Symmetric fights as a measure of escalation potential in a symbiotic,territorial snapping shrimp

Summary The shrimp Alpheus armatus territorially defends the sea anemone it occupies, using as a weapon its large, specially modified snapping claw. This behavior was studied in experimental contests which were symmetric (matched individuals) with respect to sex, size, and residency. Characteristics of these contests were compared for two size-classes of male and female shrimp.There were no significant differences between these groups of shrimp in the number of bouts required to establish dominance or in the number of snaps exchanged. Large females had shorter contests than either small or large males, and losers of contests between large females were injured more frequently and more severely.This distinctiveness of large female contests could be interpreted as evidence that (i) controlling anemones is more important for large females, (ii) injuries are less important for large females, or (iii) large females lose the ability to assess one another because their contests are less frequent.If injuries are an accurate measure of the most important costs associated with fighting, then these results indicate that short contests are not necessarily the least costly, and that females can be more aggressive than males, as measured by escalation potential, in sexually selected species.     

Aggression and fighting ability are correlated in the swordtail fish Xiphophorus cortezi: the advantage of being barless

Males of the swordtail fish Xiphophorus cortezi are polymorphic for the pigment pattern vertical bars. Previous studies indicate that barred males exhibit higher levels of aggression towards males with bars than those without, while barless males fail to exhibit differential levels of aggression to either morph. In this study I matched barred and barless males for size and paired them in dyadic contests in order to determine if either morph was more dominant and if so, if dominance was the result of higher aggression levels. I found that barless males had higher bite frequencies and were able to win a majority of the contests while barred males consistently escalated to biting first, even though in most cases they were ultimately the losers. In order to determine whether the observed aggression levels and fighting abilities were inherent to being barless or a consequence of responses to the bars themselves, the dyads were re-paired once after barless males were given temporary bars and once after barred males had their bars removed. Thus, each morph encountered his opponent in both a barred and barless state. Regardless of bar state, naturally barless males continued to be more aggressive and more dominant than their barred counterparts. In addition, naturally barred males only won contests in which they bit more. These results indicate that for this species, aggression is an important component of winning contests when opponents are roughly the same size. As a result, naturally barless males as a whole appear to have higher resource holding potential (RHP) than naturally barred males of the same size because of their greater aggression levels.     

Payoff asymmetries in contests among male brown-headed cowbirds

Summary Morphometric analyses and behavioral observations of a marked population of brood parasitic brown-headed cowbirds (Molothrus ater) revealed several male attributes correlated with dominance. Males that won larger percentages of the contests in which they were involved tended to be older and larger than other males. Dominant males arrived earlier on the breeding grounds and became engaged in more contests than subordinate males. If a female was present during a contest among males, the male which she was accompanying (her consort, and mate in most cases) was more likely to win the contest than were any of his opponents. Multiple regression analyses were performed to determine any significant partial effects of these attributes on male dominance. Age and involvement in more contests exerted significant partial effects in some analyses, but only the presence of a female consort did so consistently. I suggest that the female is a resource of greater value to the male with whom she is consorting than to other males. Her value is greater in terms of present and future benefits because he has already expended some or all of the effort necessary to establish a pair bond. He will have to expend less future effort to remain her mate than would another male to become her mate. Her presence during contests induces him to persist (or escalate) and to dominate his male opponents.     

Assessment and individual recognition of opponents in the pygmy swordtails Xiphophorus nigrensis and X. multilineatus

We examined factors that determine the outcome of agonistic encounters between male pygmy swordtail fish. Xiphophorus nigrensis and X. multilineatus males formed dominance relationships based on body size in staged laboratory encounters. There was a significant negative correlation between size asymmetry and fight intensity, suggesting that males assessed size in the encounters. However, a significant proportion of the variation in fight intensity in contests that escalated to bites could not be explained by size asymmetry. Aggressive motivation may also influence the outcome of contests and could be assessed in agonistic encounters. Theory suggests that signals of aggressive intention will be evolutionarily stable if individuals can recognize opponents and encounter one another repeatedly. In addition, individual recognition is one way that dominance hierarchies can be maintained. Here we demonstrate that males from both species can recognize individuals. In addition, at least some X. nigrensis males were site-faithful in the field, suggesting males encounter the same opponents repeatedly.Communicated by G.M. Klump     

Agonistic relationships between female mountain gorillas (Gorilla gorilla beringei)

Females in some mountain gorilla groups can be ranked on the basis of decided, non-aggressive approach-retreat interactions. However, data on four gorilla groups show that females do not form clear agonistic dominance hierarchies. Most aggressive interactions between females were undecided, and most dyads had undecided dominance relationships (Fig. 2). Females ignored aggression or responded to it aggressively more often than they responded submissively (Fig. 3). Moreover, directional inconsistency in aggression (including escalated contests) was high, and in some groups females showed significant bidirectionality in initiation of aggression (i.e., the more often they received aggression from partners, the more often they directed aggression at those partners; Fig. 4). Assigning ranks on the basis of aggressive interactions has limited power to explain variation in rates of initiating aggression and in responses and outcomes to aggression. Aggression was most common during feeding, but usually did not interrupt feeding bouts. It was proportionately more common in contexts that seem to involve competition for access to males (Figs 5, 6). This is consistent with the argument that females depend crucially on male services. As predicted by current models of female primate social relationships, agonistic relationships between gorilla females are unlike those typical in cercopithecine primates. They resemble those of some other primates and some equids in which contest competition for food is relatively unimportant and ecological costs of female transfer are low.     

Rank differences in energy intake rates in white-faced capuchin monkeys, Cebus capucinus: the effects of contest competition

The effect of aggressive competition over food resources on energy intake rate is analyzed for individuals of three groups of 25–35 white-faced capuchin monkeys, Cebus capucinus, living in and near Lomas Barbudal Biological Reserve, Costa Rica. An individuals energy intake rate on a given food species was affected by its rank and the number of agonistic interactions within the feeding tree. Dominant group members had higher energy intake rates relative to subordinate group members whether or not there was agonism within the feeding tree. Low- and mid-ranked individuals had lower energy intake rates in trees with higher amounts of aggression, while energy intake rate of high-ranked individuals was not affected by the amount of aggression in the feeding tree. Energy intake was not influenced by the sex of the individual when rank was held constant statistically. Energy intake was positively correlated with total crown energy (measured in kilojoules) within the feeding tree for two of three study groups. This difference may be explained by the quality of each groups territory. Finally, high-ranked individuals are responsible for the majority of agonism within feeding trees and target middle- and low-ranked individuals equally. These findings fit the predictions of current socioecological models for within-group contest competition over food resources. The results of this study suggest that within-group competition affects energy intake rate in white-faced capuchin monkeys.     

No reduction in aggression after loss of a broodmate: a test of the brood size hypothesis

In some vertebrate species, parents create a large brood or litter then, in the event of unfavourable ecological conditions, apparently allow the number of offspring to be adaptively reduced through siblicide. But how is sibling aggression regulated so that deaths occur only in unfavourable conditions? One proposed mechanism is brood size-dependent aggression. Two experiments tested for this mechanism by reducing three-chick broods of blue-footed boobies either during or after the period of dominance hierarchy establishment. In neither experiment did aggression of the two eldest and highest ranking chicks decline after removal of the youngest broodmate, in comparison with controls. These results suggest that dominant booby chicks do not become less aggressive to each other after disappearance of their youngest broodmate and that this species does not show brood size dependent aggression. Elder blue-footed booby chicks increase their attacks on broodmates when they receive less food, and this mechanism may be sufficient to tailor brood size to food availability.     

Aggression in female Red-Winged Blackbirds: A strategy to ensure male parental investment

Summary Female Red-winged Blackbirds (Agelaius phoeniceus) are often aggressive towards conspecific females during the breeding season. We hypothesize that the function of female-female aggression in this species is to guard the nonshareable portion of the male's parental investment.The investment-guarding hypothesis predicts that a female should be more aggressive toward another female evincing interest in mating with the territory-owning male than toward a female simply perching within the male's territory. Results of mount presentations to females with active nests confirmed this prediction. Nesting females attacked a stuffed conspecific female mounted in a precopulatory, soliciting posture significantly more often than a mount in a normal, perched posture.The male's nonshereable parental care consists of provisioning his young, and most of this care is invested in the brood of his primary (first-to-nest) female. It is therefore predicted that primary females should be more aggressive than secondary (later-nesting) females. Female mount presentations also confirmed this prediction. Primary females attacked the soliciting mount significantly more often than secondary females.     

Signaling effort does not predict aggressiveness in male spring field crickets

The relationship between signaling and aggression is still unclear despite several decades of research. However, there is a growing interest in studying signals that predict aggressive behavior or fighting ability. The goals of our study were threefold: we investigated the relationship between signaling effort and aggression, the relationship between body condition and aggression, and the effect of fighting experience on subsequent signaling behavior in wild-caught and laboratory-reared male spring field crickets (Gryllus veletis). We found that aggressive behavior was not related to signaling effort, body size, or body condition. For contest winners, wild-captured males were more aggressive than laboratory-reared males. Signaling effort was highly repeatable within individuals, but aggressive behavior had low repeatability. We found no evidence for a winner or loser effect on signaling; there was no change in signaling effort when we compared contest winners and losers before and after they participated in aggressive contests. Long-distance acoustic signaling and aggressive behavior appear to be independent of one another in spring field crickets, perhaps serving different functions in female attraction and male–male competition, respectively.     

The role of size and aggression in intrasexual male competition in a social lizard species, Egernia whitii

Competition between males is a key component of the agonistic intrasexual interactions that influence resource acquisition, social system dynamics, and ultimately reproductive success. Sexual selection theory predicts that traits that enhance success in intrasexual competition (particularly male–male competition) should be favored. In vertebrates, this often includes body size and aggression, with larger and/or more aggressive males outcompeting smaller or less aggressive conspecifics. The majority of studies consider aggression as a flexible trait which responds to local social or environmental conditions. However, aggression frequently shows considerable within-individual consistency (i.e., individuals have identifiable aggressive behavioral types). Little is known about how such consistency in aggression may influence competition outcomes. We integrated a detailed field study with a laboratory experiment to examine how a male’s aggressive phenotype and his size influence competitive interactions in Egernia whitii, a social lizard species which exhibits strong competition over resources (limited permanent shelter sites and basking sites). Individual aggression and size did not predict competition outcome in the laboratory nor did they predict home range size, overlap, or reproductive success in the field. However, winners of laboratory trial contests maintained consistent aggressive phenotypes while consistency in aggression was lost in losers. We suggest that aggression may be important in other functional contexts, such as parental care, and that alternative traits, such as fighting experience, may be important in determining competition outcome in this species.     

Surplus territory contenders in male red-winged blackbirds: where are the desperados?

Summary We tested whether payoff asymmetries could explain why floater red-winged blackbirds seldom escalate contests to acquire territories. We removed territorial male \ldowners\rd and allowed floater replacements to claim territories. We then released owners to see how three currencies of payoff asymmetries (replacement occupancy) owners' likelihood and speed of reclaiming their territories (owner success). Neither the duration of an owner's internment nor the amount of time that a floater replacement had held the territory affected owner success (Figs. 1 and 2). Owner success was also not affected by the number of neighbors that they had (Fig. 3). Finally, owner success was the same irrespective of whether or not he was the likely sire of offspring on the territory, or the size of his harem (Fig. 4). Although these results are consistent with the proposal that payoff asymmetries will be irrelevant in contests for valuable resources, they are inconsistent with the proposal's corollary that excluded individuals should become \lddesperados\rd and escalate or even fight to the death in contests for those resources. Expected payoffs for passive acquisition of territories in this species may be higher than from a more aggressive desperado strategy.Correspondence to: P.J. Weatherhead     

Pitch shifts and song structure indicate male quality in the dawn chorus of black-capped chickadees

The fee-bee song of male black-capped chickadees (Poecile atricapillus) is considered a single-type song that singers transpose up and down a continuous frequency range. While the ability to shift song pitch in this species provides a mechanism for song matching as an aversive signal in male-male territorial song contests, the functional significance of this behaviour during the solo performances of males during the dawn chorus is unclear. We analysed the dawn chorus songs and singing behaviour of males whose winter-flock dominance status we determined. We used correlation analysis to show that pitch shifts were accompanied by changes to other fine structural characteristics in song, including temporal and relative amplitude parameters. We also found that songs of socially dominant males and songs of their most subordinate flockmates could be distinguished using these methods by the way they performed a between-note frequency measure accompanying pitch shifts. That is, a ratio measure of the internote frequency interval remained constant for songs of high-ranking birds despite changes in absolute pitch, while low-ranking males sang a smaller ratio as they shifted to higher absolute pitches. These findings identify previously unrecognised variation in the songs of black-capped chickadees. More importantly, they indicate a mechanism by which pitch shifting during the dawn chorus of black-capped chickadees could provide a reliable indicator of relative male quality.Communicated by I. Hartley     

The effect of intrauterine position on the survival,reproduction and home range size of female house mice (Mus musculus)

Summary In laboratory studies using albino house mice, a female's prior intrauterine position can affect many postnatal physiological, morphological and behavioral characteristics. Females flanked by males in utero (2M females) exhibit more aggressive dominance than females flanked by females (OM females). Thus, wild 2M females may be most successful during peak population densities when their aggressive nature would allow them to displace other females from limited resources. 2M and 0M females and males delivered by cesarean section were individually marked and released as young adults on two occasions onto a highway island (the area enclosed by exit and entrance ramps at an interchange) to determine whether 2 M females have a competitive advantage over 0 M females in the field. Males were included to create realistic population structure; their intrauterine position was not a treatment. Feeding stations afforded individuals an opportunity to exhibit their dominance by maintaining home-ranges at or near the stations. The populations were monitored by periodic live-trapping and reproductive success was determined using field body weights and by post-mortem examination for uterine implantation scars. Survival and capture rates were estimated, using a modified Jolly-Seber markrecapture program, for each of four intervals between trapping occasions over the course of 7 weeks. There were no overall differences in survivorship between 2M and 0M females, neither type of female was caught more frequently at feeding stations and they did not differ in measures of reproductive success. However, 2M females had significantly larger home-range sizes than 0M females and thus space use may be a trait masculinized by prior intrauterine position. Although there are a number of life-history characteristics that differ between 0M and 2M females in the laboratory that we did not test specifically in the field, our findings and other features of wild house mouse biology suggest that prior intrauterine position does not have a strong effect on survival and reproduction in the wild. Offprint requests to: W.J. Zielinski at the present address     

Patterns of participation and free riding in territorial conflicts among ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Cooperation in animal social groups may be limited by the threat of free riding, the potential for individuals to reap the benefits of other individuals actions without paying their share of the costs. Here we investigate the factors that influence individual contributions to group-level benefits by studying individual participation in territorial defense among female ringtailed lemurs (Lemur catta). To control for potentially confounding factors, particularly group size, we studied two semi-free-ranging groups at the Duke University Primate Center. First, we used a combination of experimental and observational methods to investigate the costs and benefits of territorial defense for individual lemurs. We found three indications of costs: physical contact occurred during inter-group encounters, participation in territorial defense was negatively correlated with ambient temperature, and rates of self-directed behaviors increased during encounters. Benefits were more difficult to quantify, but observational and experimental tests suggested that individuals shared the gains of territorial defense by foraging in defended territories. Thus, during experiments in which one of the groups was prevented from defending its territory, the free-ranging group made more frequent incursions into the other groups territory. Second, we examined variation in participation in territorial defense. Individuals varied significantly in their rates of aggression and genital marking during inter-group encounters. The extensive variation documented among individuals was partially accounted for by dominance rank, kinship and patterns of parental care. However, we found no evidence to suggest that participation was enforced through punishment (policing) or exchange of benefits involving grooming. In conclusion, this study provides further insights into cooperative behavior in mammalian social groups by revealing how the costs and benefits of territoriality influence patterns of individual participation in the context of shared (collective) goods.Communicated by P. Kappeler     

Kinship and aggression: do house sparrows spare their relatives?

Kin-selection theory predicts that relatedness may reduce the level of aggression among competing group members, leading to indirect fitness benefits for kin-favoring individuals. To test this hypothesis, we investigated whether relatedness affects aggressive behavior during social activities in captive house sparrow (Passer domesticus) flocks. We found that sparrows did not reduce their aggression towards kin, as neither the frequency nor the intensity of fights differed between close kin and unrelated flock-mates. Fighting success was also unrelated to kinship and the presence of relatives in the flock did not influence the birds’ dominance rank. These results suggest that the pay-offs of reduced aggression towards kin may be low in non-breeding flocks of sparrows, e.g. due to competition among relatives as predicted by a recent refinement of kin-selection theory. Our findings indicate that the significance of kin selection may be restricted in some social systems such as winter aggregations of birds. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The role of body size and fighting experience in predicting contest behaviour in the black field cricket, <Emphasis Type="Italic">Teleogryllus commodus</Emphasis>

Body size strongly predicts fighting behaviour and outcome in many species, with the larger opponent usually winning contests. However, recent fighting experience can have a strong influence on the establishment of dominance hierarchies, with recent winners being more likely to win subsequent contests, while recent losers are more likely to lose. Recent fighting experience therefore has the potential to modify the effect of body size in determining contest behaviour and outcome. Here, we investigate whether recent fighting experience weakens the role of body size in predicting contest behaviour in the black field cricket, Teleogryllus commodus. We compared the role of body size in determining contest outcome during initial non-physical encounters and escalated aggressive physical encounters (grapples), as well as the probability of escalation occurring, in contests involving either a naïve or experienced smaller male against a naïve larger male. We found that recent fighting experience only affected contest outcome during non-physical encounters. Once a contest had escalated into grappling, the effect of previous wins and losses was no longer apparent and body size strongly predicted contest outcome. Thus, once males can directly assess their opponent's fighting ability, recent fighting experience did not alter the effect of body size on contest behaviour and outcome.     

Dominance and fertility in a functionally monogynous ant

Summary Colonies of some leptothoracine ants may contain several inseminated but sterile females in addition to a single, fertile queen (functional monogyny). We here report the first observations on the behavior of these supernumerary females in the nearctic ant Leptothorax sp.A, a species belonging to the L. muscorum complex.In four colonies, each with up to eight intermorphic females, ritualized or openly aggressive interactions between individuals were observed, similar to those among workers of some other leptothoracine ants. The responses of individual females during encounters with nestmates apparently reflect the existence of linear dominance hierarchies. In each colony, the highest ranking individual was fed and groomed significantly more often than other females, and was the only one to oviposit after hibernation and to become fully physogastric during the first weeks of spring. When these -females were removed from three colonies, several other females started to lay eggs. However, in each colony only the highest ranking individual remained fertile; the others either were pushed out of the nest and finally killed by the workers, or their ovaries degenerated again.Aggressive interactions among females may also play an important role in the foundation of new colonies either by inducing intermorphs to leave the maternal nest to found new colonies solitarily or by inducing colony fission. Offprint requests to: J. Heinze at his new address     

When to be neighbourly: differential agonistic responses in the lizard Platysaurus broadleyi

Differential treatment of neighbours and strangers (the dear enemy phenomenon) can reduce the costly effects of territory defence. Individual recognition, and by extension the dear enemy phenomenon, was examined in the cordylid lizard Platysaurus broadleyi. I had no knowledge of familiarity between individuals and therefore classed intruders as neighbours or non-neighbours and tested for differences in agonistic response consistent with the dear enemy phenomenon. In the dear enemy phenomenon, levels of aggression are inversely related to degree of familiarity such that residents are more tolerant of immediate neighbours than distant territory holders. A manipulative field experiment in conjunction with field observations of known territory holders revealed that resident males allowed neighbours to approach more closely than non-neighbours before challenging them, residents were more aggressive towards non-neighbours than neighbours, contests between neighbours were significantly longer than between neighbours and non-neighbours and contests between residents and non-neighbours were likely to result in a win for the resident, while neighbours frequently drew contests. These results suggest a level of recognition consistent with the dear enemy phenomenon. I also manipulated front leg colour to test the hypothesis that it alone could serve as a cue for individual recognition. I found no support for this hypothesis and suggest that multiple cues operate in individual recognition. Received: 4 March 1999 / Received in revised form: 11 April 1999 / Accepted: 19 April 1999     

The appeasement effect of sterility signaling in dominance contests among Bombus terrestris workers

The establishment of dominance hierarchies through aggressive interactions is very common in insect societies. In many cases, it is also mediated through pheromone emissions that enable individuals to evaluate the reproductive quality and level of aggressiveness of the dominant individual, thereby reducing the number and intensity of costly fights. Here, we studied these processes in the primitively eusocial bee Bombus terrestris, using a paired bee system. Specifically, we investigated the behavioral, reproductive, and pheromonal correlates of dominance establishment. Workers were shown to establish dominance hierarchies using overt aggression within 3–4 days. Thereafter, the aggression drastically decreased, and dominance was maintained mostly by ritualized agonistic behavior. The behaviorally dominant bee lost the ester compounds that workers produce in their Dufour's gland (the so-called “sterility signal”) concomitantly with the development of her ovaries. The other bee announced as subordinate by continuously producing high amounts of those esters. The hypothesis that sterility signaling serves as an appeasement signal to pacify the dominant bees is supported by the negative correlation found between the proportion of these esters and the level of aggression that the subordinate received from the dominant worker. Physical interactions, and presumably also the ensuing overt aggression between the bees, were essential for the above pheromonal change to take place and enabled the dominant workers to develop their ovaries and to lay eggs. The subordinate bee’s signaling of non-reproductive status may minimize energy expenditure in costly fights and help stabilize the reproductive division of labor among workers.     

A game model for dominance relations among group-living animals

We present here an attempt to understand behaviors of dominant individuals and of subordinate individuals as behavior strategies in an asymmetric “hawk-dove” game. We assume that contestants have perfect information about relative fighting ability and the value of the resource. Any type of asymmetry, both relevant to and irrelevant to the fighting ability, can be considered. It is concluded that evolutionarily stable strategies (ESSs) depend on the resource value (V), the cost of injury (D), and the probability that the individual in one role will win (x). Different ESSs can exist even when values of V, D, and x are the same. The characteristics of dominance relations detected by observers may result from the ESSs that the individuals are adopting. The model explains some characteristics of dominance relations, for example, the consistent outcome of contests, the rare occurrence of escalated fights, and the discrepancy between resource holding potential (RHP) and dominance relations, from the viewpoint of individual selection. Received: 7 May 1997 / Accepted after revision: 17 October 1997