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We used blood samples of the Blue-footed Booby, considering sex (female and male) and age-class (adult and chick) of individuals at different breeding stages during two breeding seasons (2010-2011 and 2011-2012) in Isla El Rancho, Sinaloa, to determine lead, cadmium, and mercury concentrations. Lead and cadmium concentrations were below our detection limit (0.05 and 0.36ppm, respectively). A higher concentration of mercury was found in early stages of breeding, likely related to changes in mercury environmental availability. Mercury concentrations in adults did not relate with their breeding output. Males and adults had higher mercury concentration than females and chicks. We provide information of temporal, sex and age-related variations in the concentrations of mercury in blood of the Blue-footed Booby.

The evolution of remating and prolonged pair bonds in animals has generally been explained in terms of improved coordination and cooperation between familiar individuals, but costs of mate familiarity have rarely been considered. A possible cost for males is increased risk of losing paternity if familiarity enables females to detect when alternative sires are desirable, evade mate guarding or invest more in infidelity. To test whether this familiarity cost exists, we examined whether extrapair paternity increases with bond length in the socially monogamous blue-footed booby, Sula nebouxii, using microsatellite-based analysis of 384 broods. Extrapair paternity increased from 9.4% of broods in first pairings to 21.6% in second pairings, then declined to 7.5% in third through eighth pairings. On their first remating with a female, males faced enhanced risk of losing paternity, but thereafter the risk was no greater than on first matings. However, after loss of paternity, males were no more likely to divorce. Effects of familiarity on extrapair paternity could influence the evolution and taxonomic distribution of remating and prolonged pair bonding in socially monogamous animals.

Zinc is essential for animal metabolism, but the variation in Zn concentrations within seabird populations has been seldom explored. We collected Blue-footed booby (Sula nebouxii) eggs during 2012 and 2013 and blood samples from nestlings and adults during 2011 and 2012 in Sinaloa, Mexico, to evaluate differences in Zn concentrations among years, ages, sexes, and breeding stages. Zinc concentrations in eggs ranged between 27.3 and 64.9 ppm (dry weight), whereas Zn levels in the blood of nestlings and adults ranged between 16.1 and 53.1 ppm (dry weight). Egg Zn concentrations did not differ due to developmental stage, although Zn concentrations were significantly higher in 2013 than in 2012. Nestling Zn concentrations differed between years and decreased gradually with age. Chicks had significantly higher Zn concentrations than those of adults. Once nestlings stopped growing, females presented higher Zn concentrations than males. Adult Zn concentrations were higher during the pre-laying period than during the other breeding stages. Our results suggest that growth, sex, and breeding stage affect blood Zn concentrations, which may be due to stage-specific requirements. When compared with those of other seabird species, the Zn concentrations reported here are intermediate, and we did not detect adverse effects on either nestlings or adults.

To compete for parental food deliveries nestling birds have evolved diverse behaviors such as begging displays and sibling aggression. Testosterone has been suggested to be an important mechanism orchestrating such competitive behaviors, but evidence is scarce and often indirect. Siblicidal species provide an interesting case in which a clear dominance hierarchy is established and the dominant chicks lethally attack siblings. We experimentally elevated testosterone in chicks of a facultatively siblicidal species, the black-legged kittiwake, Rissa tridactyla, and showed that testosterone-treated chicks were more aggressive toward their sibling than were control chicks. In such facultatively siblicidal species, chicks normally exhibit intense aggression only when threatened by starvation. Indeed, we found that chicks in relatively poorer condition were more aggressive than were chicks in better condition, even among testosterone-treated chicks, suggesting the action of an additional signal modulating aggression. Relatively larger siblings were also more aggressive than were relatively smaller siblings, confirming the importance of size advantage in determining dominance hierarchies within the brood. In addition, testosterone increased aggression toward a simulated predator, indicating that in kittiwakes testosterone can increase aggression in contexts other than siblicide. Testosterone promoted aggression-mediated dominance, which increased begging although testosterone treatment did not have a significant separate effect on begging. Therefore, testosterone production in the kittiwake and most likely other siblicidal species seems an important fitness mediator already early in life, outside the sexual context and not only manifesting itself in aggressive behavior but also in dominance-mediated effects on food solicitation displays toward parents.

We aimed to elucidate the factors that influence the frequency of adoption and infanticide in the Blue-footed Booby (Sula nebouxii), a seabird with biparental chick care and high chick-rearing costs. We obtained data from two colonies during four breeding seasons. In addition, we removed parents from their nests for short periods of time (10 min) and fitted their chicks with polyethylene foam protectors, in order to record chick behavior, particularly interactions with neighboring adults. Adoption and infanticide occurred mostly in zones of high nest density and during years of high breeding success. The chicks adopted were 20–40 days of age, the age at which they acquire mobility and parents still experience a certain inability to recognize them. Adopted chicks were slightly younger than chicks in the broods into which they were adopted. In most cases, adoptees were younger than their siblings in their natal broods. We also recorded short-term (lasting only a few hours) alloparental care for chicks older than 40 days. When we excluded parents from nests and their chicks were attacked, it was female neighbors that physically attacked them, whereas males were more prone to brood or preen them; therefore, the sex with certainty of parenthood was more prone to practice infanticide, presumably to avoid adoption. The lack of aggressiveness by males might be due to the possibility of paternity, given the occurrence of extra-pair copulations in this species.

Meraz, J., Ancona, S., Rodríguez, C., and Drummond, H. 2013. Reproduction of the blue-footed booby predicts commercial fish abundance in the eastern tropical Pacific. – ICES Journal of Marine Science, 70: 1263–1272. To establish whether reproduction in a colony of the blue-footed booby in the eastern tropical Pacific predicts local abundance of prey fish several months later, 13 years of data were analysed. Eight reproductive variables assessed during the period of January–May, grouped in two factors, were related to commercial catches in the surrounding 6600 km2 area during the following June–December. The first factor explained 33% of interannual variance in fish captures per unit effort during June–December (future FCUE), and 63% when only El Niño years were considered. Also, the proportion of large clutches present on three single-day censuses in the spring explained 51, 46 and 35% of variance in future FCUE among all years, and 78, 85 and 82% of variance among El Niño years. In contrast, sea surface temperatures in March did not explain variance in future FCUE. Proportion of large clutches is a moderately good predictor of the abundance of commercial fish during the subsequent seven months and can be satisfactorily and cheaply measured on a single day.

While studies of mate choice based on male color pattern are ubiquitous, studies of mate choice based on ornamental color traits in sexually monomorphic species are less common. We conducted manipulative field experiments on two color ornaments of king penguins (Aptenodytes patagonicus), the size of auricular patches of orange feathers and degree of UV reflectance from beak spots, to determine how the degree of ornamentation influenced pairing rate. In a reduction of auricular patch size, females paired significantly more quickly than males in both control and experimental samples. When this bias was taken into account statistically, pairing of individuals with reduced auricular patches was significantly delayed. We also reduced, but did not eliminate, UV reflectance from beak spots by applying a UV filter; no sex difference in pairing rate was evident in this experiment. Treated birds paired significantly more slowly than untreated control individuals, taking more than a week longer to pair on average than their unmanipulated counterparts, a result that was significant for males and approached significance for females. Our results may indicate mutual mate choice via UV reflectance of the beak spot. Given that this is a species where breeding is extremely slow and considerable investment by both males and females is required for successful reproduction, our results support the hypothesis that in such species, sexual selection might act on the same ornament in both sexes.

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.

The relationships between breeding site location in forest habitat and age, behaviour and reproductive performance of Blue-footed Boobies Sula nebouxii were examined in two different plots on the northeast corner of Isla Isabel, Mexico. Birds nesting closer to the forest edge, where nest density is highest, laid their clutches earlier and fledged more chicks. In plot A, proximity to the forest edge was associated with more wakefulness and wing drooping in chicks and more interactions with other species in female breeders. In plot B, ages of breeders increased closer to the forest edge. The patterns of nest density, laying date and age distribution suggest that, on the northeast corner of Isla Isabel, the boobies prefer to nest close to the forest edge, possibly because the benefits, including easy landing and take-off, are greater than the costs associated with inclement weather, high density of conspecifics and allospecifics, and negative social interactions.

Life-history theory predicts a trade-off between current and future reproduction, such that long-lived species should not increase their reproductive effort (RE) at a cost to their own survival. In species with long-term pair bonds and biparental care, each parent must balance its reproductive investment against that of its partner. Although the effects of "handicapping" studies on the focal individual are sometimes difficult to interpret, they are a powerful approach for investigating compensatory responses of the partner. In the present experiment, we manipulated flying ability of one parent in long-lived Cape gannets (Morus capensis), thereby indirectly increasing the demands on RE of the unmanipulated partner. Handicapped birds doubled their foraging trip duration and reduced nest attendance. Their partners showed behavioral compensation via increased nest attendance for chicks younger than 30 days and increased foraging trip frequency for older chicks. The behavioral responses of partners did not fully compensate for the reduced care of handicapped adults. For manipulated nests, overall foraging trip frequency was 21% lower, chicks were left unattended at 5 days younger, and their growth and survival was reduced compared with control nests. Handicapped adults lost 10% of their body mass during the experiment, but their partners showed no decrease in body mass. Our results show that long-lived Cape gannets can increase current RE when needed, without negative effects on body condition or survival. The reduced care of one parent was partly compensated for by its partner, and remaining costs were borne by the chick. Copyright 2009, Oxford University Press.

To determine whether solicitation by blue-footed booby chicks accurately encodes their need for food, we independently manipulated the body condition and recent food ingestion of singleton chicks and recorded three measures of begging. Variations in the three measures of begging covaried only partially, but in general, chicks begged more intensely when they were in poor body condition and also when suffering recent food deprivation. Effects of body condition and recent food deprivation on begging were broadly additive, although deprived chicks in poor condition failed to beg more intensely than those in good condition. Protracted short-term deprivation may create such a high level of need that the body condition component of need becomes temporarily unimportant. Parents more frequently fed chicks that begged more intensely, chicks in poor condition, and chicks suffering food deprivation. Deprived chicks in poor condition received more food than deprived chicks in normal condition even though they did not beg more intensely, possibly because parents responded not only to current begging but also to begging earlier in the day, or to other cues to body condition. These results support the hypothesis that begging of boobies represents honest signaling of need.

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.

The larger sex is often more vulnerable, in terms of development and survival, to poor conditions during early life. Differential vulnerability has implications for parental investment strategies such as sex ratio theory. When males are larger, it is not possible to separate the effects of larger size per se and other aspects of the male phenotype on vulnerability. Furthermore, offspring competition might favor the larger sex and thereby mask intrinsic, size-related effects. We studied sex-specific mortality in a bird species with reversed size dimorphism, the great skua Stercorarius skua, under natural and experimentally created poor conditions. Small eggs from extended laying sequences were used to create poor early conditions for the offspring, which were raised as singletons. Daughters had a lower survival in all treatment groups. Survival in natural broods was additionally affected by hatch date and position. Hatch weight was not different for sons and daughters but was lower in experimental than in natural nests. In natural nests, daughters fledged 10% heavier than sons, but in experimental nests, they did not reach a higher mass. The average survival difference between sons and daughters was not increased in experimental broods. However, hatch weight had a strong sex-specific effect. Very light females never survived, and survival probability of daughters increased with increasing hatch weight. By contrast, survival of sons over the same range of hatch weights was not related to weight. These findings support the hypothesis that larger (final) size per se is related to sex-specific offspring vulnerability during early life. Copyright 2005.

We report the predation by a coyote (Canis latrans jamesi) on a foraging, adult blue-footed booby (Sula nebouxii) during the early afternoon on 2 December 2009 on the west coast of Isla Tiburon in the Gulf of California, Mexico. The observed event took place while a flock of blue-footed and brown boobies (Sula leucogaster) was plunge-diving close to shore. The food habits of coyotes on Isla Tiburon are not known, and capture of live adult boobies by coyotes has not previously been documented in the literature.

We tested the challenge hypothesis for the hormonal regulation of aggression in chicks of the black-headed gull, Larus ridibundus. Chicks of this species are highly aggressive toward conspecifics, but never to peers that hatched from the same clutch (modal clutch size is three). Therefore, in the first experiment small families were housed together in large groups (challenged condition) and compared to families kept isolated (nonchallenged condition). As expected, in the challenged condition during the initial stage of territory establishment basal levels of testosterone M were clearly higher than those in the nonchallenged condition. In the second experiment we tested the effect of a short social challenge on short-term T-fluctuations. The design was based on an earlier experiment, showing that after temporary T-treatment chicks become very sensitive to social challenges while having low basal T-levels. We now show that these social challenges induce brief elevations in plasma T-levels. These peaks are similar to those in previously untreated chicks but untreated chicks do not respond with aggression to a challenge. Therefore, we conclude that the initial exposure to elevated T-levels increases the sensitivity to brief changes in T induced by social challenges. In this way exposure to T, that may be detrimental for development, is minimized while birds remain able to defend territories. This is the first report showing that the challenge hypothesis as established for adult birds, is also applicable for aggressive behavior in young birds outside the sexual context. Furthermore we suggest that a phase of priming with T is necessary to obtain the high behavioral responsiveness to a challenge. (C) 2002 Elsevier Science (USA).

Animal integumentary coloration plays a crucial role in visual communication and camouflage, and varies extensively among and within species and populations. To understand the pressures underlying such diversity, it is essential to elucidate the mechanisms by which animals have created novel integumentary coloration. Colours can be produced by selective absorption of light by skin pigments, through light scattering by structured or unstructured tissues, or by a combination of pigments and nanostructures. In this review, we highlight our current understanding of the interactions between pigments and structural integumentary tissues and molecules. We analyse the available evidence suggesting that these combined mechanisms are capable of creating colours and optical properties unachievable by either mechanism alone, thereby effectively expanding the animal colour palette. Moreover, structural and pigmentary colour mechanisms frequently interact in unexpected and overlooked ways, suggesting that classification of colours as being of any particular type may be difficult. Finally, we discuss how these mixtures are useful for investigating the largely unknown genetic, developmental and physical processes generating phenotypic diversity. This article is part of the themed issue ‘Animal coloration: production, perception, function and application’.

Thirteen Blue-footed Boobies (Sula nebouxii) born on Isla Isabel, Nayarit, Mexico, nested on Isla El Rancho, Sinaloa. This natal dispersal of about 450 km was unknown for this species. Recruitment of these breeders appeared to happen at between four and seven years of age, slightly higher than the recruitment age reported for Isla Isabel birds that remain there. Age at first detection did not differ between males and females. Immigration to Isla El Rancho could relate to the recent establishment of this colony, and its continuous growth during the last five years. Since there doesn’t seem to be a shortage of nesting sites at Isla Isabel, long distance natal dispersal may be a normal trait in this species of booby.

On islands in the Gulf of California in 2003 and 2004, we recorded three Blue-footed Booby (Sula nebouxii) and one Brown Booby (S. leucogaster) nests that were actively used by two females and one male. Although our data are limited, and cooperation cannot be completely ruled out, we suggest that these nests were the result of bigamous behavior.

Abstract Reversed sexual size dimorphism in avian species (females larger than males) may be an adaptive consequence of different roles of males and females in parental care. We examined the alleged division of labour in two-chick broods of the blue-footed booby, using behavioural observation and frequent weighing of chicks. In the first week of parental care, males and females fed broods at similar frequencies and provided similar masses of food, but females brooded more than males when broods were 5-10 d old. Subsequently, females provided a greater mass of food and frequency of feeds than males until chicks were at least 35 d old (mass) and 60 d old (frequency), while attending the brood for just as much time as males until chicks were at least 35 d old. Males and females did not differ in the tendency to feed (frequency and mass) the first-hatched chick differentially. In nearly all components of parental care examined here, and in other studies, the female's contribution is equal to or greater than the male's. Only in clutch attendance and nest defence does the male contribute more than the female, but his small size seems unlikely to enhance performance in these activities. Overall, small size appears potentially to limit male provisioning of the brood, and is unlikely to be an adaptation for division oflabour in parental care. This result casts doubt on the relevance of the division-of-labour hypothesis for adult size dimorphism.

International audience; 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. ; p. 413-420.