The cost of defeat: Capuchin groups travel further,faster and later after losing conflicts with neighbors

Although competition between social groups is central to hypotheses about the evolution of human social organization, competitive interactions among group‐mates are thought to play a more dominant role in shaping the behavior and ecology of other primate species. However, few studies have directly tested the impact of intergroup conflicts in non‐human primates. What is the cost of defeat? To address this question, the movements of six neighboring white‐faced capuchin (Cebus capucinus) social groups living on Barro Colorado Island, Panama were tracked simultaneously using an Automated Radio Telemetry System (ARTS), for a period of six months. Groups moved 13% (441 m) further on days they lost interactions compared with days they won interactions. To cover these larger distances, they traveled faster, stopped less frequently, and remained active later in the evening. Defeat also caused groups to alter their patterns of space use. Losing groups had straighter travel paths than winning groups, larger net displacements and were more likely to change their sleeping site. These results demonstrate that losing groups pay increased travel costs and suggest that they forage in low‐quality areas. They provide some of the first direct evidence that intergroup conflicts have important energetic consequences for members of competitively unsuccessful primate social groups. A better understanding of how intergroup competition impacts patterns of individual fitness is thus needed to clarify the role that this group‐level process plays in shaping the evolution of human‐ and non‐human primate behavior. Am J Phys Anthropol 152:79–85, 2013. © 2013 Wiley Periodicals, Inc.     

The disruption of dominance hierarchies by a non-native species: an individual-based analysis

We studied the effects of the exotic rainbow trout (Oncorhynchus mykiss) on the performance and the dominance hierarchy of native Atlantic salmon (Salmo salar) at the group and individual level using laboratory and semi-natural experiments. At the group level, we compared the effects of interspecific and intraspecific competition (substitutive and additive design) on behavioural responses and growth of young-of-the-year Atlantic salmon. At the individual level, the same design was used to evaluate: (1) the temporal consistency of behavioural responses, dominance hierarchy and growth rate of Atlantic salmon; (2) the pattern of correlations between behaviours; and (3) the relationship between individual growth rate and behaviour. In the laboratory, group-level analyses revealed a weak but similar effect of rainbow trout and intraspecific competition on the behaviour and growth of Atlantic salmon. In contrast, individual-based analyses demonstrated that rainbow trout (but not intraspecific competition) strongly affected behavioural strategy, dominance hierarchy and growth trajectory of individual Atlantic salmon. Specifically, behaviours, dominance status and growth rate of salmon were temporally consistent in the intraspecific environment, while these patterns were disrupted when rainbow trout were present. Similarly, we found that rainbow trout strongly affected behavioural correlations and the relationships between individual growth rate and behaviour. The semi-natural experiments confirmed these results as interspecific competition affected relationships between individual growth rate of salmon, initial weight and activity index. Overall, individual-based analyses highlighted important mechanisms that were concealed at the group level, and that may be crucial to understand ecological and evolutionary consequences of exotic species. Moreover, these results demonstrated that competition with an exotic species disrupts the hierarchical relationship among native individuals and may therefore represent a potential for a shift in selective pressure.     

Body condition,live weight and success in agonistic encounters in mixed parity groups of sows during gestation

Group housing of gestating sows benefits their welfare by allowing them freedom of movement and the opportunity for social interaction. However, social life could also bring disadvantages for individuals who receive direct aggression or are displaced from the feeder. The aim of this study was to investigate associations between social behaviour, body condition and live weight. Gestating sows (n=298) were investigated on a commercial farm. Sows were housed in mixed parity groups where two single space, ad libitum trough feeders served 12 animals. Sows were weighed, body condition scored and had their back fat layer measured at mixing, 4 weeks after insemination and again before farrowing. Social status was estimated based on the numbers of won and lost agonistic interactions at mixing and at the end of gestation. In addition, tear staining was scored before the farrowing and reproductive performance data were collected. With the aid of video recordings, 100 to 150 interactions per group were observed. Winning percentage at mixing and at the end of gestation were associated (P<0.05) and appeared relatively stable within individuals. Tear staining scores and litter sizes were not associated with winning percentage at the end of gestation. However, live weight, relative weight, body condition and back fat thickness were associated with winning percentage (P<0.05), giving heavier animals an advantage. Low winning percentage related to lower live weight gain, probably due to poorer success in competition for feed. Live weight within a mixed parity group could be used as a proxy measure for social status. Sows with low body condition score and submissive sows might need special attention with regard to group dynamics and housing to alleviate the effects of competition in group housing.     

An individual-oriented model on the emergence of support in fights, its reciprocation and exchange

Complex social behaviour of primates has usually been attributed to the operation of complex cognition. Recently, models have shown that constraints imposed by the socio-spatial structuring of individuals in a group may result in an unexpectedly high number of patterns of complex social behaviour, resembling the dominance styles of egalitarian and despotic species of macaques and the differences between them. This includes affiliative patterns, such as reciprocation of grooming, grooming up the hierarchy, and reconciliation. In the present study, we show that the distribution of support in fights, which is the social behaviour that is potentially most sophisticated in terms of cognitive processes, may emerge in the same way. The model represents the spatial grouping of individuals and their social behaviour, such as their avoidance of risks during attacks, the self-reinforcing effects of winning and losing their fights, their tendency to join in fights of others that are close by (social facilitation), their tendency to groom when they are anxious, the reduction of their anxiety by grooming, and the increase of anxiety when involved in aggression. Further, we represent the difference in intensity of aggression apparent in egalitarian and despotic macaques. The model reproduces many aspects of support in fights, such as its different types, namely, conservative, bridging and revolutionary, patterns of choice of coalition partners attributed to triadic awareness, those of reciprocation of support and 'spiteful acts' and of exchange between support and grooming. This work is important because it suggests that behaviour that seems to result from sophisticated cognition may be a side-effect of spatial structure and dominance interactions and it shows that partial correlations fail to completely omit these effects of spatial structure. Further, the model is falsifiable, since it results in many patterns that can easily be tested in real primates by means of existing data.     

Patterns of space utilization and group interactions among free-rangingMacaca mulatta

Patterns of space utilization and group interactions were studied using four social bands of free-ranging rhesus macaques at La Parguera, Puerto Rico. There were 0.26 group interactions/hr. of observation; 90% of the observed encounters occurred at feeder stations. Nineteen percent of the interactions involved actual fighting between monkeys from opposing social bands, 58% involved visual and/or vocal threats, 20% were group displacements without aggression and on 3% of the occasions the two groups mixed amicably. Group size was an important factor affecting both the dominance position of the social group and the group's use of space; larger groups were dominant, moved freely about the island, and used less of the total space than smaller lower-ranking troops. Smaller troops were continually forced to move about the island. A new troop of intermediate size added to the island colony provided an experimental test and validated these conclusions regarding group size, dominance and use of space. Patterns of space utilization for each group shifted seasonally, annually, and possibly in response to the addition of the new social troop.     

Consequences of lethal intragroup aggression and alpha male replacement on intergroup relations and home range use in tufted capuchin monkeys (Cebus apella nigritus)

In conflicts between primate groups, the resource-holding potential (RHP) of competitors is frequently related to group size or male group size, which can remain relatively constant for long periods of time, promoting stable intergroup dominance relationships. Demographic changes in neighboring groups, however, could introduce uncertainty into existing relationships. Among tufted capuchin monkeys (Cebus apella nigritus), dominant male replacement is a relatively infrequent demographic event that can have a profound effect on both the composition and size of the social group. Here, we report such a case and the consequences for home range use and intergroup aggression. Between June 2008 and August 2010, we periodically followed two neighboring groups (Macuco and Rita) in Iguazú National Park, recording daily paths (N = 143) and encounters between the groups (N = 28). We describe the events leading to a change in the male dominance hierarchy in the larger group (Macuco), which resulted in the death or dispersal of all adult males, followed by the succession of a young adult male to the dominant position. This takeover event reduced the numerical advantage in number of males between the two groups, although the ratio of total group sizes remained nearly constant. Following this shift in numerical asymmetry, the degree of escalation of intergroup aggression increased, and we observed reversals in the former intergroup dominance relationship. These changes in behavior during intergroup encounters were associated with changes in the use of overlapping areas. In the 6 months following the takeover, the area of home range overlap doubled, and the formerly dominant group's area of exclusive access was reduced by half. These results suggest that RHPin tufted capuchin monkeys is related to male group size. Furthermore, they highlight the importance of considering rare demographic events in attempts to understand the dynamics of aggression between primate groups.     

You are Not Welcome: Social Exchanges between Female Spider Monkeys (<Emphasis Type="Italic">Ateles geoffroyi</Emphasis>)

Group living leads to competition for food between group members. Two types of intragroup food competition may occur: scramble competition, in which all group members use the same resource, such that feeding opportunities are equal for everyone; and contest competition, in which some group members monopolize resources through aggression and dominance. In species in which females disperse from the natal group and immigrate into other groups, immigrant females increase group size and thus possibly food competition. Under these circumstances, other females may use aggression to discourage new females from joining the group. We assessed the distribution of aggression, embraces, and kisses among female spider monkeys (Ateles geoffroyi) in relation to group tenure. We recorded social interactions during 1688 10-min focal animal samples on 11 females in Santa Rosa, Costa Rica. We found that aggression was rare between long-term resident females and aggression rates were not higher during feeding than in other contexts, suggesting there was little contest competition. Long-term residents and less recently immigrant females showed higher aggression rates toward the most recent immigrants than toward other females, especially during the first months after a female immigrated, which coincided with the dry season. We did not find similar patterns for embrace and kiss. These results suggest that other females target aggression toward the most recent immigrants to reduce scramble competition. This finding suggests that group tenure should be included in socioecological models for species with female dispersal.     

Patterns and drivers of plant functional group dominance across the Western Hemisphere: a macroecological re‐assessment based on a massive botanical dataset

Plant functional group dominance has been linked to climate, topography and anthropogenic factors. Here, we assess existing theory linking functional group dominance patterns to their drivers by quantifying the spatial distribution of plant functional groups at a 100‐km grid scale. We use a standardized plant species occurrence dataset of unprecedented size covering the entire New World. Functional group distributions were estimated from 3 648 533 standardized occurrence records for a total of 83 854 vascular plant species, extracted from the Botanical Information and Ecology Network (BIEN) database. Seven plant functional groups were considered, describing major differences in structure and function: epiphytes; climbers; ferns; herbs; shrubs; coniferous trees; and angiosperm trees. Two measures of dominance (relative number of occurrences and relative species richness) were analysed against a range of hypothesized predictors. The functional groups showed distinct geographical patterns of dominance across the New World. Temperature seasonality and annual precipitation were most frequently selected, supporting existing hypotheses for the geographical dominance of each functional group. Human influence and topography were secondarily important. Our results support the prediction that future climate change and anthropogenic pressures could shift geographical patterns in dominance of plant functional groups, with probable consequences for ecosystem functioning. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 141–160.     

Stable group size in cooperative breeders: the role of inheritance and reproductive skew

Recent studies of reproductive skew have revealed great variationin the distribution of direct fitness among group members, yetthere have been surprisingly few attempts to explore the consequencesof such variation for stable group size, and none that takeinto account the future benefits of group membership to nonbreeders.This means that the existing theory is not suited to explainthe group size of most cooperatively breeding vertebrates andprimitively social insects in which group membership involvessubstantial future benefits. Here we model the group size ofsuch species as social queues in which nonbreeders can inherita breeding position if they outlive those ahead of them in thequeue. We demonstrate, however, that the results can be generalizedto systems in which inheritance occurs via scramble competition,rather than via a strict queue. The model predicts that stablegroup size will depend on the number of breeding positions inthe group and the mortality rates of breeders and nonbreeders,but not on the distribution of reproduction among the pool ofbreeders. This is because deaths occur at random, so that eachindividual has the same chance of surviving to reach each breedingposition. We tested a specific prediction of the model usingdata on ovarian development in the paper wasp, Polistes dominulus.We found a positive correlation between group size and the proportionof females with fully developed eggs, as predicted. Our resultsclarify the interaction between the dominance structure andsize of animal groups and add to the growing recognition ofthe potential for inheritance as a major determinant of bothindividual behavior and group-level characteristics of animalsocieties.     

Dominance hierarchies in group-living cleaning gobies: causes and foraging consequences

In species living in social groups, aggression among individuals to gain access to limiting resources can lead to the formation of stable social hierarchies. We tested whether dominance rank in social groups of sponge-dwelling cleaning gobies Elacatinus prochilos in Barbados was determined by physical attributes of individuals or by prior experience of dominance, and examined the foraging consequences of dominance rank. Intraspecific aggression within groups resulted in stable dominance hierarchies that were strongly correlated with fish length. Dominant individuals maintained exclusive territories while subordinate fish occupied broader home ranges. Larger, competitively dominant fish were able to monopolize areas inside the sponge lumen with the highest abundance of the polychaete Haplosyllis spp., a favoured prey item, and achieved the highest foraging rates. The removal of a territorial individual from large groups resulted in a domino-like effect in territory relocation of the remaining fish as individuals moved to the territory previously occupied by the individual just above them in the group hierarchy. Individuals added to existing groups generally failed to gain access to territories, despite being formerly dominant in their original groups. When given the opportunity to choose a location in the absence of larger competitors, gobies frequently preferred positions that were previously defended and that had abundant food. These results suggest that intraspecific competition for resources creates the observed dominance structures and provides support for the role of individual physical attributes in the formation and maintenance of dominance hierarchies.     

Estimating utilization distributions with kernel versus local convex hull methods

Estimates of utilization distributions (UDs) are used in analyses of home-range area, habitat and resource selection, and social interactions. We simulated data from 12 parent UDs, representing 3 series of increasingly intense space-use patterns (clustering of points around a home site, restriction of locations to a network of nodes and corridors, and dominance of a central hole in the UD) and compared the ability of kernel density estimation (KDE) and local convex hull (LCH) construction to reconstruct known UDs from samples of 10, 50, 250, and 1,000 location points. For KDE, we considered 4 bandwidth selectors: the reference bandwidth, least-squares cross-validation (LSCV), direct plug-in (DPI), and solve-the-equation (STE). For the sample sizes and UD patterns tested here, KDE achieved significantly higher volume-of-intersection (VI) scores with known parent UDs than did LCH; KDE also provided less biased home-range area estimates under many conditions. However, LCH minimized the UD volume that occurred outside the true home range boundary (V_out). Among the KDE bandwidth estimators, relative performance depended on the type and intensity of space use patterns, sample size, and the metric used to evaluate performance. Biologists should use KDE for UD and home range estimation within a probabilistic context, unless their objective is to exclude potentially unused areas by defining the area delimited by data. © 2011 The Wildlife Society.     

A sex difference in the context-sensitivity of dominance perceptions

Although dominance perceptions are thought to be important for effective social interaction, their primary function is unclear. One possibility is that they simply function to identify individuals who are capable of inflicting substantial physical harm, so that the perceiver can respond to them in ways that maximize their own physical safety. Another possibility is that they are more specialized, functioning primarily to facilitate effective direct (i.e., violent) intrasexual competition for mates, particularly among men. Here we used a priming paradigm to investigate these two possibilities. Facial cues of dominance were more salient to women after they had been primed with images of angry men, a manipulation known to activate particularly strong self-protection motivations, than after they had been primed with images of angry women or smiling individuals of either sex. By contrast, dominance cues were more salient to men after they had been primed with images of women than when they had been primed with images of men (regardless of the emotional expressions displayed), a manipulation previously shown to alter men's impressions of the sex ratio of the local population. Thus, men's dominance perceptions appear to be specialized for effective direct competition for mates, while women's dominance perceptions may function to maximize their physical safety more generally. Together, our results suggest that men's and women's dominance perceptions show different patterns of context-sensitivity and, potentially, shed new light on the routes through which violence and intrasexual competition have shaped dominance perceptions.     

Male-male Interactions in Heterosexual and All-male Wild Mountain Gorilla Groups

Variation in male dispersal and behavior patterns are components of intraspecific differences in social systems. A comparison of male behavior in different social settings can be useful for determining which behavioral mechanisms contribute to variability in social systems. Two heterosexual multimale groups and one all-male group of mountain gorillas (Gorilla gorilla beringei) were observed for over 1100 h at the Karisoke Research Centre, Rwanda. Data on proximity patterns, dominance relationships, aggression, agonistic interventions, affiliation, and homosexual behavior were compared among the males in these groups to examine the influence of female presence, sex ratio, group size, and kinship on male—male interactions. Males in the all-male group stayed closer together, affiliated more, exhibited more homosexual behavior, and were more aggressive toward each other than males in heterosexual groups. However, the males in heterosexual groups showed more wounding and more consistent dominance relationships. Kinship did not influence male-male relationships in the all-male group. The males in the heterosexual groups rarely interacted with one another; they may actively avoid close proximity to reduce aggression. Results suggest that the variable social system of mountain gorillas may be more strongly influenced by demographic factors, male-female social relationships, and male-male competition for mates than by any benefits of male-male relationships.     

Wistar大鼠幼年期丰富环境刺激对其社会竞争行为的影响

目的：探讨大鼠幼年期丰富环境经历对其成年后焦虑样行为和社会竞争行为的影响。方法：幼年期Wistar大鼠（生后21天）分为标准饲养组（对照组）和丰富环境刺激（EE）组。对照组为持续在标准实验室饲养条件下饲养;丰富环境组为生后21天至行为学检查时持续在丰富环境下饲养。在生后70天进行行为学检测：采用高架十字迷宫方法测试焦虑样行为;采用社会优势管道试验观察社会竞争性行为。结果：高架十字迷宫实验结果显示,与对照组相比,丰富环境组在开放臂的时间和次数显著增加;优势管道实验结果显示丰富环境刺激组大鼠的社会竞争力明显下降。结论：大鼠幼年期丰富环境刺激可改善大鼠的焦虑样情绪,但导致大鼠社会竞争力下降。     

Modulation of aggressive behaviour by fighting experience: mechanisms and contest outcomes

Experience in aggressive contests often affects behaviour during, and the outcome of, later contests. This review discusses evidence for, variations in, and consequences of such effects. Generally, prior winning experiences increase, and prior losing experiences decrease, the probability of winning in later contests, reflecting modifications of expected fighting ability. We examine differences in the methodologies used to study experience effects, and the relative importance and persistence of winning and losing experiences within and across taxa. We review the voluminous, but somewhat disconnected, literature on the neuroendocrine mechanisms that mediate experience effects. Most studies focus on only one of a number of possible mechanisms without providing a comprehensive view of how these mechanisms are integrated into overt behaviour. More carefully controlled work on the mechanisms underlying experience effects is needed before firm conclusions can be drawn.Behavioural changes during contests that relate to prior experience fall into two general categories. Losing experiences decrease willingness to engage in a contest while winning experiences increase willingness to escalate a contest. As expected from the sequential assessment model of contest behaviour, experiences become less important to outcomes of contests that escalate to physical fighting.A limited number of studies indicate that integration of multiple experiences can influence current contest behaviour. Details of multiple experience integration for any species are virtually unknown. We propose a simple additive model for this integration of multiple experiences into an individual's expected fighting ability. The model accounts for different magnitudes of experience effects and the possible decline in experience effects over time.Predicting contest outcomes based on prior experiences requires an algorithm that translates experience differences into contest outcomes. We propose two general types of model, one based solely on individual differences in integrated multiple experiences and the other based on the probability contests reach the escalated phase. The difference models include four algorithms reflecting possible decision rules that convert the perceived fighting abilities of two rivals into their probabilities of winning. The second type of algorithm focuses on how experience influences the probability that a subsequent contest will escalate and the fact that escalated contests may not be influenced by prior experience. Neither type of algorithm has been systematically investigated.Finally, we review models for the formation of dominance hierarchies that assume that prior experience influences contest outcome. Numerous models have reached varied conclusions depending on which factors examined in this review are included. We know relatively little about the importance of and variation in experience effects in nature and how they influence the dynamics of aggressive interactions in social groups and random assemblages of individuals. Researchers should be very active in this area in the next decade. The role of experience must be integrated with other influences on contest outcome, such as prior residency, to arrive at a more complete picture of variations in contest outcomes. We expect that this integrated view will be important in understanding other types of interactions between individuals, such as mating and predator-prey interactions, that also are affected significantly by prior experiences.     

Functions of fights in territory establishment

Fights are often observed when prospective territory owners settle in patches of vacant habitat, but the function of these fights in space acquisition is obscure. This study tests two hypotheses about the effect of fights on subsequent space use patterns: first, that settlers win space by winning fights and, second, that fights encourage the establishment of mutually exclusive home ranges between opponents (i.e., "fights make neighbors"). The behavior of juvenile Anolts aeneus lizards was recorded as they established territories in patches of habitat in the field. In support of the fights-make-neighbors hypothesis, opponents whose last aggressive interaction was a fight were six times more likely to have mutually exclusive home ranges at the end of the settlement period than were otherwise equivalent dyads whose last encounter was a chase. Contra the hypothesis that settlers win space by winning fights, most last fights ended in a draw, and there was no discernable relationship between the outcome of last fights and the subsequent space use of the contestants. These and previous analyses of settlement behavior in this species suggest that fights during the settlement period encourage the formation of symmetrical social and spatial relationships between neighboring settlers.     

Social behaviors within a group of captive female Hippopotamus amphibius

Grouping is known to occur in many species of mammals, and the structure of groups can range along a continuum from basic aggregations to complex social systems. Any social patterns that may occur within the group must be determined in order to understand the adaptive nature of the group. Female Hippopotamus amphibius are known to aggregate in the wild, but their social behaviors are still not understood. Our objective was to determine if captive female hippos display social structure within an aggregation by examining their interactions, and if kinship, familiarity, and dominance influence these interactions. Behavioral data, using continuous focal animal sampling and scan sampling, were collected on a group of captive female hippos housed at Disney’s Animal Kingdom and were used to analyze their interactions, association patterns based on kinship and familiarity, and a dominance hierarchy. Our results support the hypothesis that hippos exhibit social patterns due to the attraction to particular individuals. There were more associations between kin than non-kin and also between individuals that were more familiar. Dominance patterns were also found among these hippos. These results may aid in the general understanding of hippopotamus behavior and provide a framework for future research.     

Social organization of a stable natal group of captive Guyanese squirrel monkeys (<Emphasis Type="Italic">Saimiri sciureus sciureus</Emphasis>)

Socioecological models suggest competition for food, foraging efficiency, predation, infanticide risk, and the costs of dispersal regulate primate social structure and organization. Wild populations of squirrel monkeys (Saimiri spp.) appear to conform to the predictions of the predation/competition socioecological model (Sterck et al. in Behav Ecol Sociobiol 41:291–309, 1997) and the dispersal/foraging efficiency model (Isbell in Kinship and behavior in primates. Oxford University, New York, pp 71–108, 2004). However, squirrel monkeys in captivity are reported to maintain patterns of social behavior observed in their wild conspecifics despite different food distribution, predation risk, and dispersal options. This behavioral similarity suggests squirrel monkeys’ social behavior has limited flexibility to respond to environmental changes. In this study, we experimentally evaluated the flexibility of social behavior within a captive group of S. sciureus. First, we determined whether dominance and affiliative relationships observed under normal laboratory conditions (with abundant, widely distributed, food; no dispersal option; and no predators) better matched published reports of relationships among wild conspecifics or the predictions of the predation/competition model. Second, we made preferred food items defensible to determine whether dominance interactions would become more frequent and linear, as predicted by the model. The model correctly predicted rates of dominance behavior in both conditions and a linear hierarchy in the defensible food condition but did not predict the consistent affiliative relationships and linear dominance hierarchy observed in normal lab conditions. Although hierarchies were linear and male dominant, manipulating food distribution changed the dominant individual within each sex. Our findings suggest interaction rates adapt more rapidly than social structure to environmental changes in Saimiri and recommend caution in interpreting tests of socioecological models.     

Changes in Heart Rate Associated with Contest Outcome in Agonistic Encounters in Lobsters

Agonistic contests between lobsters housed together in a confined space progress through encounters of increasing intensity until a dominance relationship is established. Once this relationship is established, losing animals continually retreat from the advances of winners.These encounters are likely to consume much energy in both winning and losing animals. Therefore, one might expect involvement of many physiological systems before, during and after fights. Here, we report effects of agonistic encounters on cardiac frequency in winning and losing adult lobsters involved in dyadic interactions.The results show that: (i) small but significant increases in heart rate are observed upon chemical detection of a conspecific; (ii) during agonistic interactions, further increases in heart rate are seen; and (iii) ultimate winners exhibit greater increases in heart rate lasting longer periods of time compared to ultimate losers. Heart rate in winners remains elevated for at least 15 min after the contests have ended and animals have been returned to their home tanks. Reduced effects are seen in second and third pairings between familiar opponents.The sustained changes in heart rate that we observe in winning lobsters may result from hormonal modulation of cardiac function related to the change in social status brought about by contest outcome.