Parental care of the Common Tern Sterna hirundo

Parental care activities of male and female Common Terns Sterna hirundo were recorded over two breeding seasons. Males and females exhibited distinct parental roles throughout a breeding bout. Courtship feeding by males was extensive prior to and during egg-laying, but declined with the onset of incubation. Females performed significantly more incubation behaviour than males although both sexes spent equal time attending at the nest site. During the chick stage, females spent significantly more time on the territory than did males. Chick feeding was largely the responsibility of the male; males fed chicks at a rate approximately three times higher than that of females. In addition, whereas females showed no trend in the size of fish delivered to chicks relative to chick age, the size of fish delivered by males increased with chick age. Courtship feeding activities and extensive chick feeding contributions by male Common Terns appear to outweigh parental contributions by females, contrary to predictions for a monogamous species.     

Parental behaviour in the Lapwing Vanellus vanellus

Parental behaviour of monogamous and polygynous Lapwings was studied during incubation and brood care. Both parents attended the nest in 86% of monogamous pairs ( n = 29 pairs). In 14% of pairs, only the male parent continued incubation until the eggs hatched, whereas the female deserted the clutch before or at the end of incubation. There was a clear division of parental roles during incubation. Females spent more time incubating (64% of time) than their mates (27%), whereas males spent more time defending the nest (3%) than females (>1%). Time spent incubating did not differ between monogamous and polygynous males. However, polygynous females spent more time incubating (primary females: 95%; secondary females: 97%) than monogamous females. Biparental care was the most common pattern of post-hatching care, although in some broods either the male or the female parent deserted before the chicks fledged. Division of sex roles was less pronounced in brood care than during incubation. Females spent more time brooding (21%) than males (3%), and females attended their chicks more closely than males. Nevertheless, males and females spent similar amounts of time defending the brood from predators and conspecifics. We suggest that the apparent division of parental roles may be explained by sexual selection, i.e. the remating opportunities for male Lapwings might be reduced if they increase their share in incubation. However, the different efficiency of care provision, for example in ability to defend offspring, may also influence the roles of the sexes in parental care.     

Breeding behaviour of the Bearded Vulture Gypaetus barbatus: minimal sexual differences in parental activities

Monogamous biparental care is expected to occur when opportunities for extra-pair copulations are rare, and both parents are required to raise the chick. Bearded Vultures Gypaetus barbatus fulfill these conditions. Contributions by male and female Bearded Vulture to nest building, nest defence, incubation, nest attendance and chick feeding were studied over five years in eight pairs from the Pyrenees [Catalonia, northeast Spain). Overall, the sexes show equal investment, although the degree of parental effort developed differs depending on the specific activities. During pre-laying, males were significantly more active than females in supplying material to the nest and in territorial defence behaviour, which increased (in both sexes) as the season advanced. Incubation was shared equally both by day and by night. During chick-rearing, the nest was attended by both sexes and the presence of both parents at the nest decreased in parallel with the growth of the chick. Activities related to chick feeding were also equally divided. These results are discussed in the context of female selection of mates and the particular ecology of this species.     

Reversed sexual size dimorphism and parental care in the Red-footed Booby Sula sula

The 'division-of-labour' hypothesis predicts that males and females perform different roles in parental care and that natural selection acts differently on each sex so as to produce different body size optima suited to their particular roles. Reversed sexual size dimorphism in avian species (females larger than males) may therefore be an adaptive consequence of different roles of males and females in parental care. We investigated patterns of nest attendance, brooding, foraging and provisioning rate in a tropical seabird, the Red-footed Booby Sula sula , a species showing a reversed sexual size dimorphism. During incubation, females attended the nest more often than males, and spent more time brooding the small chick than did males during daytime. Males and females did not differ in the average duration of their foraging trips. During incubation, there was a positive relationship between nest attendance and the duration of foraging trips in males, but not in females. During the small-chick stage, for the same time spent at the nest, males spent significantly more time than females at sea. On average, females fed the chick more often than did males. In males, there was a significant and positive relationship between the probability of feeding the chick and the duration of the foraging trip, whereas in females, this probability was much less dependent on the duration of the foraging trip. Overall, female Red-footed Boobies achieved slightly, but significantly, more parental commitment than did males. However, these sexual differences in parental participation were small, suggesting a minimal division of labour in the Red-footed Booby. Our results suggest that the division of labour hypothesis is unlikely to explain fully the adult size dimorphism in Red-footed Boobies.     

Individual variation in field metabolic rate of kittiwakes (Rissa tridactyla) during the chick-rearing period

Field metabolic rate (FMR), using the doubly labelled water (DLW) method, was measured in free-ranging adult kittiwakes (Rissa tridactyla) early and late in the chick-rearing period at Svalbard, Norway. Individual variation in FMR was analysed by comparing FMR with body mass, sex, nest attendance, chick age, brood size, and basal metabolic rate (BMR). Mean FMR of kittiwakes during the chick-rearing period was 27.0+/-0.9 (SE) W kg(-1), while the individual variation (calculated as coefficient of variation [CV]) in FMR was 24%. Sex, time spent away from the nest, age of the chicks, and brood size contributed significantly to FMR and explained 65% of the variation in FMR. The FMR increased by 32% from early until late in the chick-rearing period. This occurred simultaneously with an increase in the time spent away from the nest. In 15 of 20 pairs, one of the mates had 15% or higher (mean of the 15 pairs, 22%+/-8%) FMR (W kg(-1)) than their partner, even though the mates spent equal amounts of time away from the nest. This indicates an intrapair conflict in FMR. The variation in total FMR of pairs was 40% less than the individual variation, and total FMR of pairs increased with age of the chicks. This indicates that the mates adjust their energy expenditure within a relatively constant FMR determined by the energy needs of the chicks. Individual variation in FMR could not be explained by variation in body mass or BMR. BMR measured late in the chick-rearing period was 26% lower than previous measurements of BMR from the prebreeding and incubation periods. The increase in FMR and simultaneous decrease in BMR caused a 40% increase in metabolic intensity (FMR/BMR) of kittiwakes during the chick-rearing period. It is suggested that the metabolic intensity is not a proper measure of the metabolic load in seabirds.     

Post-fledging brood and care division in the roseate tern (Sterna dougallii)

Extended post-fledging parental care is an important aspect of parental care in birds, although little studied due to logistic difficulties. Commonly, the brood is split physically (brood division) and/or preferential care is given to a subset of the brood by one parent or the other (care division). Among gulls and tern (Laridae), males and females generally share parental activities during the pre-fledging period, but the allocation of parental care after fledging is little documented. This study examined the behaviour of male and female roseate terns (Sterna dougallii) during the late chick-rearing and early post-fledging periods, and in particular the amount of feeds and the time spent in attendance given to individual chicks/fledglings. Pre-fledging parental care was biparental in all cases. Post-fledging parental care was dependent on the number of fledglings in the brood. Males and females continued biparental care in clutches with one surviving fledgling, while in two-fledgling clutches, males fed the A-fledgling while females fed the B-fledgling. Overall, there was no difference in attendance, only in feeds. This division of care may be influenced by the male only being certain of the paternity of the A-chick but not by chick sex.     

Relationship between reversed sexual dimorphism,breeding investment and foraging ecology in a pelagic seabird,the masked booby

Reversed sexual dimorphism (RSD) may be related to different roles in breeding investment and/or foraging, but little information is available on foraging ecology. We studied the foraging behaviour and parental investment by male and female masked boobies, a species with RSD, by combining studies of foraging ecology using miniaturised activity and GPS data loggers of nest attendance, with an experimental study where flight costs were increased. Males attended the chick more often than females, but females provided more food to the chick than males. Males and females foraged during similar periods of the day, had similar prey types and sizes, diving depths, durations of foraging trips, foraging zones and ranges. Females spent a smaller proportion of the foraging trip sitting on the water and had higher diving rate than males, suggesting higher foraging effort by females. In females, trip duration correlated with mass at departure, suggesting a flexible investment through control by body mass. The experimental study showed that handicapped females and female partners of handicapped males lost mass compared to control birds, whereas there was no difference for males. These results indicate that the larger female is the main provisioner of the chick in the pair, and regulates breeding effort in relation to its own body mass, whereas males have a fixed investment. The different breeding investment between the sexes is associated with contrasting foraging strategies, but no clear niche differentiation was observed. The larger size of the females may be advantageous for provisioning the chick with large quantities of energy and for flexible breeding effort, while the smaller male invests in territory defence and nest guarding, a crucial task when breeding at high densities. In masked boobies, division of labour appears to be maximal during chick rearing—the most energy-demanding period—and may be related to evolution of RSD.     

Parental behaviour of Spanish Imperial Eagles Aquila adalberti: sexual differences in a moderately dimorphic raptor

Capsule The sexes make significantly different contributions in the tasks of nest-building, incubation and food provisioning to chicks.

Aims To determine the division of parental activities during breeding (nest-building, incubation, brooding, food provisioning and feeding).

Methods Between 1991 and 1998 focal observations were made at nests using 20–60× telescopes. During the incubation period, 11 pairs were studied involving a total of 2812 observation hours over 269 days. During the chick-rearing period, seven pairs were studied involving a total of 5499 observation hours spread over 503 days. The birds were sexed and identified individually on the basis of size, position during copulation, vocalization and plumage differences.

Results Significant differences were found between the sexes in parental behaviour. Males scarcely participated in incubation and contributed significantly more than the females only in food provisioning, both during incubation (food transfers to feed the female) and during chick-rearing (to feed the chicks). Females were responsible for most of the incubation, nest-building (during incubation and chick-rearing), brooding, shading and feeding of chicks. Nest attendance by females decreased over time. In both sexes, food provisioning increased with brood size.

Conclusion Intersexual differences are discussed in the context of the reversed sexual size dimorphism and parental investment strategies. We suggest that in Spanish Imperial Eagles reversed sexual size dimorphism is best explained by the prey capture difficulty hypothesis, than by parental role division during reproduction.     

Male chicks are more costly to rear than females in a monogamous seabird, the Common Murre

Feeding rates and mass loss during chick rearing were comparedfor individually marked parents of male and female Common Murre(Uria aalge) chicks at Great Island, Newfoundland, Canada, from1997–2001. Both parents in this socially monogamous seabirdspecies share parental care duties until colony departure, afterwhich the single chick is fed only by its father. Because murresprovision their single chicks with one clearly visible fishper trip, it is possible to accurately determine whether parentsdifferentially feed male and female chicks. Based on slightlygreater mass of males in adulthood, possibly favored by sexdifferences in breeding roles, we predicted that male nestlingswould be fed more than females. Fathers' feeding rate to sons,but not daughters, increased with chick age, whereas maternalfeeding rate increased with chick age for both sexes. When year-correctedfeeding rates of pairs rearing both sexes were compared, bothmothers and fathers fed their sons significantly more than theirdaughters in the later part of the chick-rearing period. Moreover,parents rearing male chicks lost mass at a significantly higherrate than those rearing females. There was no difference infledging age for sons and daughters. These results indicatethat differential parental allocation occurs and has measurablecosts even in a species with only slight adult sexual dimorphism.     

Sex roles in the monogamous Purple Sandpiper Calidris maritima in Svalbard

The Purple Sandpiper Calidris maritima, an Arctic shorebird, displays unusual sex roles during breeding. In a 5-year study in Svalbard, in the Norwegian high Arctic, data were collected on the roles of the sexes in this species. Purple Sandpipers were similar to most shorebirds in that males actively courted females, established territories and vigorously defended their territories from intruders. Both sexes shared incubation duties approximately equally over the entire 21-day incubation period. Males incubated very little initially, but increased their effort significantly during the first 11 days of incubation to over 50% of the time from days 11–21. However, Purple Sandpipers contrasted with most other shorebird species in that females discontinued their breeding efforts at hatching. In nearly all cases, broods were attended solely by the male until the chicks reached fledging age (or even longer). Nonetheless, the pair bond must be described as monogamous because neither males nor females were found to re-mate or lay a second clutch during a season. This pattern of parental care, which is found only in very few other shorebird species, is discussed in an evolutionary context.     

Sex differences in Little Auk Alle alle parental care: transition from biparental to paternal-only care

Understanding differences in male and female care in biparental care systems can help interpret the selective pressures that shape parental strategies. We examined Little Auk Alle alle parental care at a breeding colony during the chick-rearing and fledging periods by conducting observations on marked, known-sex pairs, and by examining the sex ratio of birds carrying food to the colony. Little Auks transitioned from biparental to mostly paternal-only care during late chick-rearing. Males delivered more meals and spent more time at the colony than females during late chick-rearing. Very few females were present at the colony by the end of chick-rearing and through the fledging period, and all marked parents observed accompanying their chick to sea were male. Chick mass loss prior to fledging was associated with the lack of provisioning by the female parent, rather than a reduction in feeding frequency by both parents. The occurrence of paternal-only care during and after fledging is discussed in relation to physiological, ecological and phylogenetic constraints.     

Modulation of prolactin but not corticosterone responses to stress in relation to parental effort in a long-lived bird

We tested the hypothesis that parental effort modulates the magnitude of corticosterone and prolactin responses to stress in a long-lived bird, the Black-legged kittiwake (Rissa tridactyla). To do so, we compared corticosterone and prolactin responses to capture/restraint stress between chick-rearing birds and failed breeders (no parental effort). We predicted that (1) the increase in plasma corticosterone levels in response to stress should be lower in chick-rearing birds, (2) the decrease in plasma prolactin levels in response to stress should be lower in chick-rearing birds, and (3) as both sexes care for the chick, there should be no sex difference in the hormonal response to stress. Baseline plasma corticosterone and prolactin levels were higher in chick-rearing birds and were not influenced by body condition. Failed breeders were in better condition than chick-rearing individuals. Corticosterone response to stress was unaffected by parental effort as both chick-rearing and failed birds exhibited a robust corticosterone increase. Prolactin response to stress was however clearly influenced by parental effort: chick-rearing birds showed a modest 9% prolactin decrease whereas in failed birds prolactin concentrations fell by 41%. Body condition did not influence hormonal responses to stress. When facing stressful condition, breeding kittiwakes attenuate their prolactin response to stress while enhancing their secretion of corticosterone. Increasing corticosterone secretion triggers foraging efforts and diminishes nest attendance whereas an attenuation of prolactin response to stress maintains parental behavior. We suggest that this hormonal mechanism facilitates a flexible time-budget that has been interpreted as a buffer against environmental variability.     

Influence of adult breeding experience on growth and provisioning of Wandering Albatross Diomedea exulans chicks at South Georgia

For most seabirds, reproductive performance improves with age; in albatrosses this is thought not to be so (experience being acquired before starting breeding) but only one study (of chick growth in a single season at one site) has specifically addressed this. We compared the provisioning performance and growth rates of chicks of Wandering Albatrosses Diomedea exulans breeding for the first (IN), second and third (LE) and fourth or more times (EE) on Bird Island, South Georgia in the austral winters of 1996 and 1997. Eggs from EE adults were significantly heavier than the other two categories and these chicks had a greater mass and longer wings up to 160 days of age and longer culmen and tarsus up to 115 days old. However chicks from all categories fledged at the same average mass, size and age. No significant differences between categories in feeding frequency or meal size were detected but experienced adults made shorter long foraging trips and spent more time at the nest than less experienced birds. Adults that remained at the nest gave chicks smaller meals than those that left immediately after feeding the chick. Although provision of smaller but more frequent meals by experienced adults promotes more rapid chick growth, the resulting differences do not persist into the late chick-rearing period. Our results were very similar to those from Iles Crozet in the Indian Ocean, supporting the hypothesis that when Wandering Albatrosses start to breed they are fully competent foragers but that it takes a while, during early chick-rearing, for birds breeding for the first time to adapt to the additional demands of provisioning a chick.     

Sexual differences in reproductive effort: time-activity budgets of monogamous killdeer,Charadrius vociferus

By measuring the time allocated to various reproductive and somatic activities it was possible to look at differences in the strategies of the sexes for killdeer, a monogamous species. Killdeer time-activity budgets supported the general hypothesis that males and females maximize fitness in different ways. Males allocated more time than females to mating activities: mate attraction and territorial defence. However, the parental roles of the sexes were not those predicted for a monogamous species; males allocated more time than females to parental activities throughout a nesting attempt. Males appeared to be less time-constrained, and better able to perform parental activities. Females allocated more time to foraging than did males. It is suggested that the amount of time females allocated to parental activities is limited by the high energy demands of egg production. Further, high nest-failure rates may select for females maintaining good condition for re-nesting. The costs of desertion for killdeer were considerable. Adult removal experiments supported the hypothesis that two parents were necessary for successful reproduction. No individuals managed to raise offspring alone successfully. Lone males, however, were able to hatch chicks while females were not. The need for continuous parental attentiveness may results in selection for biparental care for killdeer.     

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.     

Riverside wren pairs jointly defend their territories against simulated intruders

Duets are a jointly produced signal where two or more individuals coordinate their vocalizations by overlapping or alternating their songs. Duets are used in a wide array of contexts within partnerships, ranging from territory defence to pair bond maintenance. It has been proposed that pairs that coordinate their songs might also better coordinate other activities, including nest building, parental care and defending shared resources. Here, we tested in the riverside wren (Cantorchilus semibadius), a neotropical duetting species that produces highly coordinated duet songs, whether males and females show similar responses to playback. During territorial disputes in songbird species, individuals tend to direct their attention towards same‐sex territorial intruders, but this bias might be less pronounced in duetting species. We performed a dual‐speaker playback experiment to examine how mated individuals respond to speakers broadcasting female‐versus‐male duet contributions. We found that riverside wrens have high levels of converging behaviour by duetting and remaining in close proximity to one another when responding to simulated paired intruders. Males and females spent more than 80% of their time less than 1 m apart while defending their territory. Both individuals in a pair aggressively engaged with both male and female simulated trespassers by approaching equally close and spending equal time near the two speakers. These results suggest that both sexes perceive a paired territorial intrusion as a similar threat and that both partners are highly invested in defending the shared resources. This study is one of the few to demonstrate equal attention and aggression from mated pairs towards simulated same‐sex and opposite‐sex intruders. We suggest that pairs responding together, in close proximity to one another, might be favourable in duetting species when defending the territory because maintaining a close distance between partners facilitates the extreme coordination of their joint territorial signals.     

Responses of naive yellow warblers to a novel nest predator

Naive parental yellow warblers, Dendroica petechia, were tested with a mounted eastern grey squirrel, Sciurus carolinensis, a novel predator on the study site, during the nest-building, egg-laying, incubation and nestling stages of the nesting cycle. Testing parents only once avoided the possibility that the nest defence behaviour of the parents was modified by reinforcement and loss of fear. In 96% of 80 trials, the female parent responded first to the model. Males rarely showed nest defence behaviour. Female nest defence as measured by defensive vocalizations (i.e., ‘chipping’), closest and modal distance of approach to the model, distraction displays, strikes, and close passes or hovers, did not increase significantly over the breeding season. ‘Chipping’ frequency by both sexes tended to increase after the incubation period but not significantly so. Distraction displays increased significantly only between nest-building and nestling stages. Age, clutch or brood size, and nest success were not correlated with the intensity of nest defence. Females defending exposed nests gave more distraction displays than those defending cryptic nests.     

Reproductive investment and parental roles in Sabine's gulls<Emphasis Type="Italic"> Xema sabini</Emphasis>

More than 90% of avian species exhibit biparental care, though parental activities are often shared unequally between the members of a pair. Among gull species (Laridae), males and females generally share parental activities, although there appear to be considerable differences between species in the relative contribution of each sex. This study examined the behaviour of male and female Sabine's gulls (Xema sabini) during the incubation period and immediately post-hatch; particularly the amount of time they each invested in breeding activities. Although considered an aberrant species in aspects of behaviour and biology, the Sabine's gull showed a high reproductive investment by both sexes, as other gull species do. Males fed females prior to egg laying and contributed equally to incubation and chick provisioning, and females contributed equally to nest defence. Overall, there was no difference between the sexes in the extent of their contributions to parental care, although there was considerable individual variation within pairs. Sabine's gulls are Arctic breeders and the extent of their contributions to parental activities could have been influenced by their extreme breeding environment and short breeding season.     

Nest Defence of Nesting Chinstrap Penguins (Pygoscelis antarctica) against Intruders

We tested some predictions of parental investment theory by studying the aggressive behaviour of colonial nesting chinstrap penguins (Pygoscelis antarctica) against human intruders into their nesting territories. We tested for differences in the aggressive behaviour of penguins according to offspring age (eggs vs. chicks), offspring number, nest location in the colonies (central vs. peripheral) and sex. Offspring age was the main factor influencing nest defence, although nest location and sex were also important. Chicks were defended more strongly than eggs, in accordance with changes in the reproductive value of offspring, and this increase in aggressiveness was not related to revisitation of the same individuals. The level of aggression of penguins breeding in central sites was higher than that of peripheral birds, a difference that could be due to the lower residual reproductive value of central-nesting, probably older, birds. The stronger aggressiveness of males could be due to a combination of factors related to sexual selection and life-history traits. Offspring number did not affect the level of nest defence.     

Habitat selection and potential fitness consequences of two early‐successional species with differing life‐history strategies

Habitat selection and its relationship to fitness is a fundamental concept in ecology, but the mechanisms driving this connection are complex and difficult to detect. Despite the difficulties in understanding such intricate relationships, it is imperative that we study habitat selection and its relationship with fitness. We compared habitat selection of least terns (Sternula antillarum) and piping plovers (Charadrius melodus) on the Missouri River (2012–2014) to examine the consequences of those choices on nest and chick survival. We hypothesized that plovers and terns would select habitat that minimized the chance of flooding and predation of eggs, chicks, and adults, but that plovers would also select habitat that would provide foraging habitat for their chicks. We developed an integrated habitat selection model that assessed selection across multiple scales (sandbar and nest scales) and directly modeled the effect of selection on nest and chick survival. In general, the species selected habitat in keeping with our hypotheses, such that predation and flooding, in particular, may have been reduced. Sandbar selection had either a negative or no appreciable effect on nest survival for both species across years. Nest‐site selection in 2012 had a generally positive effect on nest survival and chick survival for both terns and plovers, and this trended toward a negative effect by 2014. This result suggested that early selection decisions appeared to be adaptive, but we speculate that relatively high site fidelity and habitat degradation led to reduced benefit over time. Our results highlight the complex nature of habitat selection and its relationship to fitness.