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Parental overproduction is hypothesized to hedge against uncertainty over food availability and stochastic death of offspring and to improve brood fitness. Understanding the evolution of overproduction requires quantifying its benefits to parents across a wide range of ecological conditions, which has rarely been done. Using a multiple hypotheses approach and 30 years of data, we evaluated the benefits of overproduction in the Blue-footed booby, a seabird that lays up to three eggs asynchronously, resulting in an aggressive brood hierarchy that facilitates the death of last-hatched chicks under low food abundance. Results support the resource-tracking hypothesis, as low prey abundance (estimated from sea surface temperature and chlorophyll-a concentration) led to rapid brood reduction. The insurance hypothesis was supported in broods of three, where last-hatched chicks' survival increased after a sibling's death. Conversely, in broods of two, results suggested that parents abandoned last-hatched chicks following first-hatched chicks' deaths. No direct evidence supported the facilitation hypothesis: the presence of a last-hatched chick during development did not enhance its sibling's fitness in the short or long term. The value of last-hatched offspring to parents, as "extra" or "insurance" varied with indices of food abundance, brood size, and parental age. Ninety percent of overproduction benefits came from enabling parents to capitalize on favorable conditions by fledging additional offspring. Our study provides insight into the forces driving overproduction, explaining the adaptiveness of this apparently wasteful behavior and allowing us to better predict how overproduction's benefits might be modified by ocean warming., Competing Interests: All authors declare to have no competing interests., (© The Author(s) 2024. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology.)

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

Weather conditions can profoundly affect avian reproduction. It is known that weather conditions prior to and after the onset of reproduction can affect the breeding success of birds. However, little is known about how seasonal weather variability can affect birds' breeding performance, particularly for species with a slow pace of life. Long-term studies are key to understanding how weather variability can affect a population's dynamics, especially when extreme weather events are expected to increase with climate change. Using a 32-year population study of the Blue-footed booby (Sula nebouxii) in Mexico, we show that seasonal variation in weather conditions, predominantly during the incubation stage, affects offspring survival and body condition at independence. During most of the incubation period, warm sea surface temperatures were correlated with low hatching success, while rainfall in the middle of the incubation stage was correlated with high fledging success. In addition, chicks from nests that experienced warm sea surface temperatures from the pre-laying stage to near-fledging had lower body condition at 70 days of age. Finally, we show that variable annual SST conditions before and during the incubation stage can impair breeding performance. Our results provide insight into how seasonal and interannual weather variation during key reproductive stages can affect hatching success, fledging success, and fledgling body condition in a long-lived neotropical seabird., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Haemosporidian parasites are common in birds but are seldom reported in seabirds. The absence of vectors or genetic resistance to infection have been proposed to explain this pattern. However, screening of blood parasites in many seabirds has been done only by visual inspection of blood smears, which can miss low-intensity infections, and molecular detection of blood parasites must be supported by detection in blood smears to confirm the presence of haemosporidians and avoid false positive cases. Here, we tested for the presence of blood parasites of the genera Plasmodium, Haemoproteus and Leucocytozoon, combining inspection of blood smears and PCR-based detection methods in a highly philopatric colony of blue-footed boobies (Sula nebouxii) in the Tropical North Pacific. Our results indicate that adults in this colony are likely free of these blood parasites, probably due to unsuitable conditions for insect vectors in booby breeding sites, although potential genetic resistance of blue-footed boobies to infection deserves examination. Apparent absence of blood parasites in Isla Isabel boobies indirectly adds to the growing evidence of variation in parasite infections among avian host species that coexist locally., (© 2022. The Author(s).)

Females and males often exhibit different survival in nature, and it has been hypothesized that sex chromosomes may play a role in driving differential survival rates. For instance, the Y chromosome in mammals and the W chromosome in birds are often degenerated, with reduced numbers of genes, and loss of the Y chromosome in old men is associated with shorter life expectancy. However, mosaic loss of sex chromosomes has not been investigated in any non-human species. Here, we tested whether mosaic loss of the W chromosome (LOW) occurs with ageing in wild birds as a natural consequence of cellular senescence. Using loci-specific PCR and a target sequencing approach we estimated LOW in both young and adult individuals of two long-lived bird species and showed that the copy number of W chromosomes remains constant across age groups. Our results suggest that LOW is not a consequence of cellular ageing in birds. We concluded that the inheritance of the W chromosome in birds, unlike the Y chromosome in mammals, is more stable.

Assortative mating by telomere lengths has been observed in several bird species, and in some cases may increase fitness of individuals. Here we examined the relationship between telomere lengths of Blue-footed Booby (Sula nebouxii) mates, long-lived colonial seabirds with high annual divorce rates. We tested the hypothesis that interactions between maternal and paternal telomere lengths affect offspring and parental survival. We found that relative telomere lengths (RTL) were strongly positively correlated between members of a breeding pair. In addition, RTL of both parents interacted to predict fledgling recruitment, although fledglings with two very long-RTL parents performed only averagely. Telomere lengths also predicted adult survival: birds with long telomeres were more likely to survive, but birds whose mate had long telomeres were less likely to survive. Thus, having long telomeres benefits survival, while choosing a mate with long telomeres benefits reproductive output while penalizing survival. These patterns demonstrate that while a breeder's RTL predicts offspring quality, assortative mating by RTL does not enhance fitness, and a trade-off between different components of fitness may govern patterns of assortative mating by telomere length. They also illustrate how testing the adaptive value of only one parent's telomere length on either survival or reproductive success alone may provide equivocal results.

In this study, we explored chemosensory, ingestive and prey-catching responses of neonate Mexican Black-bellied Gartersnakes ( Thamnophis melanogaster ) to crayfish ( Cambarellus montezumae ). By comparing snakes from a recently discovered crayfish-eating population and a typical non-crayfish-eating population, we asked which behavioral components change as a species enlarges its feeding niche. In the crayfish-eating population chemosensory responsiveness to crayfish was not enhanced but its heritability was higher. Neonates of both populations showed similar preference for freshly-molted versus unmolted crayfish, and whereas the tendency to ingest both crayfish stages remained stable between ages 15 and 90 days in the non-crayfish-eating population, in the crayfish-eating population it actually decreased. Techniques to catch and manipulate molted crayfish were similar in the two populations. We discuss the possibility that there is no increase in the behavioral response to eat crayfish by the neonates of the crayfish-eating populations, possibly due to the absence of ecological and spatial isolation between the two T. melanogaster populations. The crayfish ingestion in some population of T. melanogaster can be explained by environmental differences between populations, or by recent origin of crayfish ingestion in T. melanogaster ., Competing Interests: The authors declare there are no competing interests., (©2020 Manjarrez et al.)

Understanding and modeling population change is urgently needed to predict effects of climate change on biodiversity. High trophic-level organisms are influenced by fluctuations of prey quality and abundance, which themselves may depend on climate oscillations. Modeling effects of such fluctuations is challenging because prey populations may vary with multiple climate oscillations occurring at different time scales. The analysis of a 28-yr time series of capture-recapture data of a tropical seabird, the Nazca Booby (Sula granti), in the Galápagos, Ecuador, allowed us to test for demographic effects of two major ocean oscillations occurring at distinct time-scales: the inter-annual El Niño Southern Oscillation (ENSO) and inter-decadal oscillations. As expected for a tropical seabird, survival of fledgling birds was highly affected by extreme ENSO events; by contrast, neither recruitment nor breeding participation were affected by either ENSO or decadal oscillations. More interesting, adult survival, a demographic trait that canalizes response to environmental variations, was unaffected by inter-annual ENSO oscillations yet was shaped by the Pacific Decadal Oscillation and small pelagic fish regime. Adult survival decreased during oceanic conditions associated with higher breeding success, an association probably mediated in this species by costs of reproduction that reduce survival when breeding attempts end later. To our knowledge, this is the first study suggesting that survival of a vertebrate can be vulnerable to a natural multidecadal oscillation., (© 2018 by the Ecological Society of America.)

In wild long-lived animals, analysis of impacts of stressful natal conditions on adult performance has rarely embraced the entire age span, and the possibility that costs are expressed late in life has seldom been examined. Using 26 years of data from 8541 fledglings and 1310 adults of the blue-footed booby ( Sula nebouxii ), a marine bird that can live up to 23 years, we tested whether experiencing the warm waters and food scarcity associated with El Niño in the natal year reduces recruitment or survival over the adult lifetime. Warm water in the natal year reduced the probability of recruiting; each additional degree (°C) of water temperature meant a reduction of roughly 50% in fledglings' probability of returning to the natal colony as breeders. Warm water in the current year impacted adult survival, with greater effect at the oldest ages than during early adulthood. However, warm water in the natal year did not affect survival at any age over the adult lifespan. A previous study showed that early recruitment and widely spaced breeding allow boobies that experience warm waters in the natal year to achieve normal fledgling production over the first 10 years; our results now show that this reproductive effort incurs no survival penalty, not even late in life. This pattern is additional evidence of buffering against stressful natal conditions via life-history adjustments., Competing Interests: We have no competing interests.

Morphological convergence is expected when organisms which differ in phenotype experience similar functional demands, which lead to similar associations between resource utilization and performance. To consume prey with hard exoskeletons, snakes require either specialized head morphology, or to deal with them when they are vulnerable, for example, during molting. Such attributes may in turn reduce the efficiency with which they prey on soft-bodied, slippery animals such as fish. Snakes which consume a range of prey may present intermediate morphology, such as that of Thamnophiine (Natricinae), which may be classified morphometrically across the soft-hard prey dietary boundary. In this study, we compared the dentition and head structure of populations of Thamnophis melanogaster that have entered the arthropod-crustacean (crayfish)-eating niche and those that have not, and tested for convergence between the former and two distantly related crayfish specialists of the genus Regina ( R . septemvittata and R. grahamii ). As a control, we included the congener T. eques . Multivariate analysis of jaw length, head length, head width, and number of maxillary teeth yielded three significant canonical variables that together explained 98.8% of the variance in the size-corrected morphological data. The first canonical variable significantly discriminated between the three species. The results show that head dimensions and number of teeth of the two Regina species are more similar to those of crayfish-eating T. melanogaster than to non-crayfish-eating snakes or of T. eques . It is unclear how particular head proportions or teeth number facilitates capture of crayfish, but our results and the rarity of soft crayfish ingestion by T. melanogaster may reflect the novelty of this niche expansion, and are consistent with the hypothesis that some populations of T. melanogaster have converged in their head morphology with the two soft crayfish-eating Regina species, although we cannot rule out the possibility of a morphological pre-adaptation to ingest crayfish.

Explaining the remarkable variation in socially monogamous females' extrapair (EP) behaviour revealed by decades of molecular paternity testing remains an important challenge. One hypothesis proposes that restrictive environmental conditions (e.g. extreme weather, food scarcity) limit females' resources and increase EP behaviour costs, forcing females to reduce EP reproductive behaviours. For the first time, we tested this hypothesis by directly quantifying within-pair and EP behaviours rather than inferring behaviour from paternity. We evaluated whether warmer sea surface temperatures depress total pre-laying reproductive behaviours, and particularly EP behaviours, in socially paired female blue-footed boobies (Sula nebouxii). Warm waters in the Eastern Pacific are associated with El Niño Southern Oscillation and lead to decreased food availability and reproductive success in this and other marine predators. With warmer waters, females decreased their neighbourhood attendance, total copulation frequency and laying probability, suggesting that they contend with restricted resources by prioritizing self-maintenance and committing less to reproduction, sometimes abandoning the attempt altogether. Females were also less likely to participate in EP courtship and copulations, but when they did, rates of these behaviours were unaffected by water temperature. Females' neighbourhood attendance, total copulation frequency and EP courtship probability responded to temperature differences at the between-season scale, and neighbourhood attendance and EP copulation probability were affected by within-season fluctuations. Path analysis indicated that decreased EP participation was not attributable to reduced female time available for EP activities. Together, our results suggest that immediate time and energy constraints were not the main factors limiting females' infidelity. Our study shows that El Niño conditions depress female boobies' EP participation and total reproductive activity. In addition to increasing general self-maintenance and reproductive costs, warm waters may increase costs specific to EP behaviours including divorce, reduced male parental care, or pathogen exposure. Our results suggest that female boobies strategically refrained from EP behaviours to avoid these or other longer-term costs, rather than being compelled by immediate constraints. This study demonstrates that current environmental conditions affect females' mating decisions, contributing to variation in EP behaviours, even in a long-lived, iteroparous species that can buffer against temporary adversity., (© 2016 The Authors. Journal of Animal Ecology © 2016 British Ecological Society.)

The recent trend for journals to require open access to primary data included in publications has been embraced by many biologists, but has caused apprehension amongst researchers engaged in long-term ecological and evolutionary studies. A worldwide survey of 73 principal investigators (Pls) with long-term studies revealed positive attitudes towards sharing data with the agreement or involvement of the PI, and 93% of PIs have historically shared data. Only 8% were in favor of uncontrolled, open access to primary data while 63% expressed serious concern. We present here their viewpoint on an issue that can have non-trivial scientific consequences. We discuss potential costs of public data archiving and provide possible solutions to meet the needs of journals and researchers., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

When animals potentially occupy diverse microhabitats, how can camouflage be achieved? Here we combine descriptive and experimental methods to uncover a novel form of phenotypic plasticity in the camouflage of bird eggs that may be present in other avian taxa. Soil from the bare substrate adheres to the blue-footed booby's (Sula nebouxii's) pale eggs, which parents manipulate both under and on top of their webs. Analysis of digital images confirmed that dirtiness increases progressively during the first 16 days of the incubation period, making eggs more similar to the nest substrate. Observations of 3,668 single-egg clutches showed that the probability of egg loss declines progressively over the same time frame and then remains low for the rest of the 41-day incubation period. An experiment showed that when chicken eggs are soiled and exposed in artificial booby nests, they are less likely to be taken by Heermann's gulls (Larus heermanni) than clean eggs.

It is widely expected that the quality of offspring will vary with the age of their parents and that this variation should influence animals' choice of mates. However, theoretical predictions for age effects are contradictory and, to our knowledge, we do not know for any wild animal how the quality of offspring is affected by both parents' ages across their lifespans, or whether mothers' and fathers' ages interact. We tackled this question using long-term data on a highly philopatric, insular population of the blue-footed booby (Sula nebouxii). In this species extra-pair paternity is most common in pairs of two young parents or two old parents, implying that these age combinations might prejudice offspring quality. Analysis of the viability of 3,361 offspring of parents up to 21 years old revealed that fledglings with two young parents or two old parents were least likely to become breeders, whereas fledglings with one young parent and one old parent were most likely to do so. For young parents of either sex, offspring viability increased with age of the other parent; for very old parents, it decreased. These effects could be mediated by parents flexibly modifying their investment in offspring in response to their own and their partners´ ages, but evidence for this was lacking. In 5,343 breeding attempts, although mothers' and fathers' ages independently affected four heavily care-dependent breeding traits at the clutch and nestling stages, their interaction did not affect any trait. The effects of parental age combinations on viability could also be mediated by genes: fledglings with one young parent and one old parent could benefit from greater heterozygosity or better genes.

Prolonged pair bonds have the potential to improve reproductive performance of socially monogamous animals by increasing pair familiarity and enhancing coordination and cooperation between pair members. However, this has proved very difficult to test robustly because of important confounds such as age and reproductive experience. Here, we address limitations of previous studies and provide a rigorous test of the mate familiarity effect in the socially monogamous blue-footed booby, Sula nebouxii, a long-lived marine bird with a high divorce rate. Taking advantage of a natural disassociation between age and pair bond duration in this species, and applying a novel analytical approach to a 24 year database, we found that those pairs which have been together for longer establish their clutches five weeks earlier in the season, hatch more of their eggs and produce 35% more fledglings, regardless of age and reproductive experience. Our results demonstrate that pair bond duration increases individual fitness and further suggest that synergistic effects between a male and female's behaviour are likely to be involved in generating a mate familiarity effect. These findings help to explain the age- and experience-independent benefits of remating and their role in life-history evolution., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

Food shortage and other challenges associated with El Niño Southern Oscillation (ENSO) experienced early in life may have long-term impacts on life history traits, but these potential impacts remain virtually unexplored. By monitoring 2556 blue-footed boobies from 11 cohorts, we showed that birds facing warm water ENSO conditions (and probably low food availability) in the natal year were underweight at fledging, recruited earlier and bred less frequently, but showed no deficit in longevity or breeding success over the first 10 years. Life history impacts of ENSO were substantial when experienced in the prenatal year, the natal year, or the second year of life, and absent when experienced in the third year of life, implying that harsh conditions have greater effects when experienced earlier in life. Sexual differences in impacts depended on the age when warm water conditions were experienced: pre-natal and natal experience, respectively, induced early recruitment and influenced the relationship between age and laying date only in females, whereas second year experience reduced total breeding success only of males. Most surprising were positive transgenerational impacts in females: daughters of females that experienced ENSO conditions in their natal year showed improved breeding success. Developmental plasticity of boobies thus enables them to largely neutralize potential long-term impacts of harsh climatic conditions experienced early in life.

Despite frequent suggestions that dominance-subordination relationships in infancy can affect subsequent agonistic potential during adult life, to our knowledge no explicit test has been made. Experiments have shown that adverse conditions during early development can have long-term effects on a variety of traits ranging from growth to competitive behaviour. In many vertebrate species, the main social setting in which the infant develops is a sibling group where competition is often mediated by a dominance hierarchy. Here, we show in a long-lived marine bird that subordination to an aggressive sibling throughout infancy does not compromise aggressiveness years later during adult life. Former junior and senior chicks of the blue-footed booby, whose typical brood of two chicks exhibits a consistent dominance-subordination relationship with strong 'trained winner' and 'trained loser' conditioning effects, did not differ in their aggressiveness while defending their nest against a conspecific intruder stimulus. Our results suggest that aggressive subordination and associated food deprivation, poor growth and elevated stress hormone during infancy do not prejudice aggressiveness of adult boobies during at least the first 13 years of life. Development of important traits such as aggressive tendencies may be buffered against the normal and predictable challenges of infancy.

A survey of the helminth fauna of the blue-footed booby, Sula nebouxii, on Isla Isabel, off the Pacific coast of México, is presented. Eight parasite species were found: 4 digeneans (Galactosomum puffini , Mesostephanus microbursa, Opisthometra planicollis, and Renicola thapari), 3 nematodes (Contracaecum sp., Porrocaecum sp., and Tetrameres sp.), and 1 cestode (Tetrabothrius sp). All these species are reported for the first time in the blue-footed booby, and they represent 8 new locality records as well. Species accumulation curves suggest these 8 parasite species comprise the total helminth fauna of this population of boobies. The most frequent and abundant parasite was R. thapari, inhabiting the kidney of its host. In male boobies, a significant negative correlation was found between abundance of R. thapari and host body condition.

1. There is increasing interest in the impacts of El Niño Southern Oscillation (ENSO) on reproduction of apical predators such as seabirds and marine mammals. Long-term studies documenting ENSO effects on reproduction of seabirds in the warm tropics are scarce, and differential sensitivity of breeding parameters to ENSO has rarely been explored. 2. Analysis of 18 years of breeding data from a colony of the blue-footed booby Sula nebouxii (Milne-Edwards) showed a delay in onset of breeding when the global Southern Oscillation Index was negative; each unit of the atmospheric pressure differential (hPa) across the Pacific Ocean meant a delay of 7 days. 3. ENSO conditions also produced declines in breeding participation, clutch size, brood size, hatching success and fledging success, especially when surface waters surrounding the colony were warmer during winter and spring. Each additional degree (°C) of water temperature produced a reduction of 0.45 fledglings per nest. Different breeding parameters were sensitive to ENSO indices in different blocks of months. 4. Warming of local waters during the winter was associated with decline in ocean productivity in the current year and the following year, consistent with ENSO impacts on breeding parameters being mediated by effects on local productivity and prey availability. However, there was no evidence of lagged effects of ENSO on any breeding parameter. 5. Comparison of 5 years revealed that when local surface waters were warm, chicks grew more slowly, but no effects of ENSO on weight and size of eggs were evident in data of 9 and 7 years, respectively. 6. Our findings extend evidence of impacts of ENSO on seabird reproduction to the eastern tropical Pacific and indicate that several breeding parameters of blue-footed boobies (but not egg size) are affected in the short term by ENSO conditions, particularly by local anomalies in sea surface temperature associated with decline in ocean productivity., (© 2011 The Authors. Journal of Animal Ecology © 2011 British Ecological Society.)

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.

Recent studies of wild populations provide compelling evidence that survival and reproduction decrease with age because of senescence, a decline in functional capacities at old ages. However, in the wild, little is known about effects of parental senescence on offspring quality. We used data from a 21-year study to examine the role of parental age on offspring probability of recruitment in a long-lived bird, the blue-footed booby (Sula nebouxii). Offspring probability of recruiting into the breeding population varied over the life of parents and effects age were similar in mothers and fathers. Offspring recruitment was high when parents were roughly 6-12 years old and low before and after then. Effects of parental age on offspring recruitment varied with lifespan (parental age at last reproduction) and previous breeding experience. Offspring recruitment from young and old parents with long reproductive lifespans was greater than that of offspring from parents with short lifespans at young and old ages. For parents with little previous breeding experience recruitment of offspring decreased with their hatch date, but experienced parents were no similarly affected. We found evidence of terminal effects on offspring recruitment in young parents but not in older parents, suggesting that senescence is more likely a gradual process of deterioration than a process of terminal illness. Failure to recruit probably reflects mortality during the first years after independence but also during the fledgling transition to full independence. Our results show effects of parental age and quality on offspring viability in a long-lived wild vertebrate and support the idea that wild populations are composed of individuals of different quality, and that this individual heterogeneity can influence the dynamics of age-structured populations.

Somatic deterioration in ageing animals may arise from allocation of resources to reproduction at the expense of repair and maintenance. Thus, accumulated reproductive effort is likely to progressively limit the expression of sexual ornaments at older ages. We analysed the effect of age and reproductive effort on the sexual attractiveness (foot colour) of male blue-footed boobies. Using a long-term dataset, we found that, as animals age and accumulate reproductive events, the expression of foot colour deteriorates. In addition, after non-breeding events males displayed more colourful feet compared with males that reproduced the year before, suggesting that sabbatical years facilitate recovery. Our results indirectly support the idea that allocation of resources to reproduction limits sexual attractiveness and that animals could cope with the negative effects of senescence on sexual ornaments by skipping some breeding events.

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.

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.

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-footed 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 11 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.

In many parentally fed species, siblings compete for food not only by begging and scrambling, but also by violently attacking each other. This aggressive competition has mostly been studied in birds, where it is often combined with dominance subordination, aggressive intimidation, and siblicide. Previous experimental and theoretical studies proposed several life-history, morphological, and behavioral variables that may facilitate the evolution of broodmate aggression, and explain its taxonomic distribution. Here we apply phylogenetic comparative analyses for the first time to test the influence of five hypothesized facilitators of the evolution of broodmate aggression, analyzing 69 species in seven avian families using two quantitative measures of aggression: incidence and intensity. We show that incidence and intensity of aggression increase with long nestling periods and indirect feeding, and small brood size is associated with intense aggression. Large food parcels were not correlated with either the incidence or intensity of aggression. Our study suggests that indirect feeding, long nestling periods, and small broods, possibly in combination with other factors, have tended to favor the evolution of aggressive broodmate competition.

1. Understanding the effects of individual and population factors on variation in breeding dispersal (the movement of individuals between successive breeding sites) is key to identifying the strategies behind breeders' movements. Dispersal is often influenced by multiple factors and these can be confounded with each other. We used 13 years of data on the locations, mates, breeding success and ages of individuals to tease apart the factors influencing breeding dispersal in a colonially breeding long-lived seabird, the blue-footed booby Sula nebouxii. 2. Breeding dispersal varied among and within years. Males dispersed further in years of higher population density, and late breeding males and females dispersed further than early breeders. This temporal variation related to changes in competition for territory was taken into account in all tests of individual factors influencing breeding dispersal. 3. Individuals that retained their mates from the previous year dispersed shorter distances than those that changed their mates. 4. The effect of previous breeding success depended on mate fidelity. Unsuccessful breeding induced greater dispersal in birds that changed their mates but not in birds that retained their mates, indicating that breeders who change mates may take their own previous breeding experience into account during habitat selection. Faithful individuals may have to stay close to their previous sites to encounter their mates. 5. Male divorcees dispersed over shorter distances than their former mates, possibly because males contribute more than females to establishing territories. 6. Dispersal of males and females declined with increasing age over the first 10-11 years of life, then increased in old age, possibly due to senescent decay in the ability to compete for mates and territories.

This study reports an experimental confirmation of the terminal investment hypothesis, a longstanding theoretical idea that animals should increase their reproductive effort as they age and their prospects for survival and reproduction decline. Previous correlational and experimental attempts to test this hypothesis have yielded contradictory results. In the blue-footed booby, Sula nebouxii, a long-lived bird, after initial increase, male reproductive success declines progressively with age. Before laying, males of two age classes were challenged with lipopolysaccharide to elicit an immune response, which induced symptoms of declining survival prospects. Reproductive success of immune-challenged mature males fell, while that of immune-challenged old males showed a 98% increase. These results demonstrate that senescent males with poor reproductive prospects increase their effort when those prospects are threatened, whereas younger males with good reproductive prospects do not.

Drawing on the concepts and theory of dominance in adult vertebrates, this article categorizes the relationships of dominance between infant siblings, identifies the behavioral mechanisms that give rise to those relationships, and proposes a model to explain their evolution. Dominance relationships in avian broods can be classified according to the agonistic roles of dominants and subordinates as "aggression-submission," "aggression-resistance," "aggression-aggression," "aggression-avoidance," "rotating dominance," and "flock dominance." These relationships differ mainly in the submissiveness/pugnacity of subordinates, which is pivotal, and in the specificity/generality of the learning processes that underlie them. As in the dominance hierarchies of adult vertebrates, agonistic roles are engendered and maintained by several mechanisms, including differential fighting ability, assessment, trained winning and losing (especially in altricial species), learned individual relationships (especially in precocial species), site-specific learning, and probably group-level effects. An evolutionary framework in which the species-typical dominance relationship is determined by feeding mode, confinement, cost of subordination, and capacity for individual recognition, can be extended to mammalian litters and account for the aggression-submission and aggression-resistance observed in distinct populations of spotted hyenas and the "site-specific dominance" (teat ownership) of some pigs, felids, and hyraxes. Little is known about agonism in the litters of other mammals or broods of poikilotherms, but some species of fish and crocodilians have the potential for dominance among broodmates.

Do aggressive dominance and subordination in vertebrate broods and litters affect development? We examined 1,167 fledglings from two-chick broods of the blue-footed booby (Sula nebouxii), a species in which the first-hatched chick dominates with violent attacks throughout the nestling period and subordinates suffer lower fledging success, but if both broodmates survive, they grow to the same size. There was little evidence that dominant fledglings were more likely to recruit into the breeding population than were subordinate fledglings, and there was no evidence that dominant and subordinate recruits differed in their age, date, brood size, or nest success at first reproduction or in their summed brood sizes or total nest success over the first 5 yr or first 10 yr of life. Compared with dominants, subordinate fledglings were less prejudiced by late hatching and established clutches earlier over the first 10 yr, and subordinate recruits had 33% larger broods over the first 5 yr. However, in broods where both chicks fledged, accumulated reproductive success for chicks up to age 5 yr was similar for dominants and subordinates. Exercising dominance throughout infancy apparently does not fortify a chick for the future and may incur a long-term cost, and suffering violent subordination throughout infancy has little or no prejudicial effect and may even steel a chick for adult life.