Group structure,within-group conflict and reproductive tactics in cooperatively breeding Galápagos mockingbirds,Nesomimus parvulus

Reproductive conflict within groups can be an important feature of cooperative breeding systems, especially when more than one individual of a sex breeds within a social group. Relationships between group structure, dominance, within-group conflict and reproductive tactics of cooperatively breeding Galápagos mockingbirds were examined on Isla Genovesa. Territorial groups of 2–24 adults included up to three breeding females, with 42% of the groups containing more than one (plural groups); females in most plural groups nested separately. Territory size increased with group size, but the area available per pair in plural groups was smaller than in singular groups (groups with only one breeding pair). Most pairings were monogamous, and males usually outnumbered females; high-ranking males obtained mates more frequently than subordinate males. In 3 relatively dry years, but not in a wet El Niño year, subordinate pairs in plural groups fledged fewer young than dominant pairs or pairs breeding in singular groups. Interference by dominant breeders, often leading to abandonment of nests by subordinate pairs, appears to account for these differences: through nest disruption in drier years, dominant individuals may reduce the cost of sharing their territories and increase the chances of recruiting helpers. Dominant males in plural groups may also father young through extra-pair copulations with subordinate females. Despite costs imposed by within-group conflict, subordinate breeders have higher long-term reproductive success than birds that defer breeding. Plural group structure is maintained because unpredictable climatic variation favours opportunistic breeding by subordinates.     

Helping effort and future fitness in cooperation animal societies

Little attention has been paid to a conspicuous and universal feature of animal societies: the variation between individuals in helping effort. Here, we develop a multiplayer kin-selection model that assumes that subordinates face a trade-off because current investment in help reduces their own future reproductive success. The model makes two predictions: (i) subordinates will work less hard the closer they are to inheriting breeding status; and (ii) for a given dominance rank, subordinates will work less hard in larger groups. The second prediction reflects the larger pay-off from inheriting a larger group. Both predictions were tested through a field experiment on the paper wasp Polistes dominulus. First, we measured an index of helping effort among subordinates, then we removed successive dominants to reveal the inheritance ranks of the subordinates: their positions in the queue to inherit dominance. We found that both inheritance rank and group size had significant effects on helping effort, in the manner predicted by our model. The close match between our theoretical and empirical results suggests that individuals adjust their helping effort according to their expected future reproductive success. This relationship has probably remained hidden in previous studies that have focused on variation in genetic relatedness.     

INTRASPECIFIC VARIATION IN SEX ALLOCATION IN A SIMULTANEOUS HERMAPHRODITE: THE EFFECT OF INDIVIDUAL SIZE

Within a population of simultaneous hermaphrodites, individuals may vary in both their current reproductive investment (biomass invested in gonads) and in how they allocate that investment between male and female function. In the chalk bass, Serranus tortugarum, estimates of both reproductive allocation and reproductive success as a male and a female can be made for individuals of different sizes. As individuals increase in size, their investment in gamete production increases, and there is a shift in allocation to a stronger female bias. Spawning frequency as a female in pair spawnings and as a male in both pair spawning and streaking (an alternative mating tactic) does not vary with individual size. As a result, larger individuals should release more sperm or eggs per spawn. Size-assortative pair spawning in this species leads to larger individuals having higher potential returns in total male reproductive success than smaller individuals, which should lead to increases in absolute levels of sperm production in larger individuals when individuals compete for fertilizations through sperm competition. However, smaller individuals contribute a smaller proportion of the sperm released in spawns with multiple spawners and thus are under more intense sperm competition than larger individuals, which should select for increases in male allocation in smaller individuals, all else equal. A local-mate-competition (LMC) model predicts that these factors select for increasing absolute male and female investment with individual size but a relative shift to more female-biased allocation as individual size increases. These predictions are supported by gonadal data. The predictions of average male allocation from the quantitative LMC model were 21.6% and 25.7%, whereas the collections averaged 21.3%. This close agreement of both the mean male allocation and its relative shift with individual size between model and data support the hypothesis that size-specific shifts in sex allocation in this species represent an adaptive response to patterns of mating success and sperm competition.     

Small Subordinate Male Advantage in the Zebrafish

Dominance is an important determinant of reproductive success in many species, and size is usually an indicator of dominance status, with larger, dominant individuals physically and physiologically preventing smaller subordinates from mating. However, small size may be advantageous in some mating contexts because enhanced manoeuvrability enables males to get closer to females during mating. Here, we determined the paternity success and testes size of dominant and subordinate male zebrafish (Danio rerio), in pairs that controlled for social status. There was no statistical difference in both body size and testes size between dominant and subordinate males. Dominant males sired significantly more offspring than subordinates, but when subordinates were small, they had a greater share of the paternity than larger subordinates. Small male advantage may be one mechanism by which variation in body size is maintained in this species.     

Maternal investment during pregnancy in wild meerkats

Maternal investment in offspring development is a major determinant of the survival and future reproductive success of both the mother and her young. Mothers might therefore be expected to adjust their investment according to ecological conditions in order to maximise their lifetime fitness. In cooperatively breeding species, where helpers assist breeders with offspring care, the size of the group may also influence maternal investment strategies because the costs of reproduction are shared between breeders and helpers. Here, we use longitudinal records of body mass and life history traits from a wild population of meerkats (Suricata suricatta) to explore the pattern of growth in pregnant females and investigate how the rate of growth varies with characteristics of the litter, environmental conditions, maternal traits and group size. Gestational growth was slight during the first half of pregnancy but was marked and linear from the midpoint of gestation until birth. The rate of gestational growth in the second half of pregnancy increased with litter size, maternal age and body mass, and was higher for litters conceived during the peak of the breeding season when it is hot and wet. Gestational growth rate was lower in larger groups, especially when litter size was small. These results suggest that there are ecological and physiological constraints on gestational growth in meerkats, and that females may also be able to strategically adjust their prenatal investment in offspring according to the likely fitness costs and benefits of a particular breeding attempt. Mothers in larger groups may benefit from reducing their investment because having more helpers might allow them to lower reproductive costs without decreasing breeding success.     

Factors influencing reproductive success and litter size in captive island foxes

A severe decline of island foxes (Urocyon littoralis) on the northern Channel Islands in the 1990s prompted the National Park Service to begin a captive breeding program to increase their numbers. Using detailed records of all the fox pairs (N = 267) that were part of the program on San Miguel, Santa Rosa, and Santa Cruz Islands from its inception in 2000 through 2007, we identified factors influencing the breeding success of pairs in captivity in the interest of formulating strategies that could increase captive productivity. We compiled a database of variables including litter size, reproductive success, distance to nearest occupied pen during the breeding season, subspecies, exposure, female age, male age, age difference, female and male origin (wild vs. captive born), same versus different origin, years paired, previous reproductive success by the pair, previous reproductive success by the female, mate aggression-related injuries, male previous involvement in a pair with mate aggression, and female previous involvement in a pair with mate aggression. We used multiple linear regression to identify factors predictive of litter size, and logistic regression to predict the probability of reproductive success. A larger inter-pen distance, older male age, less exposure, and a smaller intra-pair age difference positively affected litter size. The probabilities of reproductive success increased with fewer years paired and less exposure. Comparatively, pairs with wild born females (vs. captive born females), and previously successful pairs (vs. previously unsuccessful and new pairs) were most likely to be successful. These results indicate that the optimal situation was to pair wild-caught females with older males in sheltered pens that were as far from other pens as possible, to maintain successful pairs and repair unsuccessful ones. Published 2013. This article is a U.S. Government work and is in the public domain in the USA.     

Effects of pair migratory behavior on breeding phenology and success in a partially migratory shorebird population

In migratory systems, variation in individual phenology can arise through differences in individual migratory behaviors, and this may be particularly apparent in partial migrant systems, where migrant and resident individuals are present within the same population. Links between breeding phenology and migratory behavior or success are generally investigated at the individual level. However, for breeding phenology in particular, the migratory behaviors of each member of the pair may need to be considered simultaneously, as breeding phenology will likely be constrained by timing of the pair member that arrives last, and carryover effects on breeding success may vary depending on whether pair members share the same migratory behavior or not. We used tracking of marked individuals and monitoring of breeding success from a partially migrant population of Eurasian oystercatchers (Haematopus ostralegus) breeding in Iceland to test whether (a) breeding phenology varied with pair migratory behavior; (b) within‐pair consistency in timing of laying differed among pair migratory behaviors; and (c) reproductive performance varied with pair migratory behavior, timing of laying, and year. We found that annual variation in timing of laying differed among pair migratory behaviors, with resident pairs being more consistent than migrant and mixed pairs, and migrant/mixed pairs breeding earlier than residents in most years but later in one (unusually cold) year. Pairs that laid early were more likely to replace their clutch after nest loss, had higher productivity and higher fledging success, independent of pair migratory behavior. Our study suggests that the links between individual migratory behavior and reproductive success can vary over time and, to a much lesser extent, with mate migratory behavior and can be mediated by differences in laying dates. Understanding these cascading effects of pair phenology on breeding success is likely to be key to predicting the impact of changing environmental conditions on migratory species.     

Spacing pattern and body size composition of the protandrous anemonefish Amphiprion frenatus inhabiting colonial host anemones

The protandrous anemonefish Amphiprion frenatus often forms a group consisting of a large female, a small male, and a smaller nonbreeder at an isolated single host anemone, where home ranges of subordinates were covered with the female's home range. Within the group, the dominant individuals suppress the growth of subordinates, resulting in large size differences. The spacing pattern and body size composition of A. frenatus on colonial hosts were investigated in Ishigaki Island, Okinawa, Japan. Six breeding pairs and 14 nonbreeders inhabited a colony of 157 anemones. Each pair maintained a territory in which pair members used different hosts. Nonbreeders had unstable home ranges on the outskirts of or in the pairs' territories. Body size differences between males and females in pairs and between males and nonbreeders were small. The small size differences in the colony of hosts are caused by reduced suppression of growth of subordinates by the dominant individuals. The total area of host anemones largely affects spacing pattern and social suppression of the anemonefish.     

Within-season divorce in Blue Tits (Cyanistes caeruleus)

Divorce between breeding seasons, i.e. mate change while the old breeding partner is still alive, has been well-studied in several bird species. It can be viewed either as the result of reproductive decisions of individuals to maximize their own fitness or as the outcome of intra-sexual competition for mating opportunities. In contrast, divorce within a breeding season—also referred to as rapid mate switching—has received much less attention. Within-season divorce may occur after sudden changes in environmental conditions, such as those causing the disappearance or devaluation of nesting sites or territories, or when better breeding partners become available. Within-season divorce can be initiated by a member of the pair, or it can be the result of a take-over by an unpaired individual that competes for access to the resource. During a field experiment investigating the effects of limiting nesting sites on reproductive behaviour in Blue Tits, we recorded several cases of within-season divorce. The rate of divorce was not related to the experimental nest-site limitation, and pairs that changed their partner suffered reduced reproductive success compared to faithful pairs. Although there were no differences in the timing of breeding, clutch size or hatching success, pairs with a new partner also suffered a reduced fledging success, which was partly explained by complete brood failures. This study highlights that the pair bond prior to egg laying can be unstable when conditions force individuals to compete for a new partner or nest site and indicates the importance of the correct timing of divorce within the breeding cycle.     

Differential maternal investment counteracts for late breeding in magpies Pica pica: an experimental study

Reproductive success in many avian populations declines throughout the breeding season. Two hypotheses have gained attention to explain such a decline: the "timing" hypothesis proposes that deteriorating food availability causes the decline in reproductive success (causal effect of breeding time), whereas the "quality" hypothesis proposes that individuals of lower phenotypic quality reproduce at the end of the breeding season, causing the correlation between breeding time and breeding success. We tested both of these hypotheses in a monogamous single breeder, the magpie Pica pica , by experimentally inducing some pairs to lay a replacement clutch, after removal of the first clutch. The first clutch was left in the nest of another magpie pair (matched by laying date and clutch size), and incubated and raised by these foster parents. In this way we obtained two clutches from the same magpie pair with full siblings raised in conditions of the first and second reproductive attempts. High quality pairs (with laying dates in the first half of the breeding season) reached similar breeding success in replacement clutches as compared to first clutches of the same female. In addition, experimental pairs reared significantly more offspring of similar quality in their replacement clutches as compared to late-season first clutches, thereby suggesting that late season first clutches were produced by pairs of lower phenotypic quality. Results indicate that high quality pairs trade-off clutch size for larger eggs in replacement clutches, which could help magpie pairs to partly compensate for poorer environmental conditions associated with a delayed breeding attempt.     

Group composition affects male reproductive partitioning in a cooperatively breeding cichlid

Individuals within groups of cooperatively breeding species may partition reproduction, with the dominant pair often taking the largest share. The dominant's ability to reproductively control subordinates may depend on differences in competitive ability, due to, e.g. body size differences, but may also depend on the number of same‐sex competitors inside the group. We tested experimentally whether subordinates reproduce more when these subordinates are large or when a second subordinate of the same sex need to be controlled by the dominants, using the cooperatively breeding cichlid Neolamprologus pulcher. Dominant pairs were assisted by a large and a small unrelated subordinate; sexes of these fish were varied in a full‐factorial design (giving four treatments). Dominant males lost significantly more parentage to the large subordinate male when a small subordinate male was also present, compared to when a small subordinate female was present. However, subordinate paternity was generally low and did not significantly curb total dominant male reproductive output, which was more affected by the sizes and numbers of reproductive females present inside his group. Dominant female maternity, clutch sizes and total output did not depend on the treatments. Subordinate–subordinate reproduction was virtually absent (one out of 874 offspring). Female subordinates were more likely to provide care for their own broods. In contrast, male subordinates did not adjust their level of care to their parentage. Variability in female subordinate alloparental brood care was particularly high, with females showing more care than males in general. We also detected effects of growth rate and food ration on parentage independent of the treatments, most notably: (i) a trade‐off between dominant male growth rate and paternity; (ii) a decrease in dominant male paternity with increasing food ration; (iii) a positive effect of growth rate on paternity in small males. We conclude that dominant males should be sensitive to the number and sizes of subordinate males present in their group, particularly when these subordinates are not helpful or grow fast, and food is plentiful. Dominant females should be less sensitive, because female subordinates do not appear to impose reproductive costs and can be helpful through alloparental brood care.     

Determinants of reproductive success in dominant pairs of clownfish: a boosted regression tree analysis

1. Central questions of behavioural and evolutionary ecology are what factors influence the reproductive success of dominant breeders and subordinate nonbreeders within animal societies? A complete understanding of any society requires that these questions be answered for all individuals. 2. The clown anemonefish, Amphiprion percula, forms simple societies that live in close association with sea anemones, Heteractis magnifica. Here, we use data from a well-studied population of A. percula to determine the major predictors of reproductive success of dominant pairs in this species. 3. We analyse the effect of multiple predictors on four components of reproductive success, using a relatively new technique from the field of statistical learning: boosted regression trees (BRTs). BRTs have the potential to model complex relationships in ways that give powerful insight. 4. We show that the reproductive success of dominant pairs is unrelated to the presence, number or phenotype of nonbreeders. This is consistent with the observation that nonbreeders do not help or hinder breeders in any way, confirming and extending the results of a previous study. 5. Primarily, reproductive success is negatively related to male growth and positively related to breeding experience. It is likely that these effects are interrelated because males that grow a lot have little breeding experience. These effects are indicative of a trade-off between male growth and parental investment. 6. Secondarily, reproductive success is positively related to female growth and size. In this population, female size is positively related to group size and anemone size, also. These positive correlations among traits likely are caused by variation in site quality and are suggestive of a silver-spoon effect. 7. Noteworthily, whereas reproductive success is positively related to female size, it is unrelated to male size. This observation provides support for the size advantage hypothesis for sex change: both individuals maximize their reproductive success when the larger individual adopts the female tactic. 8. This study provides the most complete picture to date of the factors that predict the reproductive success of dominant pairs of clown anemonefish and illustrates the utility of BRTs for analysis of complex behavioural and evolutionary ecology data.     

Reproductive success depends on the quality of helpers in the endangered,cooperative El Oro parakeet (Pyrrhura orcesi)

In cooperative species, helping behaviour and reproductive success can be correlated, but understanding this correlation is often impaired by the difficulty to correctly infer causation. While helpers can incur costs by participating in brood care, it is yet unclear if their help depends on their individual quality. We address these questions in the previously unknown cooperative breeding system of the endangered El Oro parakeet (Pyrrhura orcesi). Specifically, we ask (i) whether breeders benefit directly from helpers by an enhanced reproductive success and if so, (ii) whether the amount of this potential benefit is regulated by the quality of contributing group members. Groups consist of a dominant breeding pair accompanied by helpers, but cooperation is not obligate. Microsatellite heterozygosity was used to assess individual quality; its suitability as indicator of quality was reflected in the positive relationship between offspring heterozygosity and recruitment into the population. The reproductive success of breeding pairs depended on helper (genetic) quality and the number of helpers. This relationship occurred on two different levels: clutch size and fledging success, indicating (i) that females profit from high‐quality helpers and probably adjust clutch size accordingly and (ii) that the helpers increase fledging success. Congruently, we found that offspring body condition is positively affected by helper quality, which is most probably explained by the increased feeding rates when helpers are present. We suggest a causal link between cooperation and reproductive success in this frugivorous, endangered parakeet. Further, helper (genetic) quality can be a relevant factor for determining reproductive fitness in cooperative species, particularly in small and bottlenecked populations.     

Effects of extrinsic and intrinsic factors on breeding success in a long lived seabird

There is growing concern over the impacts of climate change on animal species. Many studies have demonstrated impacts of climate change at the population level, and density dependent effects of climate are frequently reported. However, there is an increasing recognition of the differential impact of such factors on individuals since there is marked variation in individual performance. We investigated the relationships between breeding success and environmental conditions (winter NAO and one year lagged winter NAO) and intrinsic effects (colony size, pair bond duration, past breeding success rate) in the northern fulmar Fulmarus glacialis , using data from a long-term study commenced in 1950. There was a negative trend in breeding success over time, and a negative relationship with winter NAO and lagged winter NAO, which themselves had shown positive increases over the study period. The effects of lagged winter NAO remained after accounting for the linear trend. There was no evidence of density dependence, with breeding success positively related to colony size. We found strong evidence that breeding success was negatively related to pair bond duration but positively related to past breeding success rate. There was also an interaction between these two intrinsic effects such that those pairs that had historically been successful maintained success with increasing pair bond duration, whereas less successful pairs showed a decline. The prediction that there would be a differential impact of extrinsic factors among pairs was supported by an interaction between past breeding success rate and winter NAO, such that pairs with low past success rate exhibited a sharp decline in breeding success with increasing winter NAO, whereas more successful pairs did not. It is critically important to understand interactions between extrinsic factors and individual heterogeneity since a differential impact on individuals will affect population structure, and hence population dynamics.     

Helping effort in a dominance hierarchy

In many cooperatively breeding species, group members form adominance hierarchy or queue to inherit the position of breeder.Models aimed at understanding individual variation in helpingbehavior, however, rarely take into account the effect of dominancerank on expected future reproductive success and thus the potentialdirect fitness costs of helping. Here we develop a kin-selectionmodel of helping behavior in multimember groups in which onlythe highest ranking individual breeds. Each group member caninvest in the dominant's offspring at a cost to its own survivorship.The model predicts that lower ranked subordinates, who havea smaller probability of inheriting the group, should work harderthan higher ranked subordinates. This prediction holds regardlessof whether the intrinsic mortality rate of subordinates increasesor decreases with rank. The prediction does not necessarilyhold, however, where the costs of helping are higher for lowerranked individuals: a situation that may be common in vertebrates.The model makes two further testable predictions: that the helpingeffort of an individual of given rank should be lower in largergroups, and the reproductive success of dominants should begreater where group members are more closely related. Empiricalevidence for these predictions is discussed. We argue that theeffects of rank on stable helping effort may explain why attemptsto correlate individual helping effort with relatedness in cooperativelybreeding species have met with limited success.     

Group Structure,Nest Size and Reproductive Success in the Cooperatively Breeding Cichlid Julidochromis ornatus: A Correlation Study

The effect of group size on reproductive success has long been studied in cooperatively breeding species, as it might provide an adaptive explanation for group‐living in social species. Numerous studies have shown positive effects of subordinates on reproductive success (‘helper effect’), but these studies have also revealed the importance of controlling statistically, or experimentally, for the effect of other factors that might affect reproductive success. Here, we first examine the relationships between group size, body size of group members and nest size in the cooperatively breeding cichlid Julidochromis ornatus, in which unrelated helpers frequently participate in reproduction and their breeding nests inside rock crevices may be crucial for reproduction and survival of all group members. Then, we subsequently investigate the relationship between group size and reproductive success, while controlling for these factors. The results showed that group size was significantly related to body size of group members rather than nest size; and larger breeders had larger helpers. It was found that group size significantly increased group reproductive output. More importantly, reproductive success of male breeders did not depend on the presence of mature helpers, whereas female reproductive success increased when two males assisted her and tended to decrease when two females bred cooperatively. We conclude that breeding groups of J. ornatus have size hierarchical societies that relate to group size, and group composition of genetically unrelated and co‐breeding members affects their reproductive success.     

Assessing the reproductive consequences of mate retention and pair bond duration in Thorn-tailed Rayadito (Aphrastura spinicauda), a short-lived,socially monogamous neotropical bird

The adaptive value of mate retention has been studied in several socially monogamous birds but evidence of reproductive benefits for short-lived species is inconclusive. Most studies come from northern latitudes but more research on tropical birds is needed, as these species typically show higher survival rates and longer pair bonds than those from temperate regions. We gathered data on the reproductive biology of a subtropical, isolated population of Thorn-tailed Rayadito Aphrastura spinicaudaduring 2008–2017 to evaluate the reproductive consequences of mate retention. We examined data from 243 breeding attempts made by 159 breeding pairs. We found that ~30% of all breeding pairs bred together during at least two consecutive years, and some were mated for 6 years. The main cause of pair dissolution was mate loss, not divorce. Mixed-effects models provided moderate evidence for positive effects of mate retention and successive remating on reproductive success. Newly formed pairs laid eggs later and had slightly smaller clutches than remated pairs. Furthermore, clutch size seemed to increase with successive remating. Overall, our results suggest that newly formed pairs are less efficient in reproduction and that minor yearly reproductive benefits of mate retention might accumulate for birds that are able to breed with the same partner over many years. Because breeding habitat is limited in our study population, Thorn-tailed Rayaditos could benefit from remating if the number of individuals that can breed exceeds the number of available breeding positions. Profitable long-term pair bonds might be more frequent in tropical birds and therefore more studies are needed to assess the prevalence of remating-mediated effects on reproduction in relatively short-lived monogamous species breeding in tropical regions.     

Sex change in either direction by growth-rate advantage in the monogamous coral goby, Paragobiodon echinocephalus

The size-advantage model predicts the evolution of sex changeif the relative reproductive success of the sexes changes withsize or age. In the goby (Paragobiodon echinocephalus) the largesttwo fish, a male and a female matched by size, breed monogamouslyin each host coral. Because the female fecundity and male abilityof egg care increase with body size in a similar way, no size-fecundityadvantage exists. However, we found both protogyny and infrequentprotandry in a natural population of this species in Okinawa.New pairs were often formed after movement between host coralsand also sex change or sex differentiation of one or both members.In most new pairs males were larger than females, and femalesgrew much faster than their mates until breeding (growth-rateadvantage). The smaller member of a new pair should be the femalethat grows faster, because the smaller limits the reproductivesuccess of the pair. To form such a pair, the goby changed sexaccording to the sex and relative size of a new mate, as a status-dependentconditional strategy. The growth-rate advantage predicts predominanceof protogyny, but movement between host corals provides opportunitiesalso for protandry.     

Influence of density‐dependent competition on foraging and migratory behavior of a subtropical colonial seabird

Density‐dependent competition for food resources influences both foraging ecology and reproduction in a variety of animals. The relationship between colony size, local prey depletion, and reproductive output in colonial central‐place foragers has been extensively studied in seabirds; however, most studies have focused on effects of intraspecific competition during the breeding season, while little is known about whether density‐dependent resource depletion influences individual migratory behavior outside the breeding season. Using breeding colony size as a surrogate for intraspecific resource competition, we tested for effects of colony size on breeding home range, nestling health, and migratory patterns of a nearshore colonial seabird, the brown pelican (Pelecanus occidentalis), originating from seven breeding colonies of varying sizes in the subtropical northern Gulf of Mexico. We found evidence for density‐dependent effects on foraging behavior during the breeding season, as individual foraging areas increased linearly with the number of breeding pairs per colony. Contrary to our predictions, however, nestlings from more numerous colonies with larger foraging ranges did not experience either decreased condition or increased stress. During nonbreeding, individuals from larger colonies were more likely to migrate, and traveled longer distances, than individuals from smaller colonies, indicating that the influence of density‐dependent effects on distribution persists into the nonbreeding period. We also found significant effects of individual physical condition, particularly body size, on migratory behavior, which in combination with colony size suggesting that dominant individuals remain closer to breeding sites during winter. We conclude that density‐dependent competition may be an important driver of both the extent of foraging ranges and the degree of migration exhibited by brown pelicans. However, the effects of density‐dependent competition on breeding success and population regulation remain uncertain in this system.     

Living with strangers: direct benefits favour non-kin cooperation in a communally nesting bird

The greater ani (Crotophaga major), a Neotropical cuckoo, exhibits an unusual breeding system in which several socially monogamous pairs lay eggs in a single nest and contribute care to the communal clutch. Cooperative nesting is costly-females compete for reproduction by ejecting each other's eggs-but the potential direct or indirect fitness benefits that might accrue to group members have not been identified. In this study, I used molecular genotyping to quantify patterns of genetic relatedness and individual reproductive success within social groups in a single colour-banded population. Microsatellite analysis of 122 individuals in 49 groups revealed that group members are not genetic relatives. Group size was strongly correlated with individual reproductive success: solitary pairs were extremely rare and never successful, and nests attended by two pairs were significantly more likely to be depredated than were nests attended by three pairs. Egg loss, a consequence of reproductive competition, was greater in large groups and disproportionately affected females that initiated laying. However, early-laying females compensated for egg losses by laying larger clutches, and female group members switched positions in the laying order across nesting attempts. The greater ani, therefore, appears to be one of the few species in which cooperative breeding among unrelated individuals is favoured by direct, shared benefits that outweigh the substantial costs of reproductive competition.