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We describe a new disjunct population of Blue-footed Booby Sula nebouxii at Islote González in the Islas Desventuradas archipelago. Though likely much larger, the population size at the time of our survey was a minimum of 16 breeding pairs. On 14-15 December 2020, two pairs of boobies with nestlings and four pairs with empty nests were found. Likely the species is a recent arrival to the archipelago, perhaps during one of the strong El Niño events between 1970 and 2001 (especially 1982/83, when the species was first recorded in Chile). Blue-footed Booby might also nest on different areas at San Félix Island and some smaller islets around the archipelago, where they would have no nesting-site competition from the larger Masked Booby S. dactylatra. [ABSTRACT FROM AUTHOR]

Blue-footed boobies (Sula nebouxii) are socially monogamous, colonial seabirds exhibiting intra-specific nest parasitism and extra-pair copulations. Prior DNA fingerprinting assays failed to detect extra-pair offspring in the nests of congeners, and the rate of intra-specific nest parasitism has not been estimated using molecular techniques. We describe the development and characterization of 11 microsatellite DNA loci, tested using 31 individuals collected on Isla Isabel, Nayarit, México. The number of alleles per locus ranged from three to 22, averaging seven; total exclusionary power of the microsatellite panel was 0.99; no loci deviated from Hardy-Weinberg equilibrium; and we did not detect linkage disequilibrium following Bonferroni correction. This microsatellite panel will facilitate future studies of nest parasitism and extra-pair paternity in blue-footed boobies.

Populations of blue-footed boobies in the Galápagos Islands have declined by at least 50% compared with numbers recorded during the 1960s. Recently, concern has been expressed about potential adverse effects of introduced blood parasites; in particular, malarial parasites (i.e. Plasmodium sp.), which are frequently pathogenic and can cause mortality and morbidity in birds, and blood parasites of the genus Haemoproteus, which can reduce fitness. We screened blue-footed boobies at six breeding colonies within Galápagos for both genera of parasite. At two sites, we also investigated whether there was any relationship between infection status and physiological condition or breeding success. We found a high prevalence of Haemoproteus spp. in adults and chicks using PCR, although none was found on blood smears. We found no evidence of Plasmodium. Effects of parasitism on the condition of birds were limited. Also, there was no evidence that infection with Haemoproteus adversely affected breeding success or chick growth. Our results suggest that poor success of breeding birds was not strongly associated with Haemoproteus infection. However, regular colony monitoring and examination of dead birds are necessary to assess whether acute Haemoproteus infection might be responsible for some mortality, particularly of chicks. [ABSTRACT FROM AUTHOR]

Parental care in long-lived bird species involves a trade-off between the benefits of increasing the effort expended on current offspring and the costs that this represents for future reproductive output. Under regimes of high environmental variability, long-lived seabirds can adjust their breeding effort to buffer the negative effects of this variability on their offspring. However, the potential impacts of variation in breeding effort on offspring physiology in the short term and on longer-term survival are poorly understood. In this study, we manipulated brood age through a cross-fostering experiment to assess whether increasing or decreasing parental reproductive expenditure led to costs in Blue-footed Booby Sula nebouxii chicks. Specifically, we tested the consequences of altered parental reproductive expenditure on the offspring's physiological condition (plasma metabolites, heterophil to lymphocyte ratio (H/L) and body condition index ( BCI)) and survival. Offspring from broods in which parental investment was experimentally increased showed a lower BCI and lower alkaline phosphatase levels and higher H/L ratios than controls. Conversely, offspring showed the opposite pattern when reproductive expenditure was experimentally decreased. We observed no effects of manipulation of parental investment on triglyceride levels or on survival rates. Although our findings suggest that Blue-footed Booby parents have the ability to adjust their breeding effort according to the demands of their offspring, parental effort could influence the effect of hatching order by suppressing the aggressive tendency of the senior chick. [ABSTRACT FROM AUTHOR]

Individual variation in sexual fidelity and extrapair paternity (EPP) is widely attributed to environmental heterogeneity, but the only variables known to be influential are food abundance and density of conspecific breeders (potential extrapair partners). Habitat structure is thought to impact EPP but is rarely measured and, when considered, is usually confounded with food abundance and predation pressure. To sidestep these confounds, we tested whether EPP is associated with habitat structure variables and with local conspecific density in a species whose nesting habitat is not used for feeding and lacks predators. In a blue-footed booby, Sula nebouxii , colony, the probability of EPP in a female's nest was highest in parts of the study plot where there were few obstacles to locomotion, and was quadratically related to local density of sexually active males, even though local males did not sire the EP chicks. The probability of a male breeder siring EP (extrapair) chicks elsewhere was quadratically related to local density of sexually active males around his nest. From these patterns we infer that both sexes may foray for EP interactions, that males and females nesting at intermediate density are most likely to be accessed by forayers, and that obstacles in the vicinity of a female's nest constrain access of foraying males. To our knowledge, this is the first demonstration that individual variation in EPP is associated with habitat structure in the absence of confounding variation in food availability, predation pressure or breeder quality, and the first evidence that EPP opportunities of female and male breeders are reduced by high density of conspecific breeders above a particular threshold. [ABSTRACT FROM AUTHOR]

The behaviour that mediates divorce and partner change in socially monogamous species is largely unstudied and unknown, although roles and adaptive functions in some birds have been inferred from breeding outcomes, partial behavioural records and captive studies. Here, roles and functions of natural within-season mate switching of a colonial bird were characterized by describing interactions over many days during the whole daylight period. Switching occurred in 5.9% of bluefooted booby pairs and was initiated by desertion of males or, less commonly, females. Three male desertions were consistent with either the Errors of Mate Choice or Incompatibility hypotheses but were better explained by our new Unfaithful Mate hypothesis because infidelity of the mate preceded desertion. Another two male desertions were more consistent with the Better Options hypothesis because the males switched to their ongoing extra-pair (EP) partners. One female desertion was consistent with the Errors of Mate choice or Incompatibility hypotheses, and another with the Better Options hypothesis. All five deserted females switched promptly to their EP partners. Thus, most switches consisted of (1) males replacing a possibly unsatisfactory partner with a new (already identified or to be identified) partner, or (2) deserted females pairing with their EP partners. [ABSTRACT FROM AUTHOR]

In breeding birds, acoustic signalling is often an important cue for reunion between sexual partners. However, in spite of its potential interest, mate recognition has rarely been examined by comparing the two sexes. We studied the blue-footed booby, Sula nebouxii, a socially monogamous seabird, with a dramatic call sexual dimorphism suggesting two different strategies for identity coding: the female call is an amplitude-modulated sound with a harmonic series slowly modulated in frequency, while the male call is a noisy whistle strongly modulated in frequency. To compare acoustic strategies between the sexes, we (1) recorded calls of both males and females, (2) searched for an individual signature and characterized it, and (3) tested, using a playback experiment, whether the calls of males and females were equally efficient for mate recognition. Results showed that an individual signature was present in the calls of both sexes. However, the acoustic parameters involved differed: female individual identification was principally achieved by a spectral analysis of the call whereas males'' identity relied mostly on temporal cues. More than 70% of both females and males tested in playback experiments successfully recognized their mate. This suggests that the coding strategies are equally efficient in terms of individual recognition between mates. From a broader point of view, our results underline the importance of assessing both males and females within the same investigative framework. [Copyright &y& Elsevier]

In evolutionary biology, whether parents should enhance or reduce parental care according to mate ornamentation is a subject of great debate. However, the evolution of female ornaments can shed light on this question. In theory, female ornamentation should be traded off against fecundity and thus cannot be wholly informative to males without a direct indication of fecundity. Hence, direct cues of offspring quality should affect the relationship between male investment and female ornamentation. Under this hypothesis, we manipulated two direct cues of offspring quality (egg size and color) after first egg laying in the blue-footed booby and registered male incubation patterns. In this species, foot color is a dynamic signal of current condition and in females is traded off with egg size. We found that males spent more time incubating when paired with dull females but only in nests with large eggs. Males also spent less time incubating small dull eggs. Results indicate that egg size, a direct cue of reproductive value, affected the relationship between male effort and female ornamentation. Males may be willing to help females that have invested in offspring at the expense of ornamentation, which suggests compensation when females are in low condition. Another possibility is that males relax their effort when paired with highly ornamented and fecund females because they have high parenting abilities. Our findings suggest that the information conveyed by female ornaments may depend on direct cues of fecundity. Results also highlight that parental decisions are complex, modulated by a combination of information sources. [ABSTRACT FROM AUTHOR]

Theories of ageing predict that early reproduction should be associated with accelerated reproductive senescence and reduced longevity. Here, the influence of age of first reproduction on reproductive senescence and lifespan, and consequences for lifetime reproductive success (LRS), were examined using longitudinal reproductive records of male and female blue-footed boobies ( Sula nebouxii) from two cohorts (1989 and 1991). The two sexes showed different relationships between age of first reproduction and rate of senescent decline: the earlier males recruited, the faster they experienced senescence in brood size and breeding success, whereas in females, recruiting age was unrelated to age-specific patterns of reproductive performance. Effects of recruiting age on lifespan, number of reproductive events and LRS were cohort- and/or sex-specific. Late-recruiting males of the 1989 cohort lived longer but performed as well over the lifetime as early recruits, suggesting the existence of a trade-off between early recruitment and long lifespan. In males of the 1991 cohort and females of both cohorts, recruiting age was apparently unrelated to lifespan, but early recruits reproduced more frequently and fledged more chicks over their lifetime than late recruits. Male boobies may be more likely than females to incur long-term costs of early reproduction, such as early reproductive senescence and diminished lifespan, because they probably invest more heavily than females. In the 1991 cohort, which faced the severe environmental challenge of an El Niño event in the first year of life, life-history trade-offs of males may have been masked by effects of individual quality. [ABSTRACT FROM AUTHOR]

We assessed population differentiation and gene flow across the range of the blue-footed booby ( Sula nebouxii) (1) to test the generality of the hypothesis that tropical seabirds exhibit higher levels of population genetic differentiation than their northern temperate counterparts, and (2) to determine if specialization to cold-water upwelling systems increases dispersal, and thus gene flow, in blue-footed boobies compared with other tropical sulids. Work was carried out on islands in the eastern tropical Pacific Ocean from Mexico to northern Peru. We collected samples from 173 juvenile blue-footed boobies from nine colonies spanning their breeding distribution and used molecular markers (540 base pairs of the mitochondrial control region and seven microsatellite loci) to estimate population genetic differentiation and gene flow. Our analyses included classic population genetic estimation of pairwise population differentiation, population growth, isolation by distance, associations between haplotypes and geographic locations, and analysis of molecular variance, as well as Bayesian analyses of gene flow and population differentiation. We compared our results with those for other tropical seabirds that are not specialized to cold-water upwellings, including brown ( Sula leucogaster), red-footed ( S. sula) and masked ( S. dactylatra) boobies. Blue-footed boobies exhibited weak global population differentiation at both mitochondrial and nuclear loci compared with all other tropical sulids. We found evidence of high levels of gene flow between colonies within Mexico and between colonies within the southern portion of the range, but reduced gene flow between these regions. We also found evidence for population growth, isolation by distance and weak phylogeographic structure. Tropical seabirds can exhibit weak genetic differentiation across large geographic distances, and blue-footed boobies exhibit the weakest population differentiation of any tropical sulid studied thus far. The weak population genetic structure that we detected in blue-footed boobies may be caused by increased dispersal, and subsequently increased gene flow, compared with other sulids. Increased dispersal by blue-footed boobies may be the result of the selective pressures associated with cold-water upwelling systems, to which blue-footed boobies appear specialized. Consideration of foraging environment may be particularly important in future studies of marine biogeography. [ABSTRACT FROM AUTHOR]

vian eggshell color is remarkably variable among and within species and its possible function has long been controversial. Female birds use biliverdin pigment to color their eggs blue and green. Although evidence is accumulating that blue-green eggshell coloration is costly to produce, the exact mechanisms underlying its expression are little studied. Biliverdin pigment is involved in important maintenance functions such as immune and antioxidant defenses and shows similar properties to carotenoid pigments. Carotenoids play a role as immunoenhancers and have the potential to ameliorate the impact of oxidative processes, although their antioxidant function could be minor for some bird species. Important life-history components can be limited by carotenoid availability, including sexual displays and fecundity. Here we explored if biliverdin-based eggshell pigmentation was affected by carotenoid availability in a seabird, by performing a carotenoid supplementation experiment in female blue-footed boobies ( Sula nebouxii) after laying their first egg. In this species, blue eggshell color is associated with incubation patterns, and carotenoid availability affects the crucial trade-off between female foot ornamentation and fecundity. We found that under natural conditions, there was a decline in eggshell color with laying order, suggesting pigment limitation for females. However, carotenoid-supplemented females had enhanced second egg coloration compared to controls, and the color increase was independent of their previous pigment allocation to the first egg. Our results suggest that biliverdin-based eggshell coloration is costly to produce and can be alleviated by current carotenoid availability. [ABSTRACT FROM AUTHOR]

In marine ecosystems climatic fluctuation and other physical variables greatly influence population dynamics, but differential effects of physical variables on the demographic parameters of the two sexes and different age classes are largely unexplored. We analyzed the effects of climate on the survival and recruitment of both sexes and several age classes of a long-lived tropical seabird, the Blue-footed Booby (Sula nebouxii), using long-term observations on marked individuals. Results demonstrated a complex interaction between yearly fluctuations in climate (both local and global indexes, during both winter and breeding season) and the sex and age of individuals. Youngest birds' survival and. recruitment were commonly affected by local climate, whereas oldest birds' parameters tended to be constant and less influenced by environmental variables. These results confirm the theoretical prediction that sex- and age-related variation in life-history demographic traits is greater under poor environmental conditions, and they highlight the importance of including variability in fitness components in demographic and evolutionary models. Males and females showed similar variation in survival but different recruitment patterns, in relation to both age and the spatial scale of climatic influence (local or global). Results indicate different life-history tactics for each sex and different ages, with birds likely trying to maximize their fitness by responding to the environmental contingencies of each year. [ABSTRACT FROM AUTHOR]

Senescence could depress prenatal and postnatal capacities of mothers to invest in offspring. Longitudinal observations on the blue-footed booby (Sula nebouxii) revealed a quadratic effect of female age on fledgling production and cohort differences in rate of reproductive decline. By swapping clutches between females of different ages, we tested whether reproductive senescence is due to decline in egg quality or capacity to care. As laying mothers aged, egg size, ulna length of 5-day-old chicks, and ulna growth of second chicks up to age 30 days declined, and as rearing mothers aged, ulna growth and cellular mediated immune response of second chicks diminished. Oddly, senescent females (>11 years) produced more fledglings when rearing offspring of middle-aged females (8-11 years) than when rearing offspring of senescent or young females. Thus, senescence reduced egg quality and rearing capacities, and reproductive success of senescent mothers depended on prenatal effects associated with the age of the laying mother. Reproductive senescence of boobies may involve constraints on resources allocated to reproduction as well as adaptive adjustment of provision and care according to offspring value, implying that negative effects of senescence on offspring survival can be ameliorated by plasticity in postlaying or postnatal care. [ABSTRACT FROM AUTHOR]

Contradictory patterns of density-dependent animal dispersal can potentially be reconciled by integrating the conspecific attraction hypothesis with the traditional competition hypothesis. We propose a hypothesis that predicts a U-shaped relationship between density and both natal and breeding dispersal distance. Using 10 years of observations on a breeding colony of the Blue-footed Booby (Sula nebouxii), the hypothesis was confirmed by documenting simultaneous positive and negative density-dependent dispersal distances in natal and breeding dispersal of males and breeding dispersal of females within the colony. Point-pattern analyses demonstrated that the breeding sites of Blue-looted Boobies were highly aggregated in all years within a large study area. and aggregation presumably resulted in heterogeneity in patch density throughout the colony. As predicted, at moderate to high densities, dispersal distances showed positive density dependence, with individuals moving to lower density patches. In contrast, at low to moderate densities, dispersal distances showed negative density dependence, with individuals moving to higher density patches. In both sexes of the 1994 cohort, the higher the mean density in patches used by an individual over the long term (up to age I 1 years), the fewer fledglings it produced. A positive effect of density on long- term reproductive success was not detected, possibly because birds that failed during pair formation or incubation were not sampled. Density of conspecifics may be an important influence on habitat selection of breeders, and dispersal may tend to carry individuals to patches where pair formation opportunities are better and negative effects of competition on reproductive success are reduced. [ABSTRACT FROM AUTHOR]

Females are expected to vary investment in offspring according to variables that may influence the offspring fitness in a way that optimises her inclusive fitness for a particular context. Thus, when sexual ornaments signal the quality of the male, females might invest in reproduction as a function of the attractiveness of their mate. We tested whether breeding conditions and male feet colour influence reproductive decisions of blue-footed booby females. In the blue-footed booby, male feet colour is a dynamic condition-dependent sexually selected trait that is related to paternal effort. During two consecutive years, an El Niño year (poor breeding conditions) and a year with good breeding conditions, we experimentally reduced male attractiveness by modifying their feet colour after the first egg was laid and recorded female investment in the second egg. We found that, relative to the first egg in the clutch, females laid heavier second eggs during the poor year than during the good year. Females paired with males with duller feet colour reduced second-egg mass and volume and delayed the laying of the second egg, independently of the year. Absolute yolk androstenedione (A4) concentration (but not testosterone, T) in second eggs was higher during a poor year than during a good year. Only during a year with poor breeding conditions, females paired with experimental males decreased the relative A4 concentration (but not T) in the second egg compared to control females. Thus, blue-footed booby females probably favour brood reduction by decreasing egg quality and increasing size asymmetry between chicks when the breeding and the mate conditions are poor. [ABSTRACT FROM AUTHOR]

Theoreticians propose that trained winning and losing are important processes in creating linear animal dominance hierarchies, and experiments have shown that both processes can occur in animals, but their actual roles in creating natural hierarchies are unknown. We described agonism in 18 broods of three blue-footed boobies, Sula nebouxii, a species for which trained winning and losing have been demonstrated, to infer how these processes generate and maintain a natural hierarchy. Ranks in the linear hierarchy that emerged in every brood were initially assigned by asymmetries in age, size and maturity, which led to differences between broodmates in levels of expressed and received aggression and, consequently, to differences in the training of their aggressiveness and submissiveness. Later, ranks appeared to be maintained by the chicks'' acquired aggressive and submissive tendencies combined with ongoing effects of persisting differences in size and maturity. Our results suggest that trained winning and trained losing are important in the construction of booby hierarchies but that these two axes of learning are largely independent. Increase in submissiveness occurs over a period of about 10–20 days, and the level of submissiveness reached varies with the amount of aggression received. After training, submissiveness is apparently maintained by a lower level of aggression and increasing use of threats. Threats become increasingly effective as chicks age, but are never as effective as attacks. [Copyright &y& Elsevier]

Females may maximize their lifetime reproductive output by adjusting their investment in each breeding event to the perceived likelihood of success. The Blue-footed Booby (Sula nebouxii) is a long-lived seabird with facultative siblicide. We examined whether there is differential resource allocation to eggs with laying order and whether greater egg mass increases hatching probability, chick survival, hatching interval, and mass and size at hatching. We found that the relative investment in first and second eggs decreased as the season advanced: second eggs were slightly (1.5%) heavier than first eggs in early clutches; by contrast, first eggs were 2% heavier than second eggs in late clutches. Accordingly, hatching probability increased with laying date for first eggs and decreased for second eggs. The mass of the egg increased hatching probability; and no effect on chick survival was detected. Laying interval increased after a heavier egg was laid, and heavier eggs produced heavier hatchlings. Hatching intervals were positively related to laying intervals, but egg mass was unrelated to the length of the incubation period and the hatching interval. Our results suggest that egg mass influences embryo survival and that Blue-footed Booby females may adaptively allocate resources to eggs of different laying order according to breeding conditions. [ABSTRACT FROM AUTHOR]

We examined the dynamics and avoidance of mate guarding, by males and females, in the blue-footed booby, in which the two social mates are usually simultaneously present on the territory but each of them is unmonitored by the other for one-quarter of its time. Both sexes were promiscuous and liable to switch mates. Cuckolded individuals did not increase their overall presence on the territory, but in response to the extra-pair (EP) courtships of their mates, both sexes doubled their rate of intra-pair (IP) courtship and sometimes showed aggression. The male or female's presence depressed the social mate's EP activity, but intra-pair courtship had no such effect, tending even to propitiate that EP activity. Similarly, when females responded to their social mates' EP courtship with approach or aggression, disruption of EP activity was short-lived. Promiscuous females modified their diurnal pattern of attendance, as if attempting to sidestep monitoring by their mates, but cuckolded males matched the modification. Both sexes tended to perform their EP activities at a distance when their mates were present, possibly to evade monitoring or disruption by their mates. Male and female boobies cannot monitor their mates continuously, they do little to facultatively adjust their presence on territory to the risk of infidelity, and their immediate responses to overt infidelity have only the briefest impact; but the information they acquire while monitoring their mates may be critical to constraining their mates' infidelity and also to calibrating their own reproductive investment. [ABSTRACT FROM AUTHOR]

Sexual selection models suggest that female choice is based on male traits that indicate their genetic or environmental condition, consequently enhancing her reproductive success by direct or indirect benefits. We investigated the relationship between male foot colour and offspring condition in the blue-footed booby, a socially monogamous seabird, with conspicuous and variably condition-dependent coloured feet that are selected by females. In a cross-fostering experiment, we found that chick condition was related to the foot colour of the foster father and, to some extent, to the foot colour of the genetic father; thus overall, the father’s sexual ornamentation (genetic and foster) explained 32% of variance of chick condition. These data suggest that foot colour, a dynamic sexually selected trait, is mostly a signal of parental contribution. In species in which males provide parental care, females may choose mates with higher parental ability. Overall, our data suggest that colourful integuments are honest signals of parental ability. [ABSTRACT FROM AUTHOR]

Bright colours of male birds have often been shown to be the target of sexual selection through female choice, yet few studies have looked at the role of colour expressed after pairing on female motivation and behaviour. Here we analyse the role of an integumentary colour in the spectral range of 400–700 nm, the foot colour in male blue-footed boobies, Sula nebouxii, which is prominently displayed during pair courtship. Measurements early in the breeding season showed that foot colour of courting males is pale (high values of brightness) and has an aqua-blue chroma, and females in better body condition were mated to males with brighter feet. We carried out an experimental manipulation which modified the foot colour of males in courtship, making it closer to the foot colour of males in low nutritional state. We found that females paired to experimental males courted less and were less likely to copulate than females in the control group. Male behaviour was apparently unaffected by the manipulation; thus the change in female behaviour can be attributed exclusively to foot colour manipulation. These results strongly support the hypothesis of female preference for an integumentary colour and suggest that this dynamic trait is used as a male ornament after pairing. [ABSTRACT FROM AUTHOR]

Thirteen pairs of blue-footed boobies ( Sula nebouxii) were observed on their colony. Pairs courted frequently and, on average, copulated 24 times during the 30 days before laying, with 38% of those copulations occurring in the last 5 days (presumed fertile period). Males and females increased attendance at the nest site as laying approached. Seven females performed an average seven extra-pair copulations, with 1–2 paired male neighbors, but these were less concentrated in the presumed fertile period than within-pair copulations, and the last two copulations of all 7 females were with their social mates. Rates of female extra-pair copulations were six times lower when their social mate was present, and during the presumed fertile period, no female performed an extra-pair copulation in the presence of her mate. Males did not respond to infidelity of social mates with aggression, prompt copulation, retaliatory copulation, or increase in copulation. Seven of 13 males performed an average of five extra-pair copulations, with 1–3 paired female neighbors, before their own mates began egg-laying. The males' extra-pair copulations represented only 4% of their total copulations during their own mates' presumed fertile periods. Females, the larger sex, apparently control sexual access and copulate with extra males to achieve extra-pair fertilization. Males pursue a mixed strategy: they copulate with extra females, mostly outside their own mate's presumed fertile period, and they copulate increasingly with their social mate as laying approaches, probably assuring some paternity by mate guarding, involving attendance and courtship. Behavior of males and females is also consistent with other hypotheses for extensive joint nest site attendance: pairbonding, copulation access, and territory acquisition. [ABSTRACT FROM AUTHOR]