted, males increased their provisioning rates but compensated for the loss of female care only partly in two‐ and three‐chick broods. Only in small (one‐chick) broods was compensation full. We conclude that male and female Whiskered Terns adopt different reproductive strategies in the population studied here. Females invest much less in parental care than males, providing less food and deserting more frequently. Given the ready availability of food and low predation pressure, benefits appear to accrue to females that desert; selection forces may therefore not be acting against female desertion.     

Effect of clutch size on male egg-fanning behavior and hatching success in the goby, Eviota prasina (Klunzinger)

In fish that exhibit paternal care, the females often choose their mates on the basis of male traits that are indicative of the parental ability of the males. In a marine goby, Eviota prasina, males tend their eggs within their nests until hatching, and females prefer males that have longer dorsal fins and exhibit courtship behavior with a higher frequency as their mates. In order to clarify the relationship between these sexually selected traits and the parental ability of males of E. prasina, the factors affecting the hatching success of eggs within male nests and the male parental care behavior were examined in an aquarium experiment. Females spawned their eggs in male nests and the clutch size of females showed a high individual variation (range = 88-833 eggs). The hatching success of eggs within male nests showed a positive correlation with the time spent by males in fanning eggs and the clutch size. In contrast to the prediction, however, the hatching success did not show a significant correlation with the sexually selected traits, i.e., the male dorsal fin length and the frequency of courtship displays. Moreover, multiple regression analysis indicated that the time spent by the males in fanning was the most important factor affecting the survival rate of the eggs. The time spent by males in fanning behavior was influenced by the clutch size within their nests; the fanning behavior of males occurred with a higher frequency when they tended larger clutches. Males are required to invest a greater effort in egg-tending behavior to achieve a higher hatching success when they receive larger clutches, probably due to the greater reward for their parental behavior. Based on their mate choice, females may obtain other benefits such as high quality offspring.     

Nesting ecology of a naturalized population of Mallards Anas platyrhynchos in New Zealand

Investigating the reproductive ecology of naturalized species provides insights into the role of the source population's characteristics vs. post‐release adaptation that influence the success of introduction programmes. Introduced and naturalized Mallards Anas platyrhynchos are widely established in New Zealand (NZ), but little is known regarding their reproductive ecology. We evaluated the nesting ecology of female Mallards at two study sites in NZ (Southland and Waikato) in 2014–15. We radiotagged 241 pre‐breeding females with abdominal‐implant transmitters and measured breeding incidence, nesting chronology and re‐nesting propensity. We monitored 271 nests to evaluate nest survival, clutch and egg size, egg hatchability and partial clutch depredation. Breeding incidence averaged (mean ± se) 0.91 ± 0.03, clutch size averaged 9.9 ± 0.1 eggs, 94 ± 2% of eggs hatched in successful nests, partial depredation affected 6 ± 1% of eggs in clutches that were not fully destroyed by predators, and re‐nesting propensity following failure of nests or broods was 0.50 ± 0.003. Nesting season (first nest initiated to last nest hatched) lasted 4.5 months and mean initiation date of first detected nest attempts was 28 August ± 3.3 days. Smaller females were less likely to nest, but older, larger or better condition females nested earlier, re‐nested more often and laid larger clutches than did younger, smaller or poorer condition females. Younger females in Southland had higher nest survival; cumulative nest survival ranged from 0.25 ± 0.007 for adult females in Waikato to 0.50 ± 0.007 for yearling females in Southland. Compared with Mallards in their native range, the nesting season in NZ was longer, clutches and eggs were larger, and nest survival was generally greater. Different predators and climate, introgression with native heterospecifics and/or the sedentary nature of Mallards in NZ may have contributed to these differences.     

Egg burial in penduline tits, Remiz pendulinus: its role in mate desertion and female polyandry

One important component in the mating strategy of an already-matedindividual is the decision to remain with the partner and carefor the offspring or to desert. Almost all research on nestdesertion has focused on the costs and benefits of continuedparental care versus desertion of both parents. However, ifit pays both parents to desert, the timing of desertion is mostimportant. In birds, where all eggs are fertilized well beforethe last egg is laid, males should be the first to desert. Evenif females try to hide their fertile period, it is likely thatthe appearance of eggs acts as a cue that males can use to calculatethe timing of their desertion. Here we examine egg burial behaviorin penduline tits and its possible effects on parental behaviorand desertion. Penduline tits perform uniparental care fromthe earliest point of breeding, and both sexes try to becomepolygamous. We found that 36% of investigated males were polygynousand deserted as soon as the first egg appeared in their nest,and 12.5% of females became polyandrous. About 27% of the nestswere deserted by both sexes, which means high costs for femalesin terms of wasted energy in eggs and for males in terms ofwasted energy and time in building elaborate nests. Femalescover the eggs, and several facts indicate that egg coveringis a deceptive behavior of females: (1) females cover the eggsin the morning before leaving the nest for the first time, (2)females are more aggressive toward their mates during the layingperiod than before laying, (3) females try to prevent malesfrom entering the nest when eggs have been experimentally uncovered,and (4) females uncover the eggs as soon as males are experimentallyremoved. Finally, we found that a female can only desert thenest before the male deserts when she covers the eggs. We concludethat the higher the proportion of eggs a female can hide, thegreater her chance of becoming polyandrous     

Don'T put all your eggs in one nest: spread them and cut time at risk

Abstract In many egg-laying animals, some females spread their clutch among several nests. The fitness effects of this reproductive tactic are obscure. Using mathematical modeling and field observations, we analyze an unexplored benefit of egg spreading in brood parasitic and other breeding systems: reduced time at risk for offspring. If a clutch takes many days to lay until incubation and embryo development starts after the last egg, by spreading her eggs a parasitic female can reduce offspring time in the vulnerable nest at risk of predation or other destruction. The model suggests that she can achieve much of this benefit by spreading her eggs among a few nests, even if her total clutch is large. Field data from goldeneye ducks Bucephala clangula show that egg spreading enables a fecund female to lay a clutch that is much larger than average without increasing offspring time at risk in a nest. This advantage increases with female condition (fecundity) and can markedly raise female reproductive success. These results help explain the puzzle of nesting parasites in some precocial birds, which lay eggs in the nests of other females before laying eggs in their own nest. Risk reduction by egg spreading may also play a role in the evolution of other breeding systems and taxa-for instance, polyandry with male parental care in some birds and fishes.     

Sex roles, parental effort and offspring desertion in the monogamous Eurasian Curlew Numenius arquata

The reasons for female desertion of offspring and the evolution of predominantly male care among monogamous bird species are not clearly understood. We studied parental effort during the incubation and chick rearing periods in the Eurasian Curlew Numenius arquata in western Finland, and compared timing of brood desertion with other populations in Europe. Males and females contributed equally to incubation and showed no differences in the intensity of mobbing behaviour towards a potential nest predator (stuffed crow) shortly after hatching. However, females deserted their offspring approximately halfway through the brooding period ( c. 16 d after hatching), while males remained with chicks until independence ( c. 35 d). Females with late-laid clutches deserted their offspring sooner after hatching than those with clutches produced earlier in the season. Curlew females deserted younger chicks in northeast Europe, where laying dates were later, breeding seasons shorter and migration distances were longer, than in western and central Europe. We suggest that the most likely reasons for offspring desertion by females may be associated with increased female survivorship and maintenance of pairbond between years.     

Male parental care and its adaptive significance in a neotropical frog

In the Puerto Rican frog Eleutherodactylus coqui, parental care is performed exclusively by males, and consists of attending the eggs and hatchlings at a terrestrial oviposition site. The two major behavioural components of parental care are egg brooding and nest defence against conspecific egg cannibals. Defence behaviour includes aggressive calling, biting, sustained biting, wrestling, and blocking directed against nest intruders. Parental care lasts from oviposition to hatching (17–26 days) and often for several days after hatching. During pre-hatching development, males are present in their nests 97.4% of the time during the day and 75.8% of the time at night. A large portion of this time is spent brooding eggs. In a field experiment, males were removed from their nests and the fate of clutches was monitored. Compared to control clutches (males not removed), experimental clutches had significantly lower hatching success and suffered significantly greater mortality from desiccation and cannibalism. Hence, parental care yields significant benefits to male fitness via increased offspring survival.     

The bright incubate at night: sexual dichromatism and adaptive incubation division in an open-nesting shorebird

Ornamentation of parents poses a high risk for offspring because it reduces cryptic nest defence. Over a century ago, Wallace proposed that sexual dichromatism enhances crypsis of open-nesting females although subsequent studies found that dichromatism per se is not necessarily adaptive. We tested whether reduced female ornamentation in a sexually dichromatic species reduces the risk of clutch depredation and leads to adaptive parental roles in the red-capped plover Charadrius ruficapillus, a species with biparental incubation. Males had significantly brighter and redder head coloration than females. During daytime, when visually foraging predators are active, colour-matched model males incurred a higher risk of clutch depredation than females, whereas at night there was no difference in depredation risk between sexes. In turn, red-capped plovers maintained a strongly diurnal/nocturnal division of parental care during incubation, with males attending the nest largely at night when visual predators were inactive and females incubating during the day. We found support for Wallace''s conclusion that reduced female ornamentation provides a selective advantage when reproductive success is threatened by visually foraging predators. We conclude that predators may alter their prey''s parental care patterns and therefore may affect parental cooperation during care.     

Major benefits of guarding behavior in subsocial bees: implications for social evolution

Parental care is a behavior that increases the growth and survival of offspring, often at a cost to the parents' own survival and/or future reproduction. In this study, we focused on nest guarding, which is one of the most important types of extended parental care; we studied this behavior in two solitary bee species of the genus Ceratina with social ancestors. We performed the experiment of removing the laying female, who usually guards the nest after completing its provisioning, to test the effects of nest guarding on the offspring survival and nest fate. By dissecting natural nests, we found that Ceratina cucurbitina females always guarded their offspring until the offspring reached adulthood. In addition, the females of this species were able to crawl across the nest partitions and inspect the offspring in the brood cells. In contrast, several Ceratina chalybea females guarded their nests until the offspring reached adulthood, but others closed the nest entrance with a plug and deserted the nest. Nests with a low number of provisioned cells were more likely to be plugged and abandoned than nests with a higher number of cells. The female removal experiment had a significantly negative effect on offspring survival in both species. These nests frequently failed due to the attacks of natural enemies (e.g., ants, chalcidoid wasps, and other competing Ceratina bees). Increased offspring survival is the most important benefit of the guarding strategy. The abandonment of a potentially unsuccessful brood might constitute a benefit of the nest plugging behavior. The facultative nest desertion strategy is a derived behavior in the studied bees and constitutes an example of an evolutionary reduction in the extent of parental care.     

Factors affecting piping plover hatching success on Long Island,New York

Low hatching success may limit progress towards reaching productivity goals for Atlantic Coast piping plover (Charadrius melodus) recovery, despite management strategies to protect eggs from predators and decrease human disturbance of birds on nests. We measured piping plover hatching success on Eastern Long Island beaches and identified the major causes of egg failure to better understand why eggs that were otherwise intact (not depredated or destroyed by tidal flooding) failed to hatch. We documented egg and nest fates, dissected contents of unhatched eggs to determine viability, and recorded human and predator activity near a subset of plover nests on Suffolk County Parks properties. The low hatching success we recorded (0.60) in 2006 and 2007 would require higher chick survival rates than are typically observed for piping plovers to meet recovery targets for productivity. Few eggs showed signs of poor viability and overall egg hatchability was comparable to other ground nesting birds. Most egg failure was due to either depredation at unexclosed nests or nest abandonment by adults. The best predictor of nest abandonment was the maximum number of red fox tracks (Vulpes vulpes) counted on nearby transects (β = −1.16, 95% CI: −2.0 to −0.3) and we found evidence that plovers abandoned eggs in response to predation risk (e.g., a fox circling a nest exclosure). Adults from abandoned nests may have deserted eggs or been depredated. In either case, intact and viable eggs were abandoned. Nest abandonment was not related to human activity near nests, which were buffered from human disturbance by symbolic string fencing. Our results suggest that depredation and nest abandonment (e.g., desertion or death of adults) due to predator disturbance, not human disturbance or poor egg viability, contributed to the low hatching success we recorded. Active predator removal in addition to modification of predator exclosure use and design may be necessary to prevent direct (egg depredation) and indirect (nest abandonment) negative effects of predators on hatching success. © 2010 The Wildlife Society.     

The influence of parental behavior on vulnerability to nest predation in tropical thrushes of an Andean cloud forest

The Skutch hypothesis predicts that parental activity around the nest may attract the attention of predators and thus, in the tropics where predation pressure may be high, selection favors reduced parental activity. This hypothesis has been questioned by studies demonstrating that parents can decrease the risk of nest predation through nest defense. The link between parental activity and predation risk may be further confounded by nest site characteristics. We examined the effects of parental behavior and nest site on clutch survival in two sympatric tropical thrushes (Myadestes ralloides and Turdus leucops). We compared survival rates of clutches in three treatments: 1) natural nests at the incubation stage, 2) unattended nests (un‐manipulated nests of the same species, with clutches unattended by parents), and 3) exposed clutches (eggs exposed in unconcealed positions, unprotected by the nest). Parental activity had a positive effect on clutch survival, which was revealed by significantly higher survival rate of clutches in attended nests compared to unattended nests. The effect of nest site was less clear: clutches in unattended natural nests survived better than clutches in exposed sites selected by humans, but results were insignificant. We propose that parent birds can exclude a group of opportunistic predators, that are able to destroy unattended clutches. Nest site characteristics may be less important in determining clutch survival in the tropics, where predator guilds are more diverse, making completely safe sites difficult to find. Our results challenge Skutch's hypothesis and point to the need for more data from tropical latitudes.     

Evidence of selection for egg crypsis in conspicuous nests

ABSTRACT The value of egg coloration as crypsis, once accepted as a general principle, has recently been questioned because most experiments have failed to show that egg coloration deters predation. The nest‐crypsis hypothesis postulates that, among species that build conspicuous nests, selection for egg crypsis is relaxed or absent because visually searching predators detect nests prior to eggs. I tested the nest‐crypsis hypothesis using the large, relatively conspicuous nests of American Robins (Turdus migratorius), and eggs that differed markedly in color that were collected from the nests of Red‐winged Blackbirds (Agelaius phoeniceus), Brewer's Blackbirds (Euphagus cyanocephalus), and Yellow‐headed Blackbirds (Xanthocephalus xanthocephalus). Each nest (N= 22) received a clutch of each species during three sequential predation trials that were 16 d in duration. The order of clutch presentation was randomized for each nest. Survival trends for Brewer's and Yellow‐headed Blackbirds were similar, and higher than those for clutches of Red‐winged Blackbirds. By the end of trials, overall survival of the three clutch types was roughly equivalent. However, clutches of Red‐winged Blackbird eggs, the most conspicuous egg type to the human eye, were discovered sooner by predators. Because the experimental design controlled for effects of nest crypsis, nest location, and nest size, this difference in egg survival can be attributed to differences in egg pigmentation. Thus, my results support a role for egg coloration as camouflage in conspicuous nests.     

Costs and response to conspecific brood parasitism by colonial red-breasted mergansers

Costs of conspecific brood parasitism (CBP) are expected to be influenced by a species’ life history traits. Precocial birds lay large clutches, and clutches that have been enlarged by CBP can affect host fitness through a longer incubation period, displaced eggs, and lower hatching success. We examined costs and response to CBP by hosts in a population of colonial red-breasted mergansers (Mergus serrator; n?=?400 nests over 8 years) within which 29% of parasitized clutches were enlarged considerably (≥?15 eggs). Length of the incubation period did not increase with clutch size. The mean number of eggs displaced from a parasitized nest during incubation (2.8) was 2×?greater than at an unparasitized nest (1.4). Hatching success declined by 2% for each additional egg in the nest. Thus, for a nest with?≥?15 eggs, one or more fewer host eggs hatch relative to an unparasitized nest with the same number of host eggs, assuming equal probability of success for all eggs. Hosts were 40% more likely to desert nests receiving 2 or 6 experimental eggs relative to unparasitized control nests, although it is unknown whether hens deserting a nest renested elsewhere. Our study indicates that costs of CBP to hosts during nesting may be limited to those red-breasted mergansers incubating the largest clutches (≥?15 eggs), and it raises questions about the adaptive significance of deserting a parasitized clutch.     

Distribution and breeding ecology of the Ferruginous Duck Aythya nyroca in Algeria

We conducted a survey of the distribution of Ferruginous Duck Aythya nyroca across Algeria and analysed the influence of nest-site characteristics on nesting success at a Ramsar site, Lake Tonga. The species was found to occupy different wetlands (freshwater lakes, brackish marshes and salt lakes) across three major climatic belts (subhumid coastal strip, semi-arid Hauts Plateaux and arid Sahara). Mean clutch size of successful nests was 13.3 ± 6.0 eggs (N = 26) with a hatching rate of 74% for successful clutches. Nesting success was recorded for 44% of nests with clutch desertion (72.7%) accounting for the majority of failed clutches. Egg size of the study population, which breeds at the southern limit of the species’ range, was significantly smaller than that of its northerly counterparts. Clutch size was negatively related to egg size and positively associated with depth of water below the nest, suggesting that older, more experienced or high-quality birds monopolised the safest sites. Successful clutches were significantly associated with tall vegetation, suggesting that when nesting is carried out mainly on offshore floating islets, protection against aerial predators and heat stress determine nesting outcome. Conspecific brood parasitism was significantly and positively associated with deferred egg-laying and smaller egg size, suggesting an age-dependent mechanism.