Effects of foundress number on brood raids and queen survival in the fire ant Solenopsis invicta

In several ant species, colonies are founded by small groups of queens (pleometrosis), which coexist until the first workers eclose, after which all but one queen is killed. It has been hypothesized that, by producing a larger cohort of workers, cooperating queens may increase colony success during brood raids, a form of competition in which brood and workers from losing nests are absorbed into winning colonies. To test whether this benefit is sufficient to favor pleometrosis, newly mated queens of the fire ant Solenopsis invicta were assembled in groups of one, two, three, or four, reared in the laboratory until the first workers eclosed, then planted in the field in replicated assemblages. The proportion of colonies engaging in brood raids increased with average foundress number per nest and with colony density but was unaffected by variance in foundress number among interacting colonies. Within mixed assemblages of single-queen and multiple-queen colonies, queen number had no effect on the likelihood of engaging in raids or the probability of nest survival through the brood raiding period. However, following nearly 30% of raids, queens moved to new nests and displaced the resident queens. When queen relocation and subsequent mortality were accounted for, it was found that the survival of queens from four-queen groups was substantially higher than that of solitary queens. By contrast, the survival of queens from two-queen colonies was no greater than that of solitary queens. These results show that the competitive advantages of multiple-queen colonies are sufficient to counterbalance the increased mortality of queens within groups only when the number of foundresses is greater than two and when colonies are founded at high density. When colonies lose brood raids, the workers appear to abandon their mothers to join surviving colonies. However, in laboratory experiments, queens attempting to enter foreign nests were significantly more likely to displace the resident queen if their own daughters were present within the invaded nest. Thus, workers may be able to bias the probability that their mother rejoins them and displaces competing queens.     

Colony genetic diversity affects task performance in the red ant Myrmica rubra

High relatedness and low genetic diversity among individuals in a group is generally considered crucial to the evolution of cooperative behaviour. However, in about a third of social insect species, intracolonial genetic diversity is increased because of derived polyandry (multiple mating by queens) and/or polygyny (multiple reproductive queens). Several studies have shown that increased intracolonial genetic diversity can enhance task performance in honey bees, but evidence of such effect in other social insects is still lacking. Why increased genetic diversity has evolved in some, but not all species, is a fundamental question in sociobiology. In this study, we investigated the effect of intracolonial genetic diversity on the task of nest migration, using the facultatively polyandrous and polygynous red ant Myrmica rubra. Genetic diversity significantly affected migration speed, but its effects were context dependent. Migration speed correlated positively with genetic diversity in one experiment in which migrations were into a known nest site, due to quicker transfer of brood into the new nest once consensus was reached. However, in a another experiment in which migration included scouting for new nest sites, migration speed correlated negatively with genetic diversity, due to slower discovery of new nest sites and slower transfer of brood into the new nest. Our results show for the first time that genetic diversity affects task performance in a social insect other than the honeybee, but that it can produce contrasting effects under different conditions.     

Colony founding,queen dominance and oligogyny in the Australian meat ant Iridomyrmex purpureus

Summary Field observations and laboratory experiments demonstrate that in the Australian meat ant, Iridomyrmex purpureus, the modes of colony founding are remarkably diverse. New colonies can originate from single foundresses (haplometrosis), or foundress associations (pleometrosis), or by colony budding, or the adoption of newly-mated queens that dig founding chambers next to mature nests (probably their natal nests, as workers protect them and may help them dig). Readoption of foundresses and pleometrosis lead to the coexistence of several queens in one nest. We discovered a striking antagonistic behavior among coexisting queens in young colonies, in the form of ritualized antennation bouts. These interactions result in a reproductive rank order in which dominant queens inhibit egg-laying by subordinates, but escalation into physical fighting is rare. Workers ignore queen dominance interactions and treat all queens equally. The first quantitative ethogram of dominance display behavior between multiple ant queens, and its reproductive consequences, is presented. As a colony grows, queens become intolerant of each other's presence and permanently separate within the nest. Once separated, queens appear to be equal in status, laying approximately equal numbers of eggs. All queens continue to be tolerated by workers, even when the colony has reached a size of several thousand workers and begun to produce reproductives. Such mature nests of I. purpureus fulfill the criteria of oligogyny, defined by worker tolerance toward more than one queen and antagonism among queens, such that a limited number of fully functional queens are spaced far apart within a single colony. Oligogynous colonies can arise in this species by pleometrotic founding (primary oligogyny) or by adoption of queens into existing nests (secondary oligogyny). The adaptive significance of the complex system of colony founding, queen dominance and oligogyny in I. purpureus is discussed.     

Regional variation in queen and worker aggression in incipient colonies of the desert ant Messor pergandei

The fate of queen foundress associations in ants varies across taxa: in some, lethal fighting results in survival of a single queen, while in others, queens coexist long term. One hypothesis for this difference is that selection favors fighting when group sizes are small and tolerance when groups are large. In an experiment with the ant Messor pergandei, we formed small, medium, and large groups with newly mated queens from field populations that have different mean group sizes and differ in whether multiple queens occur in older established colonies. We found that whether queens are eliminated by fighting depends upon region of origin and not group size: regardless of co-foundress number, queens from sites with single-queen adult field colonies displayed agonistic behaviors and their colonies reduced to a single queen, while queens from sites with multiple-queen colonies did not fight and co-foundresses coexisted long term. Worker aggression towards and elimination of queens were also correlated with region of origin. Where fighting occurred, queens were as likely to be killed by workers as by other queens. An aggressive display was the most common form of agonistic interaction among queens, while fighting was relatively rare. We hypothesize that queen displays evolved in response to worker attacks because they increase the probability that workers will eliminate competitor queens. Our results suggest that the evolutionary interests of workers, as well as queens, could be important in determining the evolution and maintenance of queen elimination in foundress associations.     

An experimental study of pleometrotic colony founding in the fire ant, Solenopsis invicta : what is the basis for association?

Many benefits and risks of cooperative colony founding (pleometrosis) have been identified, but rarely have the proximate factors that lead to association been considered. This study examined the choices queens make during the first few hours after mating, and some of the correlates of those choices. Queens had a strong affinity for preformed holes in the soil and readily used these as their initial founding chambers. This affinity was so strong that in a field experiment, the dispersion pattern of preformed holes controlled the final dispersion of colony-founding queens. Attraction to partially formed holes is thus an important cause of pleometrosis. The excavation of complete founding chambers incurred no measurable cost on the subsequent reproductive output of queens, suggesting that the primary benefit of using preformed holes is to remove the queen quickly from exposure to predation and desiccation. In the field, pairs of queens offered five equivalent preformed holes in soil were more likely to share the same hole if the holes were shallow and close together. In these experiments, queens modified preformed soil holes so that the test holes were no longer equivalent, causing the choice of queen and hole to become confounded. Laboratory experiments in plaster arenas with unmodifiable holes confirmed the field experiments: queens were more likely to share a hole when the holes were shallow than when they were deep. Because queens entering adequately deep holes seldom reemerged, this suggested that the likelihood of sharing increased with increasing contact between queens, that is, when queens were readily and frequently detected. Such contacts will also predict the future competitive environments to be experienced by incipient colonies, and may temper the tendency of queens to associate. However, experiments in which queens were exposed to high and low densities before pairing in the choice arenas failed to show an effect on the choice to join the resident queen. Queens that joined a resident queen differed in their robustness from queens that did not join. Queens choosing their own partners did no better reproductively than those assigned partners at random. Overall, this study suggests that (1) newly mated queens are under strong selection to leave the soil surface and do so by using any available holes, whether dug by another queen or of some other origin; (2) they are attracted to other queens, and are more likely to cofound as contact with the potential cofoundress becomes more frequent and (3) they choose whether or not to cofound partly on the basis of their own reproductive characteristics. Received: 20 November 1997 / Accepted after revision: 14 March 1998     

Polydomy and sexual allocation ratios in the ant Myrmica punctiventris

Summary Colony structure and reproductive investment were studied in a population of Myrmica punctiventris. This species undergoes a seasonal cycle of polydomy. A colony overwinters in entirety but fractionates into two or more nest sites during the active season and then coalesces in the fall. Colony boundaries were determined by integrating data on spatial pattern, behavioral compatability, and genetic relatedness as revealed by protein electrophoresis. Colonies contained at most one queen. Consequently, a colony consisted of one queenright nest and one or more queenless nests. Furthermore, estimates of relatedness were fully consistent, with queens being single mated. M. punctiventris therefore has a colony genetic structure that conforms to the classical explanation of the maintenance of worker sterility by kin selection. Kin selection theory predicts that workers would favor a female-biased allocation ratio while selection on queens would favor equal investment in males and females. We predicted that in polydomous populations, queenless nests would rear more female reproductives from diploid larvae than queenright nests. There was a significant difference between queenright and queenless nests in sexual allocation; queenless nests allocated energy to reproductive females whereas queenright nests did not. At neither the nest nor colony levels did worker number limit sexual production. We also found that nests tended to rear either males or females but when colony reproduction was summed over nests, the sexes were more equally represented. The difference in allocation ratios between queenless and queenright nests was attributed solely to queen presence/absence. Our work shows that polydomy provides an opportunity for workers to evade queen control and thereby to sexualize brood.Offprint requests to: L.E. Snyder at the current address     

Brown pelican siblicide and the prey-size hypothesis

Summary We asked whether the brown pelican (Pelecanus occidentalis) practices facultative brood reduction and tested two predictions of Mock's (1985) prey-size hypothesis: (1) if chicks take food directly from the parental mouth, nestmates should compete aggressively; (2) aggression between nestmates should increase during the developmental transition from indirect feeding (parents deposit food on the substrate) to direct feeding (parents pass food from mouth to mouth). Eggs in two-egg and three-egg clutches were laid and hatched 2 days apart. Junior (second- and third-hatched) chicks, fed less than their nestmates, grew more slowly and died more frequently. Of non-hatchling victims 74% were underweight, 55% bore peck wounds and 43% were found outside the nest, implying starvation, aggression and expulsion from the nest. Aggression occurred in all observed sibships, and junior chicks were subordinate in eight of ten cases. A progressive transition from indirect to direct feeding occurred during the 30 days after hatching, but there was no evidence of an increase in pecking during the transition. In conclusion, the brown pelican practices facultative brood reduction, and chick loss is mediated by sibling aggression as predicted by the preysize hypothesis. Pecking during indirect feeds seems to contradict the prey-size hypothesis, but may serve partly to establish a relationship of dominance-subordinance.Correspondence to: H. Drummond     

Queen number, queen cycling and queen loss: the evolution of complex multiple queen societies in the social wasp genus Ropalidia

Complex, highly integrated societies have evolved from simpler societies repeatedly, and the social insects provide an excellent model system for understanding increasing complexity and integration. In the paper wasps, large societies, known as swarm-founding, have evolved repeatedly from smaller societies, known as independent-founding. Swarm-founding colonies have many more queens than independent-founding colonies, which should dramatically reduce relatedness, posing a challenge to cooperation. However, in each instance, swarm-founding species have also evolved a cyclical pattern of queen reduction which elevates relatedness despite high queen numbers. The genus Ropalidia provides an excellent system in which to study the transition to swarm-founding because it has both independent and swarm-founding species. We studied the Australian independent-founding wasp Ropalidia revolutionalis to better understand the evolution of multiple queens and their periodic reductions in swarm-founding wasps. Using microsatellite genetic markers we genotyped queens, workers and brood from 37 colonies and found that while most colonies had a single queen, three of the colonies had multiple queens at or immediately prior to the time of collection. An additional seven colonies had had multiple co-occurring queens earlier in the season. We also found that colonies experienced many queen losses, and that founding queens were gradually lost until they were replaced by a new cohort of daughter queens in many colonies. This pattern is similar to the periodic reductions and replacements in swarm-founding wasps and suggests that multiple queens and queen cycling evolved relatively early in the shift to swarm-founding in Ropalidia.Communicated by R. Page     

Polygyny,relatedness and nest founding in the polygynous myrmicine ant Leptothorax acervorum (Hymenoptera; Formicidae)

Summary There is high within-nest relatedness for functional queens (with corpora lutea), nonfunctional queens (without corpora lutea), and workers in polygynous nests of Leptothorax acervorum. The high functional queen relatedness suggests that young mated queens are adopted back to their mother nest. Functional queen relatedness does not change with the number of queens present in the nest, suggesting that the number of generations of queens, on average two to three, is rather stable. Worker relatedness decreases with increasing number of functional queens per nest (Tables 5, 6). The number of queens contributing offspring to the nest (mothers), estimated from worker and functional queen relatedness, is lower than the number of functional queens, particularly in highly polygynous nests. Estimates of number of mothers in monogynous nests indicate that these nests previously were polygynous (Table 7). There is no correlation between nest relatedness and distance between nests, and budding-off, if present, thus appears to be a rare mode of nest founding (Table 8). There are no indications of inbreeding in the two populations studied since the frequency of heterozygotes is as high as expected from random mating (Table 4). Most likely, polygyny is the rule in L. acervorum and serves to secure the presence of queens in the nest.     

How do fighting ability and nest value influence usurpation contests in <Emphasis Type="Italic">Polistes</Emphasis> wasps?

Most research on animal contests has focused on the factors that influence the intensity and outcome of aggressive contests within nonsocial species, while relatively little is known about contests in social taxa. Here, we examine contests among queens of the social paper wasp, Polistes dominulus. Queens use multiple reproductive strategies, including nesting alone, usurping established colonies, and cooperatively joining other queens. We stage contests between a nesting queen and a challenger to test how resource value (RV) and resource holding potential (RHP) influence (a) who occupies the nest at the end of the contest and (b) the extent of conflict between the queen and challenger. We found that RHP, as measured by individuals’ facial patterns and body size, influenced the outcome of the contest. Challengers with high RHP were more likely to successfully usurp the nest than challengers with low RHP. Interestingly, queens with relatively high RHP were more likely to form a cooperative association with the challenger than queens with lower RHP, suggesting that queens may evict individuals that are an aggressive threat. RV influenced the intensity of conflict. There was more aggressive conflict over large nests than over small nests. Overall, social taxa have complex contest dynamics with important parallels to contests in nonsocial taxa. Studying contests in social taxa provides an important perspective on the factors that influence individual decisions about conflict versus cooperation.     

When do helpers help? Food availability and helping in the moorhen,Gallinula chloropus

Summary The importance of food availability as a constraint upon cooperative behaviour was examined by providing supplementary food on some breeding territories of a population of moorhens (Gallinula chloropus). First brood juveniles on territories provided with food were more likely to feed their younger second brood siblings. Supplementary feeding improved the physical condition of juveniles above that of unfed conspecifics, and birds in better condition were more likely to be helpers. Helpers on fed territories also provided more food to siblings than did their counterparts on unfed territories, enabling parent birds to reduce their work rates. Results are discussed in relation to the benefits which may accrue to helpers and the energetic costs of cooperation which may act as proximate constraints upon potential helpers.     

Solitary nesting and reproductive success in the paper wasp Polistes aurifer

Female Polistes paper wasps are capable of independent nesting, yet many populations demonstrate a mixture of solitary and cooperative nest foundation. Previous studies of Polistes have found survival and/or productivity advantages of cooperative nest foundation compared to solitary nesting, and reproductive skew models have been designed to predict the dynamics of such flexible cooperation. In this paper, we examine the success of different nesting strategies in a previously unstudied population of Polistes aurifer in southern California. The colony cycle of this population is less synchronous than that of other temperate species, and the frequency of solitary nesting averages 86.2%. Our data suggest that this low rate of cooperative nest founding is adaptive, as demonstrated by the lack of survival or productivity advantages for cooperative foundress associations. Due to foundress turnover and nest foundation later in the season, many nests produce only one set of offspring. This results in a loss of the eusocial nature of some nests in the population. Data from a small sample of multifoundress nests show significant positive reproductive skew, despite concession model predictions that skew should be low in populations with low ecological constraints on independent nesting. This lack of support for the concessions skew model reflects a diminished incentive for cooperation.Communicated by L. Keller     

Assured fitness returns favor sociality in a mass-provisioning sweat bee,Megalopta genalis (Hymenoptera: Halictidae)

Assured fitness returns models for the evolution of sociality emphasize the selective value of ensuring that offspring receive adequate parental care to reach maturity. If a member of a social group dies, it can accrue returns on investment in offspring through the efforts of surviving social partners. We provide evidence that in the mass-provisioning, facultatively social sweat bee Megalopta genalis, adult presence in the nest throughout brood development provides protection from ant predation. Nests with adults present were well protected, and brood in nests with adults removed suffered higher predation. Females in observation nests showed effective defensive behavior against experimentally introduced ants, and bees in natural nests repulsed naturally occurring ant raids. Megalopta nest architecture and behavior are such that the brood of several cooperating females can be defended with little additional cost relative to solitary nesting. The benefits of cooperative defense may favor group living in mass provisioning bees. Our observations and experiments suggest that parental care throughout brood development can be adaptive in mass provisioning species, supporting the predictions of assured fitness returns models.     

Recognition of caste and mating status maintains monogyny in the ant Aphaenogaster senilis

In ants dispersing through colony fission, queens mate near their natal nest and found a new society with the help of workers. This allows potential future queens to challenge the mother queen’s reproductive monopoly. Conflicts might be resolved if the mated queen signals her presence and the workers control the developmental fate of the diploid larvae (whether they develop to worker or queen). In this study we sought to determine whether, in the fission-performing ant Aphaenogaster senilis, conflicts between queens for control of the colony are resolved by the resident queen signalling her mating status. Virgin queens were less effective than newly mated queens in inhibiting queen rearing. Moreover, potential challenger queens were recognized and heavily aggressed independent of mating status. Chemical analyses showed that mating status was associated with changes in cuticular hydrocarbon and poison gland composition, but not in Dufour’s gland composition. Cuticular dimethylalkanes were identified as potential constituents that signal both caste (present in queens only) and mating status (mated queens have higher amounts). We hypothesised that pheromone emission by virgin queens did not reach the threshold needed to fully inhibit larval development into queens but was sufficiently high to stimulate overt aggression by mated queens. These findings provide evidence for the complexity of chemical communication in social insects, in which a small number of signals may have a variety of effects, depending on the context.     

Convergent development of ecological,genetic, and morphological traits in native supercolonies of the red ant Myrmica rubra

Ant supercolonies (large networks of interconnected nests) represent the most extreme form of multi-queen breeding (polygyny) and have been found across ant lineages, usually in specific long-term stable populations. Many studies on the genetic population structure and demography of ant supercolonies have been done in recent decades, but they have lacked multicolonial control patches with separated colonies headed by a single or few queens so the origin of the supercolonial trait syndrome has remained enigmatic. Here, we set out to compare sympatric supercolonial and multicolonial patches in two natural Danish populations of the common red ant Myrmica rubra. We used DNA microsatellites to reconstruct genetic colony/population structure and obtained morphological and density measurements to estimate life history and ecology covariates. We found that supercolonies in both populations completely dominated their patches whereas colonies in multicolonial patches coexisted with other ant species. Supercolony patches had very low genetic differentiation between nests, negligible relatedness within nests, and lower inbreeding than multicolonial patches, but there were no significant morphological differences. One population also had nests that approached true outbred monogyny with larger workers and males but smaller queens than in the two other social nest types. Our results suggest that once smaller colonies start to adopt additional queens, they also gain the potential to ultimately become supercolonial when the habitat allows rapid expansion through nest budding. This is relevant for understanding obligate polygyny in ants and for appreciating how and why introduced North American populations of M. rubra have recently become invasive.     

Dispersal decisions and predispersal behavior in <Emphasis Type="Italic">Polistes</Emphasis> paper wasp ‘workers’

Social insects are popular models for studying the evolution of cooperation. Casteless taxa where individuals have the flexibility to either nest alone or cooperate are particularly valuable for understanding the causes and consequences of cooperative behavior. For example, some ‘workers’ from Polistes paper wasp nests disappear from their natal colony soon after pupal emergence and nest independently. However, little is known about dispersal behavior. In this paper, I compare predispersal behavior of wasps who leave their natal colony soon after emergence with behavior of individuals who remain on the natal colony as true workers. I found that P. dominulus females with short nest tenure behave much like gynes (reproductive-destined offspring produced at the end of the season), as wasps with short nest tenure are behaviorally selfish while on the natal colony. They spend a smaller proportion of their time foraging and a larger proportion of their time resting than workers with long nest tenure. In addition, I assessed the factors that may favor early dispersal. Nest environment strongly influenced dispersal; large colonies had a smaller proportion of females with short nest tenure. Queen turnover also increased dispersal behavior perhaps because queen turnover reduces relatedness between a colony’s current and future offspring, thereby reducing the kin-selected benefits of cooperation. Therefore, casteless social insects exhibit a surprising degree of reproductive flexibility. Individuals may use information about their internal state and nest environment to optimize their behavioral strategies.     

Sanitizing the fortress: protection of ant brood and nest material by worker antibiotics

Social groups are at particular risk for parasite infection, which is heightened in eusocial insects by the low genetic diversity of individuals within a colony. To combat this, adult ants have evolved a suite of defenses to protect each other, including the production of antimicrobial secretions. However, it is the brood in a colony that are most vulnerable to parasites because their individual defenses are limited, and the nest material in which ants live is also likely to be prone to colonization by potential parasites. Here, we investigate in two ant species whether adult workers use their antimicrobial secretions not only to protect each other but also to sanitize the vulnerable brood and nest material. We find that, in both leaf-cutting ants and weaver ants, the survival of the brood was reduced and the sporulation of parasitic fungi from them increased, when the workers nursing them lacked functional antimicrobial-producing glands. This was the case for both larvae that were experimentally treated with a fungal parasite (Metarhizium) and control larvae which developed infections of an opportunistic fungal parasite (Aspergillus). Similarly, fungi were more likely to grow on the nest material of both ant species if the glands of attending workers were blocked. The results show that the defense of brood and sanitization of nest material are important functions of the antimicrobial secretions of adult ants and that ubiquitous, opportunistic fungi may be a more important driver of the evolution of these defenses than rarer, specialist parasites.     

Brood parasitism disproportionately increases nest provisioning and helper recruitment in a cooperatively breeding bird

Obligate avian brood parasites lay their eggs in nests of other species (hosts), which raise parasitic young. Parasitic nestlings are likely to influence host’s parental behaviours as they typically beg for food more vigorously than young host for a given hunger level. However, few studies have tested this idea, with conflicting results. These prior studies were largely limited to biparental hosts, but little is known about the effect of brood parasitism on parental behaviours in hosts that breed cooperatively. We followed a multimodel approach to examine the effect of brood parasitism on nest provisioning and helper recruitment in the baywing (Agelaioides badius), a cooperative breeder parasitised by screaming (Molothrus rufoaxillaris) and shiny (Molothrus bonariensis) cowbirds. Multimodel inference results indicated that feeding visits increased with nestling age, cooperative group size and number of cowbird nestlings in the brood. Brood size had little influence on feeding visits, which further suggests that baywings adjusted their provisioning effort in response to cowbird parasitism. In addition, nests parasitised artificially with shiny cowbird eggs or hatchlings recruited more helpers than unmanipulated nests having only host or screaming cowbird young. Our results provide novel evidence that brood parasitism and cooperative breeding interact in determining the levels of nest provisioning.     

Social evolution and its correlates in bees of the subgenus Evylaeus (Hymenoptera; Halictidae)

Summary Eleven behavioural characteristics of eight species of the subgenus Evylaeus were analysed using principal components analysis. The first component axis represents social level and explains over forty percent of the total variation in the data. The following characteristics are highly correlated with social level — (i) a reduction in the proportion of males in the first brood, (ii) a reduction in the proportion of workers that mate, (iii) a reduction in the proportion of workers that have developed ovaries, (iv) an increase in the mean number of workers, (v) increased contact between adults and developing brood and (vi) an increase in the size difference between queens and workers. Because these factors appear, in general, to be under the control of the queen it is argued that parental manipulation has been an important component of social evolution in these bees. The number of worker broods and the mechanism of male production are also related to social level but are less important. Nest architecture, nest defense and polygyny seem to be unrelated to social level. Variation in nest architecture may be in response to edaphic features of the substrate. The lack of any relationship between polygyny and social level implies that the semisocial route to eusociality was not the one taken. It is likely that polygyny can only occur where the substrate is suitable for the winter hibernation of sisters in their natal nest. Multivariate methods are useful in determining the relative social level of closely related halictine species.     

Hunger and food allocation among nestlings of facultatively siblicidal ospreys

Summary Previous studies of food allocation among nestlings of siblicidal birds have focussed on the proximate link between food and aggression. However, food sharing among sibs (and brood reduction) need not involve aggression. Here I examine the role of hunger (defined operationally in terms of the amount of food eaten recently) within the broader framework of passive and aggressive food-sharing among nestlings of facultatively siblicidal ospreys (Pandion haliaetus). Feeding among brood members was hierarchical; senior, dominant sibs ate more when hungry, leaving a smaller residual share of food for junior sibs. Senior sibs did not become more generous at large meals (i.e., the prospect of a large meal did not quell the selfish tendencies of senior sibs). That individuals ate more when hungry indicates a proximate link between food and the allocation of resources among siblings. Overall, aggression was infrequent and seemed to be used when junior sibs contested food allocation at meals. Aggression was diminished and food allocation was skewed toward the junior sib in broods with artificially exaggerated hatching intervals, suggesting that senior sibs were more generous when their dominant status was not threatened.Offprint requests to the present address