1150:
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.
160:
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
151:
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,
515:
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
261:
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
1149:
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
227:
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
344:
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
332:
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 β 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
204:
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,
159:
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
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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
212:
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
150:
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
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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.
794:
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
1798:
Bonnatti, Vanessa; Luz
Paulino SimΓ΅es, ZilΓ‘; 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.
357:
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".
362:
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.
762:
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.
508:
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
996:
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
664:
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
2847:
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".
821:
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.
1850:
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.
1113:
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.
341:
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 motherβthe 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
3905:
Drummond, Hugh; Edda
Gonzalez; Jose Luis Osorno (1986). "Parent-Offspring Cooperation in the Blue-footed Booby (Sula nebouxii): Social Roles in Infanticidal Brood Reduction".
2514:
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".
3470:
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.
529:
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".
716:
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.
422:
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.
4557:
205:
agriculture). This contradicts the "egalitarian hypothesis", which predicts that status would affect reproductive success more amongst foragers than amongst nonforagers.
2310:
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".
1704:
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".
2444:
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,
691:
individuals, the opposite of what had been expected. Two core hypotheses attempt to explain this. The first suggests that higher ranking individuals exert more
1141:
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 TaΓ― National Park".
4387:
Hewitt, Stacey E.; Macdonald, David W.; Dugdale, Hannah L. (2009). "Context-dependent linear dominance hierarchies in social groups of European badgers,
1184:
Leigh, Steven R.; Setchell, Joanna M.; Charpentier, Marie; et al. (2008). "Canine tooth size and fitness in male mandrills (Mandrillus sphinx)".
349:
which live in social packs separated into male and female hierarchies, top-ranking alpha females have been observed to produce 76β81% of all litters.
3872:
897:
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
271:
2654:
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 moleβrat, Heterocephalus glaber"
4771:
3402:
Sauther, Michelle L. (1993). "Resource Competition in Wild Populations of Ringtailed Lemurs (Lemur Catta): Implications for Female Dominance".
2267:
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".
1220:
1129:
teats produce a greater quantity of milk. Once established, this teat order remains stable with each piglet tending to feed from a particular
1121:
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.; HΓΆlldobler, B. (2000). "Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator".
2170:
Owens, D.; Owens, M. (1996). "Social dominance and reproductive patterns in brown hyaenas, Hyaena brunnea, of the central Kalahari desert".
724:
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".
3435:
Young, Andrew L.; Richard, Alison F.; Aiello, Leslie C. (1990). "Female Dominance and Maternal Investment in Strepsirhine Primates".
220:
In rodents, the highest-ranking male frequently sires the most offspring. The same pattern is found in most carnivores, such as the
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123:, feeding, aggressive, and other behaviour patterns subsequently occur with relatively little influence from other group members.
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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. Dominanceβsubordination 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".
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development is not totally achieved in the worker caste, which opens the possibility of reproduction by workers. Since
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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.
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2982:"Evidence that primer pheromones do not cause social suppression of reproduction in male and female naked mole-rats"
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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
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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:
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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:
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5025:
4819:
4757:
2046:
Hogg, J. T.; Forbes, S. H. (1997). "Mating in bighorn sheep: Frequent male reproduction via a high-risk
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are suppressed, while in females it appears that the suppression involves the entire suppression of the
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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.).
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In many bird species, the dominant individuals have higher rates of food intake. Such species include
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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.
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1961:
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1019:
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38:
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2423:
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Cowlishaw, Guy; Dunbar, Robin I. M. (1991). "Dominance rank and mating success in male primates".
54:
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4408:
4369:
4320:
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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"
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1005:
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484:
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865:), they have "exceptionally high concentrations" of androgens, "particularly during gestation".
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Intruder retreats: When participants are of equal fighting ability and competing for a certain
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Drea, Christine M. (August 2009). "Endocrine Mediators of Masculinization in Female Mammals".
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1997:
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Dubuc, Constance; Muniz, Laura; Heistermann, Michael; Engelhardt, Antje; Widdig, Anja (2011).
1721:
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1201:
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4474:"Male dominance rank and reproductive success in chimpanzees, Pan troglodytes schweinfurthii"
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The asymmetries between individuals have been categorized into three types of interactions:
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972:
704:
556:
530:
475:
189:
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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".
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2718:
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studies with computer stimulated hierarchal conditions showed increased activity in the
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2500:
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1748:
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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. 35β49.
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Digby, L. I.; Kahlenberg, S. M. (2002). "Female dominance in blue-eyed black lemurs".
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1682:
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87:
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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"
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1114:
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741:
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419:
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31:
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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.
17:
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4356:
4339:
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2675:
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1863:
1820:
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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,
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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:
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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:
4718:
4683:
4614:
4558:"Sperm Competition in the Florida Manatee (Trichechus manatus latirostris)"
4507:
4365:
4316:
4243:
4164:
4125:
3965:
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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:
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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. 813β1024.
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:
GonzΓ‘lez, 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' 1927β54".
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. 156β159, 194β205, 208β209, 250β253.
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. 476β511.
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: 1391β1399.
2404:. New York: Springer. pp. 95β100.
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; HΓΆlldobler, 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. 135β152.
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 fΓΌr 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. 119β140.
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): 1533β1539.
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): 1585β1591.
2000:. pp. 254β283.
1908:2011Sci...333..357G
1813:2014NW....101...17B
1801:Naturwissenschaften
1624:2016PNAS..11310824V
1618:(39): 10824β10829.
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): 993β1000.
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. 73β74.
4350:(16): R657βR660.
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): 162β168.
3241:(6056): 697β700.
3182:(6056): 693β697.
3045:10.1644/bhe-002.1
2713:(6251): 796β797.
2576:(7014): 218β222.
1998:Blackwell Science
1718:10.1002/ajp.22424
1560:. pp. 39β54.
1485:10.1111/ibi.12447
1390:. pp. 66β85.
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:(2β3): 215β226.
2804:
2798:
2797:
2787:
2777:
2760:(9): 2945β2950.
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): 1193β217.
2307:
2301:
2300:
2275:(6): 1411β1424.
2269:Animal Behaviour
2264:
2258:
2257:
2240:(6): 1454β1456.
2229:
2223:
2222:
2202:
2196:
2195:
2172:Animal Behaviour
2167:
2158:
2157:
2147:
2115:
2109:
2108:
2106:
2097:(5): 1079β1087.
2082:
2076:
2075:
2043:
2037:
2036:
2019:(1β2): 117β170.
2008:
2002:
2001:
1989:
1983:
1982:
1964:
1944:
1938:
1937:
1927:
1902:(6040): 357β60.
1887:
1876:
1875:
1852:Insectes Sociaux
1847:
1841:
1840:
1795:
1789:
1788:
1778:
1768:
1759:(8): 1615β1627.
1744:
1738:
1737:
1701:
1695:
1694:
1662:
1656:
1655:
1645:
1635:
1603:
1597:
1596:
1579:(6): 1045β1056.
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:(3β4): 283β313.
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): 948β955.
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): 477β480.
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): 994β1003.
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:
18:Alpha (biology)
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): 131β142.
4689:
4632:
4581:
4567:
4548:
4534:
4513:
4484:(4): 873β885.
4464:
4445:
4434:(3): 273β283.
4418:
4399:(1): 161β169.
4379:
4330:
4287:
4272:
4257:
4230:(7): 847β856.
4214:
4207:
4178:
4131:
4110:(2): 703β708.
4090:
4049:
4030:(1): 431β438.
4008:
3989:
3932:
3913:(5): 365β372.
3897:
3863:
3844:(6): 425β436.
3828:
3808:
3795:10.2307/453561
3789:(4): 265β268.
3773:
3716:
3695:(1): 117β127.
3675:
3656:(4): 847β867.
3640:
3593:
3574:(4): 221β226.
3558:
3505:
3478:(4): 555β567.
3462:
3449:10.1086/285057
3443:(4): 473β488.
3427:
3420:
3394:
3367:(3): 191β199.
3351:
3302:
3276:
3225:
3166:
3139:(6): 701β709.
3123:
3108:
3079:(4): 332β340.
3051:
3038:(2): 255β264.
3013:
2992:(1): 225β230.
2969:
2920:
2893:(5): 527β531.
2874:
2855:(2): 133β142.
2832:
2799:
2740:
2697:
2646:
2627:(4): 325β332.
2611:
2560:
2541:
2506:
2487:(2): 124β132.
2471:
2452:(4): 327β334.
2436:
2407:
2392:
2367:
2337:
2302:
2259:
2224:
2213:(3): 281β322.
2197:
2178:(3): 535β551.
2159:
2110:
2077:
2048:unconventional
2038:
2003:
1984:
1962:10.1.1.319.709
1955:(3): 558β568.
1939:
1877:
1858:(2): 114β121.
1842:
1790:
1739:
1696:
1667:Macaca radiate
1657:
1598:
1563:
1529:
1510:(2): 131β154.
1491:
1478:(2): 456β462.
1458:
1447:(3): 263β274.
1428:
1409:(4): 255β264.
1393:
1384:Goat Husbandry
1374:
1361:(4): 708β715.
1341:
1330:(4): 308β335.
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:: 1299β1315.
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:
3281:
3277:
3272:
3268:
3264:
3260:
3256:
3252:
3248:
3244:
3240:
3236:
3229:
3226:
3221:
3217:
3213:
3209:
3205:
3201:
3197:
3193:
3189:
3185:
3181:
3177:
3170:
3167:
3162:
3158:
3154:
3150:
3146:
3142:
3138:
3134:
3127:
3124:
3119:
3112:
3109:
3104:
3100:
3095:
3094:2027.42/46903
3090:
3086:
3082:
3078:
3074:
3067:
3060:
3058:
3056:
3052:
3046:
3041:
3037:
3033:
3029:
3023:
3017:
3014:
3009:
3005:
3000:
2995:
2991:
2987:
2983:
2976:
2974:
2970:
2965:
2961:
2956:
2951:
2947:
2943:
2939:
2935:
2931:
2924:
2921:
2916:
2912:
2908:
2904:
2900:
2896:
2892:
2888:
2881:
2879:
2875:
2870:
2866:
2862:
2858:
2854:
2850:
2843:
2841:
2839:
2837:
2833:
2827:
2822:
2818:
2814:
2810:
2803:
2800:
2795:
2791:
2786:
2781:
2776:
2771:
2767:
2763:
2759:
2755:
2751:
2744:
2741:
2736:
2732:
2728:
2724:
2720:
2716:
2712:
2708:
2701:
2698:
2693:
2689:
2685:
2681:
2677:
2673:
2669:
2665:
2661:
2657:
2650:
2647:
2642:
2638:
2634:
2630:
2626:
2622:
2615:
2612:
2607:
2603:
2599:
2595:
2591:
2587:
2583:
2579:
2575:
2571:
2564:
2561:
2556:
2552:
2545:
2542:
2537:
2533:
2529:
2525:
2521:
2517:
2510:
2507:
2502:
2498:
2494:
2490:
2486:
2482:
2475:
2472:
2467:
2463:
2459:
2455:
2451:
2447:
2440:
2437:
2425:
2421:
2417:
2411:
2408:
2403:
2396:
2393:
2388:
2387:2027.42/77948
2384:
2380:
2379:
2371:
2368:
2363:
2359:
2352:
2350:
2348:
2346:
2344:
2342:
2338:
2333:
2329:
2325:
2321:
2317:
2313:
2306:
2303:
2298:
2294:
2290:
2286:
2282:
2278:
2274:
2270:
2263:
2260:
2255:
2251:
2247:
2243:
2239:
2235:
2228:
2225:
2220:
2216:
2212:
2208:
2201:
2198:
2193:
2189:
2185:
2181:
2177:
2173:
2166:
2164:
2160:
2155:
2151:
2146:
2141:
2137:
2133:
2129:
2125:
2121:
2114:
2111:
2105:
2100:
2096:
2092:
2088:
2081:
2078:
2073:
2069:
2065:
2061:
2057:
2053:
2049:
2042:
2039:
2034:
2030:
2026:
2022:
2018:
2014:
2007:
2004:
1999:
1995:
1988:
1985:
1980:
1976:
1972:
1968:
1963:
1958:
1954:
1950:
1943:
1940:
1935:
1931:
1926:
1921:
1917:
1913:
1909:
1905:
1901:
1897:
1893:
1886:
1884:
1882:
1878:
1873:
1869:
1865:
1861:
1857:
1853:
1846:
1843:
1838:
1834:
1830:
1826:
1822:
1818:
1814:
1810:
1806:
1802:
1794:
1791:
1786:
1782:
1777:
1772:
1767:
1762:
1758:
1754:
1750:
1743:
1740:
1735:
1731:
1727:
1723:
1719:
1715:
1711:
1707:
1700:
1697:
1692:
1688:
1684:
1680:
1676:
1672:
1668:
1661:
1658:
1653:
1649:
1644:
1639:
1634:
1629:
1625:
1621:
1617:
1613:
1609:
1602:
1599:
1594:
1590:
1586:
1582:
1578:
1574:
1567:
1564:
1559:
1555:
1548:
1546:
1544:
1542:
1540:
1538:
1536:
1534:
1530:
1525:
1521:
1517:
1513:
1509:
1505:
1498:
1496:
1492:
1486:
1481:
1477:
1473:
1469:
1462:
1459:
1454:
1450:
1446:
1442:
1435:
1433:
1429:
1424:
1420:
1416:
1412:
1408:
1404:
1397:
1394:
1389:
1385:
1378:
1375:
1369:
1364:
1360:
1356:
1352:
1345:
1342:
1337:
1333:
1329:
1325:
1321:
1314:
1311:
1306:
1302:
1298:
1294:
1293:
1288:
1281:
1278:
1274:
1269:
1265:
1261:
1257:
1253:
1249:
1245:
1241:
1234:
1231:
1226:
1222:
1215:
1212:
1207:
1203:
1199:
1195:
1191:
1187:
1180:
1177:
1170:
1166:
1163:
1161:
1158:
1157:
1153:
1151:
1144:
1142:
1140:
1136:
1132:
1128:
1124:
1120:
1116:
1111:
1107:
1098:
1091:
1089:
1087:
1083:
1079:
1078:
1073:
1069:
1065:
1061:
1057:
1056:Frans De Waal
1052:
1048:
1040:
1035:
1033:
1031:
1027:
1026:
1021:
1016:
1014:
1013:
1008:
1007:
1002:
1001:
995:
990:
988:
987:
982:
978:
974:
970:
966:
961:
959:
955:
952:
948:
944:
940:
936:
932:
928:
920:
918:
915:
910:
906:
904:
900:
896:
892:
888:
880:
875:
868:
866:
864:
860:
856:
852:
851:spotted hyena
848:
844:
840:
835:
831:
823:
820:
817:
816:
815:
812:
810:
806:
802:
798:
789:
784:
777:
774:
766:
764:
761:
757:
753:
748:
743:
739:
735:
731:
727:
719:
717:
715:
711:
710:olive baboons
706:
702:
698:
694:
690:
686:
682:
678:
675:
670:
667:
663:
659:
658:ovarian cycle
655:
651:
647:
646:
638:
633:
629:
627:
623:
619:
615:
611:
607:
603:
599:
598:
593:
590:levels after
589:
581:
579:
577:
576:
571:
570:
566:
565:carpenter ant
562:
558:
554:
550:
549:
545:
540:
538:
537:
532:
528:
527:
522:
517:
516:advertising.
514:
513:
507:
504:layer on the
503:
499:
498:
497:D. quadriceps
493:
492:
487:
486:
482:
478:
477:
473:
468:
460:
458:
456:
453:
449:
445:
441:
440:social status
437:
433:
425:
423:
421:
417:
412:
406:
404:
395:
394:common shrews
391:
387:
384:
381:
380:
379:
376:
374:
366:
364:
361:
352:
350:
348:
342:
340:
339:toque monkeys
336:
327:
325:
323:
319:
315:
311:
309:
308:bighorn sheep
304:
301:
297:
292:
285:
280:
278:
273:
265:
263:
259:
257:
253:
244:
242:
240:
239:
234:
229:
225:
223:
218:
216:
211:
208:High-ranking
206:
199:
197:
195:
191:
186:
184:
180:
172:
167:
165:
163:
154:
152:
145:
141:
134:
132:
130:
126:
122:
117:
110:
108:
105:
101:
97:
93:
89:
88:social groups
85:
81:
80:pecking order
77:
73:
69:
61:
56:
52:
48:
44:
40:
33:
19:
5048:
5036:
5024:
5002:
4957:
4953:E. O. Wilson
4923:Noam Chomsky
4889:presociality
4781:Sociobiology
4702:
4698:
4692:
4649:
4645:
4635:
4627:ResearchGate
4625:– via
4598:
4594:
4584:
4570:
4561:
4551:
4537:
4528:ResearchGate
4526:
4516:
4481:
4477:
4467:
4454:
4448:
4431:
4427:
4421:
4396:
4392:
4388:
4382:
4347:
4343:
4333:
4303:(1): 23β31.
4300:
4296:
4290:
4281:
4275:
4266:
4260:
4227:
4223:
4217:
4194:
4181:
4144:
4140:
4134:
4107:
4103:
4093:
4066:
4062:
4052:
4027:
4021:
4017:
4011:
4002:
3998:
3992:
3949:
3945:
3935:
3910:
3906:
3900:
3889:. Retrieved
3882:the original
3873:
3866:
3841:
3837:
3831:
3823:
3817:
3811:
3786:
3782:
3776:
3736:(1): 35492.
3733:
3729:
3719:
3692:
3689:Reproduction
3688:
3678:
3653:
3649:
3643:
3610:
3606:
3571:
3567:
3561:
3526:
3522:
3516:
3508:
3475:
3471:
3465:
3440:
3436:
3430:
3403:
3397:
3364:
3360:
3354:
3319:
3315:
3305:
3288:
3238:
3234:
3228:
3179:
3175:
3169:
3136:
3132:
3126:
3117:
3111:
3076:
3072:
3035:
3031:
3016:
2989:
2985:
2937:
2933:
2923:
2890:
2886:
2852:
2848:
2816:
2812:
2802:
2757:
2753:
2743:
2710:
2706:
2700:
2659:
2655:
2649:
2624:
2620:
2614:
2573:
2569:
2563:
2554:
2550:
2544:
2519:
2515:
2509:
2484:
2480:
2474:
2449:
2445:
2439:
2428:. Retrieved
2424:the original
2419:
2410:
2401:
2395:
2377:
2370:
2357:
2315:
2311:
2305:
2272:
2268:
2262:
2237:
2233:
2227:
2210:
2206:
2200:
2175:
2171:
2127:
2123:
2113:
2094:
2090:
2080:
2058:(1): 33β48.
2055:
2051:
2047:
2041:
2016:
2012:
2006:
1993:
1987:
1952:
1948:
1942:
1899:
1895:
1855:
1851:
1845:
1807:(1): 17β24.
1804:
1800:
1793:
1756:
1752:
1742:
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