Dominance hierarchy - Knowledge (XXG)

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with low-ranking group members being able to influence which group member is leader and which one is not. Sometimes dominant animals must maintain alliances with subordinates and grant them favours to receive their support in order to retain their dominant rank. In chimpanzees, the alpha male may need to tolerate lower-ranking group members hovering near fertile females or taking portions of his meals. Other examples can include Muriqui monkeys. Within their groups, there is abundant food and females will mate promiscuously. Because of this, males gain very little in fighting over females, who are, in turn, too large and strong for males to monopolize or control, so males do not appear to form especially prominent ranks between them, with several males mating with the same female in view of each other. This type of mating style is also present in manatees, removing their need to engage in serious fighting. Among female elephants, leadership roles are not acquired by sheer brute force, but instead through seniority, and other females can collectively show preferences for where the herd can travel. In hamadryas baboons, several high-ranking males will share a similar rank, with no single male being an absolute leader. Female bats also have a somewhat fluid social structure, in which rank is not strongly enforced. Bonobos are matriarchal, yet their social groups are also generally quite flexible, and serious aggression is quite rare between them. In olive baboons, certain animals are dominant in certain contexts, but not in others. Prime age male olive baboons claim feeding priority, yet baboons of any age or sex can initiate and govern the group's collective movements.

217:, the offspring of high-ranking individuals have better fitness and thus an increased rate of survival. This is most likely a function of two factors: The first is that high-ranking males mate with high-ranking females. Assuming their high rank is correlated with higher fitness and fighting ability, this trait will be conferred to their offspring. The second factor is that higher-ranking parents probably provide better protection to their offspring and thus ensure higher survival rates. Amongst rhesus macaques, higher-ranking males sired more offspring, though the alpha male was never the one to sire the most offspring, with that instead being a high-ranking but not top male. The complex relationship between rank and reproduction in this species is likely explained by the fact that rhesus macaques queue, rather than fight, for dominance, meaning that the alpha male is not necessarily the strongest or most attractive male. 277:

friendly behaviours, and hostile behaviours are predicted to be reciprocated with similar, hostile behaviours. When an individual acts in a dominant, authoritative manner in a group, this behaviour tends to prompt submissive responses from other group members. Similarly, when group members display submissive behaviour, others feel inclined to display dominant behaviours in return. Tiedens and Fragale (2003) found that hierarchical differentiation plays a significant role in liking behaviour in groups. Individuals prefer to interact with other group members whose power, or status behaviour complements their own. That is to say, group members who behave submissively when talking to someone who appears to be in control are better liked, and similarly individuals who display dominant behaviours (e.g., taking charge, issuing orders) are more liked when interacting with docile, subservient individuals.

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resources such as mates and food. Age, intelligence, experience, and physical fitness can influence whether or not an individual deems it worthwhile to pursue a higher ranking in the hierarchy, which often comes at the expense of conflict. Hierarchy results from interactions, group dynamics, and sharing of resources, so group size and composition affect the dominance decisions of high-ranking individuals. For example, in a large group with many males, it may be difficult for the highest-ranking male to dominate all the mating opportunities, so some mate sharing is likely to exist. These opportunities available to subordinates reduce the likelihood of a challenge to the dominant male: mating is no longer an all-or-nothing game and the sharing is enough to placate most subordinates. Another aspect that can determine dominance hierarchies is the environment. In populations of Kenyan

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maintains discipline and coherence of the flock. He leads the group but shares leadership on a foraging expedition with a mature she-goat who will normally outlast a succession of dominant males. However, earlier work showed that leadership orders in goats were not related to age or dominance. In sheep, position in a moving flock is highly correlated with social dominance, but there is no definite study to show consistent voluntary leadership by an individual. In birds, dominant individuals preferentially select higher perches to put themselves in the best position to detect and avoid predators, as well as to display their dominance to other members of their own species. It has been suggested that decision-taking about the actions of the group is commonly dissociated from social dominance.

375:, the study of optimal strategies during pair-wise conflict, was grounded in the false assumption that animals engaged in conflict were of equal fighting ability. Modifications, however, have provided increased focus on the differences between the fighting capabilities of animals and raised questions about their evolutionary development. These differences are believed to determine the outcomes of fights, their intensity, and animal decisions to submit or continue fighting. The influence of aggression, threats, and fighting on the strategies of individuals engaged in conflict has proven integral to establishing social hierarchies reflective of dominant-subordinate interactions. 983:) can only lay unfertilised eggs. These eggs are in general viable, developing into males. A worker that performs reproduction is considered a "cheater" within the colony, because it leaves disproportionately more descendants than do its sisters and mother. The advantage of remaining functionally sterile is only accomplished if every worker assume this "compromise". When one or more workers start reproducing, the "social contract" is destroyed and the colony cohesion is dissolved. Aggressive behavior derived from this conflict may result in the formation of hierarchies, and attempts of reproduction by workers are actively suppressed. In some wasps, such as 258:, high-ranking individuals experience higher resting metabolic rates and therefore need to consume more food in order to maintain fitness and activity levels than do subordinates in their groups. The energetic costs of defending territory, mates, and other resources can be very consuming and cause high-ranking individuals, who spend more time in these activities, to lose body mass over long periods of dominance. Therefore, their physical condition decreases the longer they spend partaking in these high-energy activities, and they lose rank as a function of age. 291:

the pecking order keeps both parties from incurring the costs of a prolonged fight. In hens, it has been observed that both dominants and subordinates benefit from a stable hierarchical environment, because fewer challenges means more resources can be dedicated to laying eggs. In groups of highly related individuals, kin selection may influence the stability of hierarchical dominance. A subordinate individual closely related to the dominant individual may benefit more genetically by assisting the dominant individual to pass on their genes.

929:, only one to few individuals members of a colony can reproduce, whereas the other colony members have their reproductive capabilities suppressed. This conflict over reproduction in some cases results in a dominance hierarchy. Dominant individuals in this case are known as queens and have the obvious advantage of performing reproduction and benefiting from all the tasks performed by their subordinates, the worker caste (foraging, nest maintenance, nest defense, brood care and thermal regulation). According to 55: 1022:. In this case, another advantage of maintaining a hierarchy is to prolong the colony lifespan. The top ranked individuals may die or lose fertility and "extra queens" may benefit from starting a colony in the same site or nest. This advantage is critical in some ecological contexts, such as in situations where nesting sites are limited or dispersal of individuals is risky due to high rates of predation. This polygynous behavior has also been observed in some eusocial bees such as 140: 1015:, however, the dominant-subordinate context in the yellow paper wasps is directly related to the exchange of food. Future foundresses within the nest compete over the shared resources of nourishment, such as protein. Unequal nourishment is often what leads to the size differences that result in dominant-subordinate position rankings. Therefore, if during the winter aggregate, the female is able to obtain greater access to food, the female could thus reach a dominant position. 677:

suppression, researchers removed the queen from the colony but did not remove her bedding. They reasoned that if a primer pheromones were on the bedding then the sub-dominant's reproductive function should continue to be suppressed. Instead however, they found that the sub-dominants quickly regained reproductive function even in the presence of the queen's bedding and thus it was concluded that primer pheromones do not seem to play a role in suppressing reproductive function.

783: 324:, young males take advantage of their underdeveloped secondary sex characteristics to engage in sneak copulations. These young males mimic all the visual signs of a female lizard in order to successfully approach a female and copulate without detection by the dominant male. This strategy does not work at close range because the chemical signals given off by the sneaky males reveal their true nature, and they are chased out by the dominant. 224:. The dwarf mongoose lives in a social system with one dominant pair. The dominant female produces all or almost all of the offspring in the living group, and the dominant male has first access to her during her oestrus period. In red deer, the males who experienced winter dominance, resulting from greater access to preferred foraging sites, had higher ability to get and maintain larger harems during the mating season. 500:. In general, aggressive interactions are ritualistic and involve antennation (drumming), abdomen curling and very rarely mandible bouts and stinging. The winner of the interaction may walk over the subordinated, that in turn assumes a prostrated posture. To be effective, these regulatory mechanisms must include traits that make an individual rank position readily recognizable by its nestmates. The composition of the 5021: 917:

elder, stronger chick almost always becomes the dominant chick. During times of food shortage, the dominant chick often kills the subordinate chick by either repeatedly pecking or by ousting the younger chick from the nest. The brood hierarchy makes it easier for the subordinate chick to die quietly in times of food scarcity, which provides an efficient system for booby parents to maximize their investment.

5045: 874: 841:, as androgens are associated with aggressive behavior in young females. Females that were "exposed to greater concentrations of maternal late in fetal development were less likely to be aggressed against postnatally, whereas females that were...exposed to greater concentrations of maternal ...were more likely to receive aggression postnatally". Dominance rank in female 1097: 5033: 262:

immunosuppressant activity, which reduces survival and presents opportunities for parasitic infestation and other health risks. This reduced fitness due to the alpha position results in individuals maintaining high rank for shorter periods of time and having an overall reduced health and longevity from the physical strain and costs of the position.

405:, for example, engage in exhausting roaring contests to exhibit their strength. However, such an activity would impose more costs than benefits for unfit stags, and compel them to retreat from the contest. Larger stags have also been known to make lower-frequency threat signals, acting as indicators of body size, strength, and dominance. 396:. If one participant believes he is the resident of the territory, he will win when the opponent is weaker or food is scarce. However, if both shrews believe they are the true territory holder, the one with the greater need for food, and therefore, the one that values the resource more, is most likely to win. 302:

and stress; over a long period of time, this can lead to decreased fitness. The lowest-ranking males also had high stress levels, suggesting that it is the beta males that gain the most fitness, avoiding stress but receiving some of the benefits of moderate rank. The mating tactics of savanna baboons

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Considerable evidence now indicates that in humans social stratification is principally based jointly on dominance (coercive capacity based on strength, threat, and intimidation) and prestige (persuasive capacity based on skills, abilities, and knowledge). Although intimidation can beget compliance,

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animals as territory owners are often dominant over all others in their own territory but submissive elsewhere, or dependent on the resource. Even with these factors held constant, perfect dominance hierarchies are rarely found in groups of any great size, at least in the wild. Dominance hierarchies

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brood of two chicks always has a dominance hierarchy due to the asynchronous hatching of the eggs. One egg is laid four days before the other, and incubation starts immediately after laying, so the elder chick is hatched four days before the younger chick and has a four-day head start on growth. The

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prefrontal cortex, one processing judgment cues and the other processing status of an individual. Other studies have determined that lesions to the prefrontal cortex (when the area is severed to disrupt functioning to observe its role in behavior) led to deficits in processing social hierarchy cues,

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have individual "facial badges" that permit them to recognize each other and to identify the status of each individual. Individuals whose badges were modified by painting were aggressively treated by their nestmates; this makes advertising a false ranking status costly, and may help to suppress such

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In wild male baboons, the highest-ranking male, also known as the alpha, experiences high levels of both testosterone and glucocorticoid, which indicates that high-ranking males undergo higher levels of stress which reduces fitness. Reduced health and longevity occurs because these two hormones have

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opportunities. Rather than fighting each time they meet, individuals of the same sex establish a relative rank, with higher-ranking individuals often gaining more access to resources and mates. Based on repetitive interactions, a social order is created that is subject to change each time a dominant

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Although many group-living animal species have a hierarchy of some form, some species have more fluid and flexible social groupings, where rank does not need to be rigidly enforced, and low-ranking group members may enjoy a wider degree of social flexibility. Some animal societies are "democratic",

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selection within groups, where competition between individuals results in differential access to resources and mating opportunities. This can be mapped across a spectrum of social organization ranging from egalitarian to despotic, varying across multiple dimensions of cooperation and competition in

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In many monogamous bird species, the dominant pairs tend to get the best territories, which in turn promote offspring survival and adult health. In dunnocks, a species of bird that experiences many mating systems, sometimes individuals will form a group that will have one dominant male who achieves

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secreted by the queen cause direct suppression of these vital reproductive hormones and functions however current evidence suggests that it is not the secretion of pheromones which act to suppress reproductive function but rather the queen's extremely high levels of circulating testosterone, which

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Subordinate individuals often demonstrate a huge reproductive disadvantage in dominance hierarchies. Among brown hyenas, subordinate females have less opportunity to rear young in the communal den, and thus have fewer surviving offspring than do high-ranking individuals. Subordinate males copulate

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Subordinate individuals suffer a range of costs from dominance hierarchies, one of the most notable being reduced access to food sources. When a resource is obtained, dominant individuals are first to feed as well as taking the longest time. Subordinates also lose out in shelter and nesting sites.

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Being subordinate offers a number of benefits. Subordination is beneficial in agonistic conflicts where rank predicts the outcome of a fight. Less injury will occur if subordinate individuals avoid fighting with higher-ranking individuals who would win a large percentage of the time — knowledge of

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As expected, the individual who emerges triumphant is rewarded with the dominant status, having demonstrated their physical superiority. However, the costs incurred to the defeated, which include loss of reproductive opportunities and quality food, can hinder the individual's fitness. In order to

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display a unique white plumage; the higher the percentage of the crown that consists of white feathers, the higher the status of the individual. For other animals, the time spent in the group serves as a determinant of dominance status. Rank may also be acquired from maternal dominance rank. In

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The interpersonal complementarity hypothesis suggests that obedience and authority are reciprocal, complementary processes. That is, it predicts that one group member's behaviours will elicit a predictable set of actions from other group members. Friendly behaviours are predicted to be met with

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In primates, a well-studied group, high rank brings reproductive success, as seen in a 1991 meta-analysis of 32 studies. A 2016 study determined that higher status increased reproductive success amongst men, and that this did not vary by type of subsistence (foraging, horticulture, pastoralism,

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Given the benefits and costs of possessing a high rank within a hierarchical group, there are certain characteristics of individuals, groups, and environments that determine whether an individual will benefit from a high rank. These include whether or not high rank gives them access to valuable

1084:. Linear ranking systems, or "pecking orders", which tend to fall in between egalitarianism and despotism, follow a structure where every member of the group is recognized as either dominant or submissive relative to every other member. This results in a linear distribution of rank, as seen in 676:

on the colony and thus "scare" the other mole-rats into submission. Research has shown that removal of the queen from the colony allows the reestablishment of reproductive function in sub-dominant individuals. To see if a priming pheromone secreted by the queen was indeed causing reproductive

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males have more access to fertile females and consequently partake in most of the matings within the group; in one population, three males were responsible for over 75% of matings. In this population, males often vary in rank. As their rank improves, they gain more exclusive time with fertile

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For many animal societies, an individual's position in the dominance hierarchy corresponds with their opportunities to reproduce. In hierarchically social animals, dominant individuals may exert control over others. For example, in a herd of feral goats it is a large male that is dominant and

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when they transition from non-breeding to breeding seasons, increasing female aggression. Androgens are greater in pregnant female lemurs, which suggests that organizational androgens might influence the developing offspring. Organizational androgens play a role in "explaining female social

620:. The size of the oocytes plays a significant role in establishing dominance in the paper wasp. Foundresses treated with 20-hydroxyecdysone showed more dominance than did those treated with juvenile hormone, so 20-hydroxyecdysone may play a larger role in establishing dominance (Roseler 408:

Engaging in agonistic behavior can be very costly and thus there are many examples in nature of animals who achieve dominance in more passive ways. In some, the dominance status of an individual is clearly visible, eliminating the need for agonistic behavior. In wintering bird flocks,

632: 857:), androgens (i.e. specifically, androstenedione and testosterone) are "implicated in the organization and activation of...nonreproductive behavioral traits, including aggression, social dominance, rough-and-tumble play, and scent marking". For aggressively dominant female 401:

minimize these losses, animals generally retreat from fighting or displaying fighting ability unless there are obvious cues indicating victory. These often involve characteristics that provide an advantage during agonistic behavior, such as size of body, displays, etc.

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Female-biased dominance occurs rarely in mammals. It occurs when all adult males exhibit submissive behavior to adult females in social settings. These social settings are usually related to feeding, grooming, and sleeping site priority. It is observed consistently in

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Bonnatti, Vanessa; Luz Paulino Simões, Zilá; Franco, Fernando Faria; Tiago, Mauricio (3 January 2014). "Evidence of at least two evolutionary lineages in Melipona subnitida (Apidae, Meliponini) suggested by mtDNA variability and geometric morphometrics of forewings".

241:, the queen seeks to maintain reproductive success by preventing workers from caring for their cells, pushing or hitting them using her antennae. Workers display aggression towards males, claiming priority over the cells when males try to use them to place eggs. 707:

are a result of social factors, particularly when the hierarchy is in transition, perhaps resulting in increased aggression and confrontation. As a result, the dominant individual fights more and has elevated glucocorticoids during this period. Field studies of

164:, high-ranking females have higher foraging success when the food resources are clumped, but when food is distributed throughout an area they lose their advantage, because subordinate females can acquire food with less risk of encountering a dominant female. 316:, which have a social order involving one dominant male controlling most access to mates, display a behavior known as sneak copulation. While one male at a carcass has a 5:1 mating advantage, subordinate males will tempt females away from the carcass with 196:. The dominant individuals in these groups fill themselves up first and fill up more quickly, so they spend less time foraging, which reduces the risk of predation. Thus they have increased survival because of increased nutrition and decreased predation. 118:

is an individual's preferential access to resources over another based on coercive capacity based on strength, threat, and intimidation, compared to prestige (persuasive capacity based on skills, abilities, and knowledge). A dominant animal is one whose

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suggesting this area is important in regulating this information. Although the prefrontal cortex has been implicated, there are other downstream targets of the prefrontal cortex that have also been linked in maintaining this behavior. This includes the

624:, 1984). Subsequent research however, suggests that juvenile hormone is implicated, though only on certain individuals. When injected with juvenile hormone, larger foundresses showed more mounting behaviors than smaller ones, and more oocytes in their 102:. Different types of interactions can result in dominance depending on the species, including ritualized displays of aggression or direct physical violence. In social living groups, members are likely to compete for access to limited resources and 908:

Wild and feral chickens form relatively small groups, usually including no more than 10 to 20 individuals. It has been shown that in larger groups, which is common in farming, the dominance hierarchy becomes less stable and aggression increases.

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Subordinate animals engage in a number of behaviors in order to outweigh the costs of low rank. Dispersal is often associated with increased mortality and subordination may decrease the potential benefits of leaving the group. In the

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through lesion studies in rats and primates which led to disruption in hierarchy, and can affect the individual negatively or positively depending on the subnuclei that is targeted. Additionally, the dorsal medial PFC-medial dorsal

648:) similarly have a dominance hierarchy dependent on the highest ranking female (queen) and her ability to suppress critically important reproductive hormones in male and female sub-dominants. In sub-dominant males, it appears that 310:, however, subordinates occasionally win a fight for a female, and they father 44% of the lambs born in the population. These sheep live in large flocks, and dominance hierarchies are often restructured each breeding season. 3604:

Grebe, Nicholas M.; Fitzpatrick, Courtney; Sharrock, Katherine; Starling, Anne; Drea, Christine M. (September 2019). "Organizational and activational androgens, lemur social play, and the ontogeny of female dominance".

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it has been shown that subordinate individuals, given the opportunity to desert, often do not due to the risk of death and the low possibility that they would establish themselves as dominant members in a new group.

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nuclei (a neurotransmitter involved with many behaviors including reward and learning). In manipulation studies of this region, there were changes in fighting and affiliative behavior in primates and crustaceans.

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of social insects is the clue used by nestmates to recognize each other in the colony, and to discover each insect's reproductive status (and rank). Visual cues may also transmit the same information. Paper wasps

933:, the reproduction costs of the worker caste are compensated by the contribution of workers to the queen's reproductive success, with which they share genes. This is true not only for the popular social insects ( 249:

There are costs to being of a high rank in a hierarchical group which offset the benefits. The most common costs to high-ranking individuals are higher metabolic rates and higher levels of stress hormones. In

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between. Conflict can be resolved in multiple ways, including aggression, tolerance, and avoidance. These are produced by social decision-making, described in the "relational model" created by the zoologist

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there are many possible queens that inhabit a nest, but only one can be queen at a time. When a queen dies the next queen is selected by an age-based dominance hierarchy. This is also true in the species

728:, a region involved with decision making and social behavior. High social rank in a hierarchical group of mice has been associated with increased excitability in the medial prefrontal cortex of pyramidal 469:

this is mainly achieved by aggressive interactions between the potential reproductive females. In eusocial insects, aggressive interactions are common determinants of reproductive status, such as in the

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and behavior and thus redirects the sub-dominant's behavior into helping the queen with her offspring, though the mechanisms of how this is accomplished are debated. Former research suggests that

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Roseler, P.F.; Roseler, I.; Strambi, A.; Augier, R. (1984). "Influence of insect hormones on the establishment of dominance hierarchies among foundresses of the paper wasp, Polistes gallicus".

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Female behavioral strategy: dominance helps females deal with the unusually high reproductive demands; they prevail in more social conflicts because they have more at stake in terms of fitness.

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Koedam, D.; Contrera, A. de O. Fidalgo; Imperatriz-Fonseca, V. L. (2004). "How queen and workers share in male production in the stingless bee Melipona subnitida Ducke (Apidae, Meliponini)".

989:, instead of not laying eggs, the female workers begin being able to reproduce, but once being under the presence of dominant females, the subordinate female workers can no longer reproduce. 1062:. In despotic systems where competition is high, one or two members are dominant while all other members of the living group are equally submissive, as seen in Japanese and rhesus macaques, 185:, the high-ranking males have the first access to vertebrate prey that has been caught by the group, and in yellow baboons the dominant males feed for longer without being interrupted. 824:

Male behavioral strategy: males defer as a parental investment because it ensures more resources in a harsh unpredictable climate for the female, and thus, the male's future offspring.

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in small herds of domestic horses are generally linear hierarchies whereas in large herds the relationships are triangular. Dominance hierarchies can be formed at a very early age.

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subordinates are often displaced from feeding sites by dominant males. Additionally, they are excluded from sleeping sites, and they suffer reduced growth and increased mortality.

572:, eggs from queens have a peculiar chemical profile that workers can distinguish from worker laid eggs. When worker-laid eggs are found, they are eaten. In some species, such as 418:, offspring gain dominance status based on the rank of the mother—the higher ranked the mother, the higher ranked the offspring will be (Yahner). Similarly, the status of a male 181:

success and access to food resources. During times of water shortage the highest-ranking vervet females have greater access than subordinates females to water in tree holes. In

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Drummond, Hugh; Edda Gonzalez; Jose Luis Osorno (1986). "Parent-Offspring Cooperation in the Blue-footed Booby (Sula nebouxii): Social Roles in Infanticidal Brood Reduction".

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Monnin, T.; Ratnieks, F. L. W.; Brandao, C. R. F. (2003). "Reproductive conflict in animal societies: hierarchy length increases with colony size in queenless ponerine ants".

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Drea, Christine M. (April 2007). "Sex and seasonal differences in aggression and steroid secretion in Lemur catta: Are socially dominant females hormonally 'masculinized'?".

1108:, dominance relationships may vary with time as individuals age, gain or lose social status, or change their reproductive condition. Dominance may also vary across space in 960:

was correlated with the individual's ranking position within a dominance hierarchy, but aggression between potential reproductives only started after the queen was removed.

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ants inhibits ovary development; the only reproductive individual of this naturally queenless genus is the one that retains its sclerite intact. This individual is called a

450:. Hence, hierarchy serves as an intrinsic factor for population control, ensuring adequate resources for the dominant individuals and thus preventing widespread starvation. 2885:

Tibbetts, E. A.; Izzo, A. S. (2009). "Endocrine mediated phenotypic plasticity: Condition-dependent effects on juvenile hormone on dominance and fertility of wasp queens".

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seem to support this, as dominant individuals had lower cortisol levels in a stable hierarchy than did subdominant individuals, but the reverse was true at unstable times.

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is determined by the rank of his family. Although dominance is determined differently in each case, it is influenced by the relationships between members of social groups.

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agriculture). This contradicts the "egalitarian hypothesis", which predicts that status would affect reproductive success more amongst foragers than amongst nonforagers.

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Chase, I. D.; Tovey, C.; Murch, P. (2003). "Two's Company, Three's a Crowd: Differences in Dominance Relationships in Isolated versus Socially Embedded Pairs of Fish".

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Widdig, Anja; Kessler, Matthew J.; Bercovitch, Fred B.; et al. (2016). "Genetic studies on the Cayo Santiago rhesus macaques: a review of 40 years of research".

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Foster, R. L.; Ameilia, B.; Verdirame, D.; O'Donnell, S. (2004). "Reproductive physiology, dominance interactions, and division of labour among bumble bee workers".

849:. Although a high rank is an advantage for females, clear linear hierarchies in female chimpanzees have not been detected. In "masculinized" female mammals like the 818:

The Energy Conservation Hypothesis: males subordinate to females to conserve energy for intense male-male competition experienced during very short breeding seasons

392:, the resident of the territory is likely to end as the victor because he values the territory more. This can be explained further by looking at the example of the 3174:

Wang, F.; Zhu, J.; Zhu, H.; Zhang, Q.; Lin, Z.; Hu, H. (2011-11-04). "Bidirectional Control of Social Hierarchy by Synaptic Efficacy in Medial Prefrontal Cortex".

604:, responsible for regulating growth and development in insects including wasps, the foundresses exhibit an increase in dominance. Further, foundresses with larger 608:, a region of the female wasp brain responsible for the synthesis and secretion of juvenile hormone, are naturally more dominant. A follow-up experiment utilized 371:

Animal decisions regarding involvement in conflict are defined by the interplay between the costs and benefits of agonistic behaviors. When initially developed,

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individuals, the opposite of what had been expected. Two core hypotheses attempt to explain this. The first suggests that higher ranking individuals exert more

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sheep revealed an almost linear hierarchy in the Merinos but a less rigid structure in the Border Leicesters when a competitive feeding situation was created.

4542: 600:). This depends on the queen (or foundress), possibly involving specific hormones. Laboratory experiments have shown that when foundresses are injected with 1058:. In systems where competition between and within the sexes is low, social behaviour gravitates towards tolerance and egalitarianism, such as that found in 3648:

Wittig, Roman M.; Boesch, Christophe (1 August 2003). "Food Competition and Linear Dominance Hierarchy Among Female Chimpanzees of the Taï National Park".

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Hewitt, Stacey E.; Macdonald, David W.; Dugdale, Hannah L. (2009). "Context-dependent linear dominance hierarchies in social groups of European badgers,

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Leigh, Steven R.; Setchell, Joanna M.; Charpentier, Marie; et al. (2008). "Canine tooth size and fitness in male mandrills (Mandrillus sphinx)".

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which live in social packs separated into male and female hierarchies, top-ranking alpha females have been observed to produce 76–81% of all litters.

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and introduced into English in 1927. In his 1924 German-language article, he noted that "defense and aggression in the hen is accomplished with the

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Hoover, S. E. R.; Keeling, C. I.; Winston, M. L.; Slessor, K. N. (2003). "The effect of queen pheromones on worker honey bee ovary development".

2930:"Hormonal and behavioural correlates of male dominance and reproductive status in captive colonies of the naked mole–rat, Heterocephalus glaber" 4771: 3402:

Sauther, Michelle L. (1993). "Resource Competition in Wild Populations of Ringtailed Lemurs (Lemur Catta): Implications for Female Dominance".

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Baker, P. J.; Robertson, C. P. J.; Funk, S. M.; Harris, S. (1998). "Potential fitness benefits of group living in the red fox, Vulpes vulpes".

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teats produce a greater quantity of milk. Once established, this teat order remains stable with each piglet tending to feed from a particular

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and, within minutes of being born, or sometimes seconds, will attempt to suckle. The piglets are born with sharp teeth and fight to develop a

901:". This emphasis on pecking led many subsequent studies on fowl behaviour to use it as a primary observation; however, it has been noted that 4206: 3419: 2415: 687:, may be implicated in dominance hierarchies. Higher ranking individuals tend to have much higher levels of circulating glucocorticoids than 2619:

Peeters, C.; Liebig, J.; Hölldobler, B. (2000). "Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator".

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Owens, D.; Owens, M. (1996). "Social dominance and reproductive patterns in brown hyaenas, Hyaena brunnea, of the central Kalahari desert".

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Several areas of the brain contribute to hierarchical behavior in animals. One of the areas that has been linked with this behavior is the

1608:"Men's status and reproductive success in 33 nonindustrial societies: Effects of subsistence, marriage system, and reproductive strategy" 4824: 772: 385:

Resource value: Animals more invested in a resource are likely to invest more in the fight despite potential for incurring higher costs.

732:, the primary excitatory cell type of the brain. High ranking macaques have a larger rostral prefrontal cortex in large social groups. 337:, which display defined linear dominance in both sexes, allow subordinate males and females decreased time of feeding at a carcass. In 4222:

Hooper, Paul L.; Kaplan, Hillard S.; Jaeggi, Adrian V. (1 March 2021). "Gains to cooperation drive the evolution of egalitarianism".

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Young, Andrew L.; Richard, Alison F.; Aiello, Leslie C. (1990). "Female Dominance and Maternal Investment in Strepsirhine Primates".

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In rodents, the highest-ranking male frequently sires the most offspring. The same pattern is found in most carnivores, such as the

3849: 4575: 123:, feeding, aggressive, and other behaviour patterns subsequently occur with relatively little influence from other group members. 303:

are correlated with their age. Older, subordinate males form alliances to combat higher-ranking males and get access to females.

4958: 1665:

Samuels, A.; Silk, J. B.; Rodman, P. (1984). "Changes in the dominance rank and reproductive behavior of male bonnet macaques (

4591:"The fourth level of social structure in a multi-level society: Ecological and social functions of clans in hamadryas baboons" 1401:

Stewart, J. C.; Scott, J. P. (1947). "Lack of correlation between leadership and dominance relationships in a herd of goats".

3233:

Sallet, J.; Mars, R. B.; Noonan, M. P.; et al. (2011-11-04). "Social Network Size Affects Neural Circuits in Macaques".

2378:

Functional Importance of Plumage Badges as Intraspecific Signals in White-Crowned Sparrows (zonotrichia Leucophrys Oriantha)

3685:"Androgens and masculinization of genitalia in the spotted hyaena (Crocuta crocuta). 2. Effects of prenatal anti-androgens" 4746:

on 'The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies' (2022)

1133:

or group of teats. Dominance–subordination relationships can vary markedly between breeds of the same species. Studies on

1118: 3816:

Schjelderup-Ebbe, T. (1975). "Contributions to the social psychology of the domestic chicken ". In Schein, M. W. (ed.).

1947:

Tiedens, Larissa; Fragale, Alison (2003). "Power moves: Complementarity in dominant and submissive nonverbal behavior".

829: 971:

development is not totally achieved in the worker caste, which opens the possibility of reproduction by workers. Since

5049: 3131:

Sapolsky, R. M. (1992). "Cortisol concentrations and the social significance of rank instability among wild baboons".

993: 465:

The suppression of reproduction by dominant individuals is the most common mechanism that maintains the hierarchy. In

46: 2205:

Dittus, W. P. J. (1977). "The Social Regulation of Population Density and Age-Sex Distribution in the Toque Monkey".

382:

Resource-holding potential: Animals that are better able to defend resources often win without much physical contact.

3065: 2982:"Evidence that primer pheromones do not cause social suppression of reproduction in male and female naked mole-rats" 4966: 4859: 4764: 4458: 4022: 1439:

Squires, V. R.; Daws, G. T. (1975). "Leadership and dominance relationships in Merino and Border Leicester sheep".

1024: 1032:, queens typically control the nest, though a "dwarf" queen will take its place in the case of a premature death. 886: 559:" is an additional mechanism that prevents reproduction by workers, found in bees and ants. Policing may involve 574: 533:, and is responsible for mutilating all the newly emerged females, to maintain its social status. Gamergates of 4917: 4881: 2479:

Hughes, C. R.; Beck, M. O.; Strassman, J. E. (1987). "Queen succession in the social wasp Polistes annularis".

1238:

Cheng, Joey T. (2020). "Dominance, prestige, and the role of leveling in human social hierarchy and equality".

684: 255: 4100:"Interaction and Behavior of Virgin and Physogastric Queens in Three Meliponini Species (Hymenoptera, Apidae)" 5037: 4472:

Wroblewski, E. E.; Murray, C. M.; Keele, B. F.; Schumacher-Stankey, J. C.; Hahn, B. H.; Pusey, A. E. (2009).

1018:

In some species, especially in ants, more than one queen can be found in the same colony, a condition called

956:. In a laboratory experiment, Clarke and Faulkes (1997) demonstrated that reproductive status in a colony of 551:, a pheromone produced by the queen mandibular glands is responsible for inhibiting ovary development in the 519:

Other behaviors are involved in maintaining reproductive status in social insects. The removal of a thoracic

4844: 4814: 4804: 3021: 2809:"Agonistic interactions and reproductive dominance in Pachycondyla obscuricornis (Hymenoptera, Formicidae)" 1104:

Dominance and its organisation can be highly variable depending on the context or individuals involved. In

967:. Queen and workers are diploid, but males develop from haploid genotypes. In some species, suppression of 832:

at this time and all three are likely to play a role. Adult female lemurs have increased concentrations of

4295:

Matsumura, Shuichi (1999). "The evolution of "egalitarian" and "despotic" social systems among macaques".

3942:"Dominance and queen succession in captive colonies of the eusocial naked mole-rat, Heterocephalus glaber" 1956: 1138: 1076: 754:

connection has been linked with maintenance of rank in mice. Another area that has been associated is the

430:

Individuals with greater hierarchical status tend to displace those ranked lower from access to space, to

120: 42: 3881: 5025: 4819: 4757: 2046:

Hogg, J. T.; Forbes, S. H. (1997). "Mating in bighorn sheep: Frequent male reproduction via a high-risk

1050: 1046: 656:

are suppressed, while in females it appears that the suppression involves the entire suppression of the

644: 568: 535: 496: 410: 213:

females; when their rank decreases, they get less time. In many primates, including bonnet macaques and

143: 4059:"Observations on the Winter Aggregates of Two Polistine Paper Wasps (Hymenoptera Vespidae Polistinae)" 2087:"Dominance status and carcass availability affect the outcome of sperm competition in burying beetles" 1992:

Pusey, A. E.; Packer, C. (1997). "The ecology of relationships". In J. R. Krebs; N. B. Davies (eds.).

188:

In many bird species, the dominant individuals have higher rates of food intake. Such species include

4799: 4794: 4653: 4070: 3953: 3737: 3323: 3242: 3183: 2761: 2714: 2663: 2577: 2488: 1903: 1808: 1619: 846: 755: 490: 443: 306:

Fighting with dominant males is a risky behavior that may result in defeat, injury or even death. In

578:, workers may try to escape policing by shuffling their eggs within the egg pile laid by the queen. 4849: 1961: 1291: 1019: 1011: 999: 649: 38: 5044: 2423: 1571:

Cowlishaw, Guy; Dunbar, Robin I. M. (1991). "Dominance rank and mating success in male primates".

54: 5076: 4937: 4866: 4722: 4618: 4408: 4369: 4320: 4247: 4168: 4139:

Klass, Keren; Cords, Marina (17 September 2015). "Agonism and dominance in female blue monkeys".

4039: 3922: 3853: 3798: 3726:"Exceptional endocrine profiles characterise the meerkat: sex, status, and reproductive patterns" 3665: 3630: 3583: 3495: 3452: 3384: 3266: 3215: 3156: 3098: 2910: 2864: 2730: 2687: 2636: 2601: 2531: 2461: 2327: 2292: 2249: 2187: 2067: 2028: 1867: 1832: 1729: 1686: 1588: 1263: 1109: 1005: 609: 596: 511: 484: 451: 389: 237: 4896: 930: 865:), they have "exceptionally high concentrations" of androgens, "particularly during gestation". 388:

Intruder retreats: When participants are of equal fighting ability and competing for a certain

139: 5071: 4982: 4714: 4679: 4610: 4503: 4361: 4312: 4239: 4202: 4190: 4160: 4148: 4121: 3979: 3763: 3706: 3622: 3566:

Drea, Christine M. (August 2009). "Endocrine Mediators of Masculinization in Female Mammals".

3548: 3487: 3415: 3376: 3341: 3292: 3258: 3207: 3199: 3148: 3003: 2959: 2902: 2789: 2679: 2593: 2284: 2149: 1997: 1974: 1929: 1824: 1780: 1747:

Dubuc, Constance; Muniz, Laura; Heistermann, Michael; Engelhardt, Antje; Widdig, Anja (2011).

1721: 1647: 1519: 1418: 1255: 1201: 985: 980: 913: 838: 808: 782: 725: 4474:"Male dominance rank and reproductive success in chimpanzees, Pan troglodytes schweinfurthii" 5003: 4992: 4854: 4743: 4706: 4669: 4661: 4602: 4493: 4485: 4435: 4400: 4351: 4304: 4231: 4152: 4111: 4078: 4031: 3969: 3961: 3914: 3845: 3790: 3753: 3745: 3696: 3657: 3614: 3575: 3538: 3530: 3479: 3444: 3407: 3368: 3331: 3250: 3191: 3140: 3088: 3080: 3039: 2993: 2949: 2941: 2894: 2856: 2820: 2779: 2769: 2722: 2671: 2628: 2585: 2523: 2496: 2453: 2382: 2361: 2319: 2276: 2241: 2214: 2179: 2139: 2131: 2098: 2059: 2020: 1966: 1919: 1911: 1859: 1816: 1770: 1760: 1713: 1678: 1637: 1627: 1580: 1557: 1511: 1479: 1448: 1410: 1362: 1331: 1300: 1247: 1193: 1105: 926: 601: 466: 378:

The asymmetries between individuals have been categorized into three types of interactions:

631: 4927: 4834: 4829: 2245: 1387: 976: 972: 704: 556: 530: 475: 189: 4426:

Houpt, K. A.; Law, K.; Martinisi, V. (1978). "Dominance hierarchies in domestic horses".

2568:

Tibbetts, E. A.; Dale, J. (2004). "A socially enforced signal of quality in paper wasp".

4657: 4074: 3957: 3741: 3327: 3246: 3187: 2765: 2718: 2667: 2581: 2492: 1907: 1812: 1623: 736:

studies with computer stimulated hierarchal conditions showed increased activity in the

4932: 4674: 4641: 4498: 4473: 3974: 3941: 3758: 3725: 3543: 3514: 2954: 2929: 2500: 2144: 2119: 1924: 1891: 1775: 1748: 1642: 1607: 1063: 975:

are seasonal and workers are wingless, workers are almost always non-breeders, and (as

950: 885:

The concept of dominance, originally called "pecking order", was described in birds by

680: 673: 605: 547: 505: 313: 221: 209: 4697:

Furuichi, T. (2011). "Female contributions to the peaceful nature of bonobo society".

4035: 3836:

Rajecki, D. W. (1988). "Formation of leap orders in pairs of male domestic chickens".

3820:. Vol. 3. Stroudsburg, Pennsylvania: Dowden, Hutchinson and Ross. pp. 35–49. 3359:

Digby, L. I.; Kahlenberg, S. M. (2002). "Female dominance in blue-eyed black lemurs".

2784: 2749: 1682: 1584: 1515: 5065: 4947: 4942: 4839: 4622: 4522: 4439: 4251: 4186: 3857: 3634: 3587: 3579: 3270: 3144: 3102: 2457: 1452: 1286: 1267: 1085: 1081: 1055: 850: 657: 564: 439: 415: 346: 307: 214: 193: 182: 161: 87: 4726: 4590: 4412: 4324: 4099: 4043: 3926: 3724:

Davies, Charli S.; Smyth, Kendra N.; Greene, Lydia K.; et al. (December 2016).

3669: 3499: 3456: 3388: 3219: 3160: 2868: 2750:"Surface Hydrocarbons of queen eggs regulate worker reproduction in a social insect" 2691: 2549:

Monnin, T. (2006). "Chemical recognition of reproductive status in social insects".

2535: 2465: 2331: 2296: 2253: 2191: 2032: 1871: 1836: 1733: 1690: 1592: 594:

may be associated with dominance hierarchies in the social order of the paper wasp (

539:

arise from aggressive interactions, forming a hierarchy of potential reproductives.

320:

and attempt to copulate before the dominant male can drive them forcefully away. In

4952: 4922: 4888: 4780: 4626: 4527: 4373: 4172: 4083: 4058: 3116:

Genuth, S. M. (1993). "The endocrine system". In Berne, R. M.; Levy, M. N. (eds.).

2914: 2734: 2640: 2605: 2120:"Flat lizard female mimics use sexual deception in visual but not chemical signals" 2071: 1159: 1122: 1114: 1029: 964: 741: 733: 709: 653: 552: 419: 393: 338: 299: 31: 4489: 4404: 699:

for energy use. This is supported by the fact that when food availability is low,

3874:

Aggressive behavior of chickens: Some effects of social and physical environments

3411: 2011:

Noe, R.; Sluijter, A. A. (1990). "Reproductive Tactics of Male Savanna Baboons".

1251: 1197: 4997: 4876: 3618: 3483: 2898: 1890:

Gesquiere, Laurence R.; Learn, Niki H.; Simao, Carolina M.; et al. (2011).

688: 591: 372: 334: 321: 4235: 3312:"History of winning remodels thalamo-PFC circuit to reinforce social dominance" 2705:

Ratnieks, F. L. W.; Visscher, P. K. (1989). "Worker policing in the honeybee".

2323: 1970: 1749:"Testing the priority-of-access model in a seasonally breeding primate species" 131:, and can be relatively easily influenced or inhibited by other group members. 4987: 4901: 4356: 4339: 3661: 3084: 2675: 2527: 1863: 1820: 1765: 1335: 1009:

also exhibits this type of hierarchy. Within the dominance hierarchies of the

873: 842: 480: 232: 67: 3296: 3203: 2232:

Creel, S. (1997). "Handling of African wild dogs and chronic stress: Reply".

1028:. In this species, multiple queens of varying sizes are present. The larger, 4871: 4198: 3336: 3311: 3254: 3195: 2774: 2218: 2103: 2086: 2085:

Pettinger, Adam M.; Steiger, Sandra; Mueller, Josef K.; et al. (2011).

2024: 1915: 1632: 1367: 1350: 1304: 1164: 759: 703:

levels increase within the dominant male. The second suggests that elevated

668: 563:

and immobilization of workers who lay eggs. In some ant species such as the

543: 471: 447: 446:

by mating more often and having more resources to invest in the survival of

317: 251: 91: 83: 17: 4718: 4683: 4614: 4558:"Sperm Competition in the Florida Manatee (Trichechus manatus latirostris)" 4507: 4365: 4316: 4243: 4164: 4125: 3965: 3767: 3626: 3552: 3534: 3491: 3380: 3345: 3262: 3211: 2945: 2906: 2793: 2683: 2597: 2280: 2183: 2153: 2135: 1978: 1933: 1828: 1784: 1725: 1651: 1422: 1259: 1205: 4193:. In Carter, Carol Sue; Lederhendler, I. Izja; Kirkpatrick, Brian (eds.). 4116: 3983: 3710: 3701: 3684: 3513:

von Engelhard, Nikolaus; Kappeler, Peter M.; Heistermann, Michael (2000).

3152: 3007: 2998: 2981: 2963: 2288: 2063: 1523: 1096: 912:

Dominance hierarchies are found in many species of bird. For example, the

665: 4809: 3523:

Proceedings of the Royal Society of London. Series B: Biological Sciences

3093: 2934:

Proceedings of the Royal Society of London. Series B: Biological Sciences

2825: 2808: 2386: 1273:

hierarchical relationships based on dominance are relatively less stable.

1126: 1067: 938: 833: 751: 746: 700: 696: 661: 613: 525: 520: 454: 402: 178: 71: 59: 4665: 3880:. 27th Annual National Breeder's Roundtable. Kansas City. Archived from 3850:

10.1002/1098-2337(1988)14:6<425::AID-AB2480140604>3.0.CO;2-#

2589: 1319: 4710: 4308: 3918: 3372: 3310:

Zhou, Tingting; Zhu, Hong; Fan, Zhengxiao; et al. (14 July 2017).

2860: 2632: 1071: 1059: 902: 878: 858: 814:

There are three basic proposals for the evolution of female dominance:

737: 625: 617: 587: 560: 359: 4606: 4543:"Humans Would be Better off if They Monkeyed Around Like the Muriquis" 4156: 3997:

West-Eberhard, M. J. (1969). "The social biology of polistine wasps".

3802: 3749: 3683:

Drea, C. M.; Weldele, M. L.; Forger, N. G.; et al. (1 May 1998).

3044: 3028:"Dominance, aggression and glucocorticoid levels in social carnivores" 3027: 2376: 1717: 1484: 1467: 2726: 1414: 1386:(4th, revised and edited by Jean Laing ed.). London and Boston: 1134: 804: 787: 729: 692: 636: 435: 295: 103: 3818:

Social Hierarchy and Dominance. Benchmark Papers in Animal Behavior

3794: 3448: 4523:"Meat-sharing as a coalition strategy by an alpha male chimpanzee" 4191:"Conflict Resolution and Distress Alleviation in Monkeys and Apes" 1095: 1003:, where the next queen is selected based on age rather than size. 968: 872: 800: 796: 781: 713: 630: 501: 138: 53: 30:"Alpha female" redirects here. For the professional wrestler, see 4640:

Crisp, Rachel J.; Brent, Lauren J. N.; Carter, Gerald G. (2021).

4098:

Nogueira-Ferreira, F. H.; Silva-Matos, E. V.; Zucchi, R. (2009).

4016:

Strassmann & Meyer (1983). "Gerontocracy in the social wasp,

1466:

Portugal, S. J.; Sivess, L.; Martin, G. R.; et al. (2017).

1130: 946: 934: 898: 811:

is observed to be the most prominent model of female dominance.

431: 4753: 4749: 2118:

Whiting, Martin J.; Webb, Jonathan K.; Keogh, J. Scott (2009).

942: 37:"Alpha male" redirects here. For the slang terms for men, see 3120:. Vol. 3. St. Louis: Mosby Year Book. pp. 813–1024. 1287:"The concept and definition of dominance in animal behaviour" 3289:

The mouse that roared: neural mechanisms of social hierarchy

963:

The social insects mentioned above, excluding termites, are

695:

and thus need higher levels of glucocorticoids to mobilize

4057:

González, J. A.; Nascimento, F. S.; Gayubo, S. F. (2002).

1434: 1432: 4642:"Social dominance and cooperation in female vampire bats" 2748:

Endler, A.; Liebig, J.; Schmitt, T.; et al. (2004).

1502:

Rowell, T. E. (1974). "The concept of social dominance".

345:

far less with females than do the high-ranking males. In

98:, and a submissive lower-ranking individual is called a 635:

The effect of relative rank on stress hormone levels in

2124:

Proceedings of the Royal Society B: Biological Sciences

1892:"Life at the Top: Rank and Stress in Wild Male Baboons" 1320:"The nature and measurement of interpersonal dominance" 790:

is one of the few mammals with female-biased dominance.

2880: 2878: 1606:

Von Rueden, Christopher R.; Jaeggi, Adrian V. (2016).

828:

In lemurs, no single hypothesis fully explains female

2842: 2840: 2838: 2836: 3066:"Dominance, cortisol and stress in wild chimpanzees" 2975: 2973: 62:

advertises his status with bright facial coloration.

4975: 4910: 4787: 2381:(Thesis). Deep Blue at the University of Michigan. 2351: 2349: 2347: 2345: 2343: 2341: 177:A benefit to high-ranking individuals is increased 4338:Watts, Heather E.; Holekamp, Kay E. (2007-08-21). 3781:Perrin, P. G. (1955). "'Pecking order' 1927–54". 2356:Huntingford, Felicity; Turner, Angela K. (1987). 1552:Huntingford, Felicity; Turner, Angela K. (1987). 881:showing feather damage from pecking by other hens 94:higher-ranking individual is sometimes called an 3515:"Androgen levels and female social dominance in 3404:Lemur Social Systems and Their Ecological Basis 2928:Clarke, F. M.; Faulkes, C. G. (7 August 1998). 1612:Proceedings of the National Academy of Sciences 1468:"Perch height predicts dominance rank in birds" 1403:Journal of Comparative Physiological Psychology 1145:Species with egalitarian/non-linear hierarchies 905:tend to leap and use their claws in conflicts. 3073:Journal of Behavioral Ecology and Sociobiology 2364:. pp. 156–159, 194–205, 208–209, 250–253. 1547: 1545: 1543: 1541: 1539: 1537: 1535: 1533: 1497: 1495: 4765: 3599: 3597: 3059: 3057: 3055: 1994:Behavioural Ecology: An Evolutionary Approach 438:opportunities. Thus, individuals with higher 8: 1949:Journal of Personality and Social Psychology 1885: 1883: 1881: 1045:Dominance hierarchies emerge as a result of 4744:Theme issue of Philosophical Transactions B 4589:Schreier, Amy L.; Swedell, Larissa (2009). 4195:The Integrative Neurobiology of Affiliation 3568:Current Directions in Psychological Science 1318:Burgoon, J.; Johnson, M.; Koch, P. (1998). 107:animal is challenged by a subordinate one. 41:. For dominance hierarchies in humans, see 4772: 4758: 4750: 683:, signaling molecules which stimulate the 4673: 4497: 4455:A Natural History of Domesticated Mammals 4355: 4282:Animal Behavior: An Evolutionary Approach 4115: 4082: 3973: 3757: 3700: 3542: 3335: 3092: 3043: 2997: 2953: 2824: 2783: 2773: 2143: 2102: 1960: 1923: 1774: 1764: 1641: 1631: 1483: 1366: 1351:"Helping effort in a dominance hierarchy" 672:cause her to exert intense dominance and 127:animals are opposite; their behaviour is 4521:T. Nishida; et al. (January 1992). 2165: 2163: 1349:Cant, Michael A.; Field, Jeremy (2005). 616:known to enhance maturation and size of 272:Interpersonal complementarity hypothesis 266:Interpersonal complementarity hypothesis 4284:. Sinauer Associates. pp. 476–511. 1176: 90:interact, creating a ranking system. A 3940:Clarke, F. M.; Faulkes, C. G. (1997). 3064:Muller, M. N.; Wrangham, R.W. (2004). 2980:Faulkes, C. G.; Abbott, D. H. (1993). 2940:(1404). The Royal Society: 1391–1399. 2404:. New York: Springer. pp. 95–100. 1100:Eringer cattle competing for dominance 3282: 3280: 2986:Journal of Reproduction and Fertility 2375:Laubach, Zachary (27 November 2012). 2246:10.1046/j.1523-1739.1997.0110061454.x 7: 5032: 4556:Champagne, Melissa (November 2008). 3650:International Journal of Primatology 78:(formerly and colloquially called a 27:Type of socially subordinate ranking 4825:Evolutionary models of food sharing 3907:Behavioral Ecology and Sociobiology 3133:Journal of Psychoneuroendocrinology 2849:Behavioral Ecology and Sociobiology 2807:Oliveim, PS; Hölldobler, B (1991). 2516:Behavioral Ecology and Sociobiology 2052:Behavioral Ecology and Sociobiology 1753:Behavioral Ecology and Sociobiology 773:List of dominance hierarchy species 86:that arises when members of animal 2501:10.1111/j.1439-0310.1987.tb00678.x 2402:Wildlife Behavior and Conservation 660:. This suppression reduces sexual 25: 3287:Wang F, Kessels HW, Hu H (2014). 2422:. 22 October 1973. Archived from 146:have a clear dominance hierarchy. 5043: 5031: 5020: 5019: 4576:"Elephants are socially complex" 3580:10.1111/j.1467-8721.2009.01640.x 2458:10.1111/j.0307-6962.2004.00388.x 228:all of the mating in the group. 4959:Sociobiology: The New Synthesis 4595:American Journal of Primatology 4269:. Pearson Education. p. 5. 4141:American Journal of Primatology 4104:Genetics and Molecular Research 3999:Mis. Publ. Zool. Univ. Michigan 2813:Psyche: A Journal of Entomology 2416:"Behavior: The Animal Watchers" 1706:American Journal of Primatology 889:in 1921 under the German terms 155:When individuals seek high rank 4084:10.1080/03946975.2002.10531162 3406:. Springer. pp. 135–152. 1: 4490:10.1016/j.anbehav.2008.12.014 4405:10.1016/j.anbehav.2008.09.022 4036:10.1016/S0003-3472(83)80063-3 1683:10.1016/s0003-3472(84)80212-2 1585:10.1016/s0003-3472(05)80642-6 1516:10.1016/S0091-6773(74)90289-2 1240:Current Opinion in Psychology 992:In some wasp species such as 4440:10.1016/0304-3762(78)90117-7 3871:Craig, J. V. (11 May 1978). 3412:10.1007/978-1-4899-2412-4_10 3145:10.1016/0306-4530(92)90029-7 3020:For example, fishes such as 1453:10.1016/0304-3762(75)90019-X 1252:10.1016/j.copsyc.2019.10.004 1198:10.1016/j.jhevol.2008.01.001 720:Brain pathways and hierarchy 5050:Evolutionary biology portal 4267:Primate Behavioural Ecology 3824:Zeitschrift für Psychologie 3619:10.1016/j.yhbeh.2019.07.002 3484:10.1016/j.yhbeh.2007.02.006 2899:10.1016/j.yhbeh.2009.09.003 2400:Yahner, Richard H. (2012). 994:Liostenogaster flavolineata 778:Female dominance in mammals 47:Alpha male (disambiguation) 5093: 4646:Royal Society Open Science 4459:Cambridge University Press 4453:Clutton-Brock, J. (1987). 4236:10.1038/s41562-021-01059-y 4189:; Aureli, Filippo (1999). 2324:10.1163/156853903771980558 1971:10.1037/0022-3514.84.3.558 1186:Journal of Human Evolution 1041:Spectrum of social systems 1025:Schwarziana quadripunctata 770: 575:Pachycondyla obscuricornis 269: 36: 29: 5015: 4699:Evolutionary Anthropology 4357:10.1016/j.cub.2007.06.002 3085:10.1007/s00265-003-0713-1 2676:10.1007/s00114-003-0462-z 2551:Annales Zoologici Fennici 2528:10.1007/s00265-003-0600-9 1864:10.1007/s00040-004-0781-x 1821:10.1007/s00114-013-1123-5 1766:10.1007/s00265-011-1172-8 1336:10.1080/03637759809376456 887:Thorleif Schjelderup-Ebbe 2446:Physiological Entomology 1324:Communication Monographs 685:fight or flight response 4845:Male warrior hypothesis 4815:Evolutionary psychology 4805:Dual inheritance theory 4562:Hcnso Student Capstones 4428:Applied Animal Ethology 3662:10.1023/A:1024632923180 3437:The American Naturalist 3337:10.1126/science.aak9726 3255:10.1126/science.1210027 3196:10.1126/science.1209951 3022:Oreochromis mossambicus 2775:10.1073/pnas.0308447101 2219:10.1163/156853977x00450 2025:10.1163/156853990x00455 1916:10.1126/science.1207120 1633:10.1073/pnas.1606800113 1441:Applied Animal Ethology 1305:10.1163/156853993X00290 367:Conflict over dominance 245:Costs of being dominant 4265:Strier, Karen (2006). 4224:Nature Human Behaviour 3966:10.1098/rspb.1997.0137 3535:10.1098/rspb.2000.1175 2946:10.1098/rspb.1998.0447 2281:10.1006/anbe.1998.0950 2184:10.1006/anbe.1996.0058 2136:10.1098/rspb.2008.1822 1382:MacKenzie, D. (1980). 1285:Drews, Carlos (1993). 1219:Yeh J (11 June 2018). 1101: 1077:Neolamprologus pulcher 1060:woolley spider monkeys 882: 791: 639: 457:enhances this effect. 411:white-crowned sparrows 147: 144:Wedge-capped capuchins 82:) is a type of social 63: 45:. For other uses, see 43:Dual strategies theory 4820:Evolution of morality 4280:Alcock, John (2018). 4117:10.4238/vol8-2kerr008 3702:10.1530/jrf.0.1130117 3607:Hormones and Behavior 3472:Hormones and Behavior 2999:10.1530/jrf.0.0990225 2887:Hormones and Behavior 2104:10.1093/beheco/arr093 2064:10.1007/s002650050361 1368:10.1093/beheco/ari051 1221:"Dominance Hierarchy" 1099: 954:Heterocephalus glaber 876: 785: 645:Heterocephalus glaber 634: 569:Camponotus floridanus 536:Harpegnathos saltator 442:tend to have greater 426:Regulation mechanisms 328:Costs to subordinates 142: 57: 4800:Challenge hypothesis 4795:Behavioural genetics 4201:. pp. 119–140. 4187:de Waal, Frans B. M. 3826:, 1922, 88:225-252.) 3032:Journal of Mammalogy 2234:Conservation Biology 949:), but also for the 847:reproductive success 756:dorsal raphe nucleus 491:Dinoponera australis 444:reproductive success 353:Mitigating the costs 298:have high levels of 200:Reproductive success 58:A high-ranking male 4850:Reciprocal altruism 4666:10.1098/rsos.210266 4658:2021RSOS....810266C 4075:2002TrZoo..15....1G 3958:1997RSPSB.264..993C 3887:on 16 February 2016 3838:Aggressive Behavior 3742:2016NatSR...635492D 3529:(1452): 1533–1539. 3328:2017Sci...357..162Z 3247:2011Sci...334..697S 3188:2011Sci...334..693W 2766:2004PNAS..101.2945E 2719:1989Natur.342..796R 2668:2003NW.....90..477H 2656:Naturwissenschaften 2590:10.1038/nature02949 2582:2004Natur.432..218T 2493:1987Ethol..76..124H 2426:on 14 December 2008 2130:(1662): 1585–1591. 2000:. pp. 254–283. 1908:2011Sci...333..357G 1813:2014NW....101...17B 1801:Naturwissenschaften 1624:2016PNAS..11310824V 1618:(39): 10824–10829. 1012:Polistes versicolor 1000:Polistes instabilis 845:is correlated with 650:luteinizing hormone 461:In eusocial animals 76:dominance hierarchy 39:Alpha and beta male 4938:Sarah Blaffer Hrdy 4867:Sex and psychology 4711:10.1002/evan.20308 4309:10.1007/BF02557699 4018:Polistes exclamans 3952:(1384): 993–1000. 3919:10.1007/bf00295710 3730:Scientific Reports 3373:10.1007/BF02629647 3026:Creel, S. (2005). 2861:10.1007/bf00299381 2826:10.1155/1991/64635 2633:10.1007/pl00001724 2091:Behavioral Ecology 1504:Behavioral Biology 1355:Behavioral Ecology 1102: 1092:Context dependency 1088:and brown hyenas. 1006:Polistes exclamans 981:laying worker bees 883: 863:Suricata suricatta 839:ring-tailed lemurs 792: 767:In specific groups 640: 610:20-hydroxyecdysone 597:Polistes dominulus 512:Polistes dominulus 485:Polistes annularis 238:Melipona subnitida 148: 64: 5059: 5058: 4983:Stephen Jay Gould 4607:10.1002/ajp.20736 4461:. pp. 73–74. 4350:(16): R657–R660. 4340:"Hyena societies" 4208:978-0-26253-158-0 4157:10.1002/ajp.22481 3750:10.1038/srep35492 3421:978-1-4899-2414-8 3322:(6347): 162–168. 3241:(6056): 697–700. 3182:(6056): 693–697. 3045:10.1644/bhe-002.1 2713:(6251): 796–797. 2576:(7014): 218–222. 1998:Blackwell Science 1718:10.1002/ajp.22424 1560:. pp. 39–54. 1485:10.1111/ibi.12447 1390:. pp. 66–85. 1139:Border Leicesters 986:Polistes fuscatus 914:blue-footed booby 809:ring-tailed lemur 726:prefrontal cortex 642:Naked mole-rats ( 347:African wild dogs 16:(Redirected from 5084: 5047: 5035: 5034: 5023: 5022: 5004:Not in Our Genes 4993:Richard Lewontin 4855:Sexual selection 4774: 4767: 4760: 4751: 4731: 4730: 4694: 4688: 4687: 4677: 4637: 4631: 4630: 4586: 4580: 4579: 4572: 4566: 4565: 4553: 4547: 4546: 4539: 4533: 4532: 4518: 4512: 4511: 4501: 4478:Animal Behaviour 4469: 4463: 4462: 4450: 4444: 4443: 4423: 4417: 4416: 4393:Animal Behaviour 4384: 4378: 4377: 4359: 4335: 4329: 4328: 4292: 4286: 4285: 4277: 4271: 4270: 4262: 4256: 4255: 4219: 4213: 4212: 4183: 4177: 4176: 4136: 4130: 4129: 4119: 4095: 4089: 4088: 4086: 4063:Tropical Zoology 4054: 4048: 4047: 4023:Animal Behaviour 4013: 4007: 4006: 3994: 3988: 3987: 3977: 3937: 3931: 3930: 3902: 3896: 3895: 3893: 3892: 3886: 3879: 3868: 3862: 3861: 3833: 3827: 3822:(Reprinted from 3821: 3813: 3807: 3806: 3778: 3772: 3771: 3761: 3721: 3715: 3714: 3704: 3680: 3674: 3673: 3645: 3639: 3638: 3601: 3592: 3591: 3563: 3557: 3556: 3546: 3510: 3504: 3503: 3467: 3461: 3460: 3432: 3426: 3425: 3399: 3393: 3392: 3356: 3350: 3349: 3339: 3307: 3301: 3300: 3284: 3275: 3274: 3230: 3224: 3223: 3171: 3165: 3164: 3128: 3122: 3121: 3113: 3107: 3106: 3096: 3070: 3061: 3050: 3049: 3047: 3018: 3012: 3011: 3001: 2977: 2968: 2967: 2957: 2925: 2919: 2918: 2882: 2873: 2872: 2844: 2831: 2830: 2828: 2819:(2–3): 215–226. 2804: 2798: 2797: 2787: 2777: 2760:(9): 2945–2950. 2745: 2739: 2738: 2727:10.1038/342796a0 2702: 2696: 2695: 2651: 2645: 2644: 2621:Insectes Sociaux 2616: 2610: 2609: 2565: 2559: 2558: 2546: 2540: 2539: 2511: 2505: 2504: 2476: 2470: 2469: 2441: 2435: 2434: 2432: 2431: 2412: 2406: 2405: 2397: 2391: 2390: 2372: 2366: 2365: 2362:Chapman and Hall 2353: 2336: 2335: 2318:(10): 1193–217. 2307: 2301: 2300: 2275:(6): 1411–1424. 2269:Animal Behaviour 2264: 2258: 2257: 2240:(6): 1454–1456. 2229: 2223: 2222: 2202: 2196: 2195: 2172:Animal Behaviour 2167: 2158: 2157: 2147: 2115: 2109: 2108: 2106: 2097:(5): 1079–1087. 2082: 2076: 2075: 2043: 2037: 2036: 2019:(1–2): 117–170. 2008: 2002: 2001: 1989: 1983: 1982: 1964: 1944: 1938: 1937: 1927: 1902:(6040): 357–60. 1887: 1876: 1875: 1852:Insectes Sociaux 1847: 1841: 1840: 1795: 1789: 1788: 1778: 1768: 1759:(8): 1615–1627. 1744: 1738: 1737: 1701: 1695: 1694: 1662: 1656: 1655: 1645: 1635: 1603: 1597: 1596: 1579:(6): 1045–1056. 1573:Animal Behaviour 1568: 1562: 1561: 1558:Chapman and Hall 1549: 1528: 1527: 1499: 1490: 1489: 1487: 1463: 1457: 1456: 1436: 1427: 1426: 1415:10.1037/h0060710 1398: 1392: 1391: 1379: 1373: 1372: 1370: 1346: 1340: 1339: 1315: 1309: 1308: 1299:(3–4): 283–313. 1282: 1276: 1275: 1235: 1229: 1228: 1225:Encyclopedia.com 1216: 1210: 1209: 1181: 1115:Domestic piglets 1106:European badgers 1082:African wild dog 927:insect societies 921:Eusocial insects 830:social dominance 602:juvenile hormone 582:Hormonal control 488:and in the ants 467:eusocial mammals 256:pied flycatchers 190:dark-eyed juncos 173:Foraging success 21: 5092: 5091: 5087: 5086: 5085: 5083: 5082: 5081: 5062: 5061: 5060: 5055: 5011: 4971: 4928:Richard Dawkins 4906: 4897:Dunbar's number 4835:Kin recognition 4830:Group selection 4783: 4778: 4740: 4738:Further reading 4735: 4734: 4696: 4695: 4691: 4639: 4638: 4634: 4601:(11): 948–955. 4588: 4587: 4583: 4574: 4573: 4569: 4555: 4554: 4550: 4541: 4540: 4536: 4520: 4519: 4515: 4471: 4470: 4466: 4452: 4451: 4447: 4425: 4424: 4420: 4386: 4385: 4381: 4344:Current Biology 4337: 4336: 4332: 4294: 4293: 4289: 4279: 4278: 4274: 4264: 4263: 4259: 4221: 4220: 4216: 4209: 4185: 4184: 4180: 4138: 4137: 4133: 4097: 4096: 4092: 4056: 4055: 4051: 4015: 4014: 4010: 3996: 3995: 3991: 3939: 3938: 3934: 3904: 3903: 3899: 3890: 3888: 3884: 3877: 3870: 3869: 3865: 3835: 3834: 3830: 3815: 3814: 3810: 3783:American Speech 3780: 3779: 3775: 3723: 3722: 3718: 3682: 3681: 3677: 3647: 3646: 3642: 3603: 3602: 3595: 3565: 3564: 3560: 3512: 3511: 3507: 3469: 3468: 3464: 3434: 3433: 3429: 3422: 3401: 3400: 3396: 3358: 3357: 3353: 3309: 3308: 3304: 3286: 3285: 3278: 3232: 3231: 3227: 3173: 3172: 3168: 3130: 3129: 3125: 3115: 3114: 3110: 3068: 3063: 3062: 3053: 3025: 3019: 3015: 2979: 2978: 2971: 2927: 2926: 2922: 2884: 2883: 2876: 2846: 2845: 2834: 2806: 2805: 2801: 2747: 2746: 2742: 2704: 2703: 2699: 2662:(10): 477–480. 2653: 2652: 2648: 2618: 2617: 2613: 2567: 2566: 2562: 2548: 2547: 2543: 2513: 2512: 2508: 2478: 2477: 2473: 2443: 2442: 2438: 2429: 2427: 2414: 2413: 2409: 2399: 2398: 2394: 2374: 2373: 2369: 2358:Animal Conflict 2355: 2354: 2339: 2309: 2308: 2304: 2266: 2265: 2261: 2231: 2230: 2226: 2204: 2203: 2199: 2169: 2168: 2161: 2117: 2116: 2112: 2084: 2083: 2079: 2045: 2044: 2040: 2010: 2009: 2005: 1991: 1990: 1986: 1946: 1945: 1941: 1889: 1888: 1879: 1849: 1848: 1844: 1797: 1796: 1792: 1746: 1745: 1741: 1703: 1702: 1698: 1677:(4): 994–1003. 1664: 1663: 1659: 1605: 1604: 1600: 1570: 1569: 1565: 1554:Animal Conflict 1551: 1550: 1531: 1501: 1500: 1493: 1465: 1464: 1460: 1438: 1437: 1430: 1400: 1399: 1395: 1388:Faber and Faber 1381: 1380: 1376: 1348: 1347: 1343: 1317: 1316: 1312: 1284: 1283: 1279: 1237: 1236: 1232: 1218: 1217: 1213: 1183: 1182: 1178: 1173: 1156: 1147: 1094: 1043: 1038: 973:nuptial flights 931:Hamilton's rule 923: 871: 855:Crocuta crocuta 780: 775: 769: 722: 705:stress hormones 681:Glucocorticoids 637:savanna baboons 584: 557:Worker policing 476:Bombus bifarius 463: 428: 369: 355: 330: 314:Burying beetles 296:savanna baboons 288: 283: 274: 268: 247: 202: 175: 170: 157: 137: 113: 50: 35: 28: 23: 22: 15: 12: 11: 5: 5090: 5088: 5080: 5079: 5074: 5064: 5063: 5057: 5056: 5054: 5053: 5041: 5029: 5016: 5013: 5012: 5010: 5009: 5008: 5007: 4995: 4990: 4985: 4979: 4977: 4973: 4972: 4970: 4969: 4964: 4963: 4962: 4950: 4945: 4940: 4935: 4933:Daniel Dennett 4930: 4925: 4920: 4914: 4912: 4908: 4907: 4905: 4904: 4899: 4894: 4893: 4892: 4886: 4885: 4884: 4869: 4864: 4863: 4862: 4852: 4847: 4842: 4837: 4832: 4827: 4822: 4817: 4812: 4807: 4802: 4797: 4791: 4789: 4785: 4784: 4779: 4777: 4776: 4769: 4762: 4754: 4748: 4747: 4739: 4736: 4733: 4732: 4705:(4): 131–142. 4689: 4632: 4581: 4567: 4548: 4534: 4513: 4484:(4): 873–885. 4464: 4445: 4434:(3): 273–283. 4418: 4399:(1): 161–169. 4379: 4330: 4287: 4272: 4257: 4230:(7): 847–856. 4214: 4207: 4178: 4131: 4110:(2): 703–708. 4090: 4049: 4030:(1): 431–438. 4008: 3989: 3932: 3913:(5): 365–372. 3897: 3863: 3844:(6): 425–436. 3828: 3808: 3795:10.2307/453561 3789:(4): 265–268. 3773: 3716: 3695:(1): 117–127. 3675: 3656:(4): 847–867. 3640: 3593: 3574:(4): 221–226. 3558: 3505: 3478:(4): 555–567. 3462: 3449:10.1086/285057 3443:(4): 473–488. 3427: 3420: 3394: 3367:(3): 191–199. 3351: 3302: 3276: 3225: 3166: 3139:(6): 701–709. 3123: 3108: 3079:(4): 332–340. 3051: 3038:(2): 255–264. 3013: 2992:(1): 225–230. 2969: 2920: 2893:(5): 527–531. 2874: 2855:(2): 133–142. 2832: 2799: 2740: 2697: 2646: 2627:(4): 325–332. 2611: 2560: 2541: 2506: 2487:(2): 124–132. 2471: 2452:(4): 327–334. 2436: 2407: 2392: 2367: 2337: 2302: 2259: 2224: 2213:(3): 281–322. 2197: 2178:(3): 535–551. 2159: 2110: 2077: 2048:unconventional 2038: 2003: 1984: 1962:10.1.1.319.709 1955:(3): 558–568. 1939: 1877: 1858:(2): 114–121. 1842: 1790: 1739: 1696: 1667:Macaca radiate 1657: 1598: 1563: 1529: 1510:(2): 131–154. 1491: 1478:(2): 456–462. 1458: 1447:(3): 263–274. 1428: 1409:(4): 255–264. 1393: 1384:Goat Husbandry 1374: 1361:(4): 708–715. 1341: 1330:(4): 308–335. 1310: 1277: 1230: 1211: 1175: 1174: 1172: 1169: 1168: 1167: 1162: 1155: 1152: 1146: 1143: 1093: 1090: 1086:spotted hyenas 1074:, the cichlid 1068:dwarf hamsters 1064:leopard geckos 1042: 1039: 1037: 1034: 951:naked mole-rat 922: 919: 870: 867: 837:dominance" in 826: 825: 822: 819: 779: 776: 771:Main article: 768: 765: 758:, the primary 721: 718: 674:aggressiveness 606:corpora allata 586:Modulation of 583: 580: 548:Apis mellifera 462: 459: 427: 424: 416:rhesus monkeys 398: 397: 386: 383: 368: 365: 354: 351: 329: 326: 287: 284: 282: 279: 270:Main article: 267: 264: 246: 243: 222:dwarf mongoose 215:rhesus monkeys 210:bonnet macaque 201: 198: 194:oystercatchers 183:chacma baboons 174: 171: 169: 166: 162:vervet monkeys 156: 153: 136: 133: 112: 109: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 5089: 5078: 5075: 5073: 5070: 5069: 5067: 5052: 5051: 5046: 5042: 5040: 5039: 5030: 5028: 5027: 5018: 5017: 5014: 5006: 5005: 5001: 5000: 4999: 4996: 4994: 4991: 4989: 4986: 4984: 4981: 4980: 4978: 4974: 4968: 4967:Robert Wright 4965: 4961: 4960: 4956: 4955: 4954: 4951: 4949: 4948:Frans de Waal 4946: 4944: 4943:Steven Pinker 4941: 4939: 4936: 4934: 4931: 4929: 4926: 4924: 4921: 4919: 4918:Anne Campbell 4916: 4915: 4913: 4909: 4903: 4900: 4898: 4895: 4890: 4887: 4883: 4880: 4879: 4878: 4875: 4874: 4873: 4870: 4868: 4865: 4861: 4858: 4857: 4856: 4853: 4851: 4848: 4846: 4843: 4841: 4840:Kin selection 4838: 4836: 4833: 4831: 4828: 4826: 4823: 4821: 4818: 4816: 4813: 4811: 4808: 4806: 4803: 4801: 4798: 4796: 4793: 4792: 4790: 4786: 4782: 4775: 4770: 4768: 4763: 4761: 4756: 4755: 4752: 4745: 4742: 4741: 4737: 4728: 4724: 4720: 4716: 4712: 4708: 4704: 4700: 4693: 4690: 4685: 4681: 4676: 4671: 4667: 4663: 4659: 4655: 4652:(7): 210266. 4651: 4647: 4643: 4636: 4633: 4628: 4624: 4620: 4616: 4612: 4608: 4604: 4600: 4596: 4592: 4585: 4582: 4577: 4571: 4568: 4563: 4559: 4552: 4549: 4544: 4538: 4535: 4530: 4529: 4524: 4517: 4514: 4509: 4505: 4500: 4495: 4491: 4487: 4483: 4479: 4475: 4468: 4465: 4460: 4456: 4449: 4446: 4441: 4437: 4433: 4429: 4422: 4419: 4414: 4410: 4406: 4402: 4398: 4394: 4390: 4383: 4380: 4375: 4371: 4367: 4363: 4358: 4353: 4349: 4345: 4341: 4334: 4331: 4326: 4322: 4318: 4314: 4310: 4306: 4302: 4298: 4291: 4288: 4283: 4276: 4273: 4268: 4261: 4258: 4253: 4249: 4245: 4241: 4237: 4233: 4229: 4225: 4218: 4215: 4210: 4204: 4200: 4196: 4192: 4188: 4182: 4179: 4174: 4170: 4166: 4162: 4158: 4154: 4151:: 1299–1315. 4150: 4146: 4142: 4135: 4132: 4127: 4123: 4118: 4113: 4109: 4105: 4101: 4094: 4091: 4085: 4080: 4076: 4072: 4068: 4064: 4060: 4053: 4050: 4045: 4041: 4037: 4033: 4029: 4025: 4024: 4019: 4012: 4009: 4004: 4000: 3993: 3990: 3985: 3981: 3976: 3971: 3967: 3963: 3959: 3955: 3951: 3947: 3946:Proc Biol Sci 3943: 3936: 3933: 3928: 3924: 3920: 3916: 3912: 3908: 3901: 3898: 3883: 3876: 3875: 3867: 3864: 3859: 3855: 3851: 3847: 3843: 3839: 3832: 3829: 3825: 3819: 3812: 3809: 3804: 3800: 3796: 3792: 3788: 3784: 3777: 3774: 3769: 3765: 3760: 3755: 3751: 3747: 3743: 3739: 3735: 3731: 3727: 3720: 3717: 3712: 3708: 3703: 3698: 3694: 3690: 3686: 3679: 3676: 3671: 3667: 3663: 3659: 3655: 3651: 3644: 3641: 3636: 3632: 3628: 3624: 3620: 3616: 3612: 3608: 3600: 3598: 3594: 3589: 3585: 3581: 3577: 3573: 3569: 3562: 3559: 3554: 3550: 3545: 3540: 3536: 3532: 3528: 3524: 3520: 3518: 3509: 3506: 3501: 3497: 3493: 3489: 3485: 3481: 3477: 3473: 3466: 3463: 3458: 3454: 3450: 3446: 3442: 3438: 3431: 3428: 3423: 3417: 3413: 3409: 3405: 3398: 3395: 3390: 3386: 3382: 3378: 3374: 3370: 3366: 3362: 3355: 3352: 3347: 3343: 3338: 3333: 3329: 3325: 3321: 3317: 3313: 3306: 3303: 3298: 3294: 3290: 3283: 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