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{{Redirect|Alpha male|the slang terms for men|Alpha and beta male|dominance hierarchies in humans|Dual strategies theory|other uses|Alpha male (disambiguation)}}
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[[File:Mandrillus sphinx (alpha male).jpg|thumb|upright=1.5|A high-ranking male [[mandrill]] advertises his status with bright facial coloration.<ref name="LeighSetchell2008">{{cite journal |last1=Leigh |first1=Steven R. |last2=Setchell |first2=Joanna M. |last3=Charpentier |first3=Marie |last4=Knapp |first4=Leslie A. |last5=Wickings |first5=E. Jean |display-authors=3 |title=Canine tooth size and fitness in male mandrills (Mandrillus sphinx) |journal=[[Journal of Human Evolution]] |volume=55 |issue=1 |year=2008 |pages=75–85 |doi=10.1016/j.jhevol.2008.01.001 |pmid=18472142 |bibcode=2008JHumE..55...75L |url=https://durham-repository.worktribe.com/output/1530435 }}</ref>]]
In the [[zoological]] field of [[ethology]], a '''dominance hierarchy''' (formerly and colloquially called a '''pecking order''') is a type of social [[hierarchy]] that arises when members of animal [[social animal|social groups]] interact, creating a ranking system. Different types of interactions can result in dominance depending on the species, including [[Ritualized aggression|ritualized displays of aggression]] or direct physical violence.<ref>{{Cite journal |last1=Tibbetts |first1=Elizabeth A. |last2=Pardo-Sanchez |first2=Juanita |last3=Weise |first3=Chloe |date=2022-02-28 |title=The establishment and maintenance of dominance hierarchies |journal=Philosophical Transactions of the Royal Society B: Biological Sciences
In social living groups, members are likely to compete for access to limited resources and [[mating]] 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 animal is challenged by a subordinate one.
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== Definitions ==
'''Dominance''' 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).<ref name="Cheng 2020 pp. 238–244">{{cite journal |last=Cheng |first=Joey T. |title=Dominance, prestige, and the role of leveling in human social hierarchy and equality |journal=[[Current Opinion in Psychology]] |volume=33 |year=2020 |doi=10.1016/j.copsyc.2019.10.004 |pages=238–244 |quote=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, hierarchical relationships based on dominance are relatively less stable. |type=Review |pmid=31794955|s2cid=208627517 }}</ref> A dominant animal is one whose [[animal sexual behavior|sexual]], feeding, aggressive, and other behaviour patterns subsequently occur with relatively little influence from other group members.<ref>{{cite journal |url=https://www.researchgate.net/publication/249576147 |title=The concept and definition of dominance in animal behaviour |first=Carlos |last=Drews |date=1993 |journal=[[Behaviour (journal)|Behaviour]] |volume=125 |number=3–4 |pages=283–313|doi=10.1163/156853993X00290 }}</ref> '''Subordinate''' animals are opposite; their behaviour is '''submissive''', and can be relatively easily influenced or inhibited by other group members.<ref>{{cite journal |last1=Burgoon |first1=J. |last2=Johnson |first2=M. |last3=Koch |first3=P. |year=1998 |title=The nature and measurement of interpersonal dominance |journal=Communication Monographs |volume=65 |issue=4 |pages=308–335 |doi=10.1080/03637759809376456 |url=http://www.tandfonline.com/loi/rcmm20|url-access=subscription }}</ref>
== Dominance ==
[[File:Weeper Capuchin 01 (cropped).JPG|thumb|[[Wedge-capped capuchin]]s have a clear dominance hierarchy.]]
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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.<ref>{{cite journal |last1=Cowlishaw |first1=Guy |last2=Dunbar |first2=Robin I. M. |s2cid=53190498 |year=1991 |title=Dominance rank and mating success in male primates |journal=Animal Behaviour |volume=41 |issue=6 |pages=1045–1056 |doi=10.1016/s0003-3472(05)80642-6 }}</ref> 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, agriculture). This contradicts the "egalitarian hypothesis", which predicts that status would affect reproductive success more amongst foragers than amongst nonforagers.<ref>{{cite journal |last1=Von Rueden |first1=Christopher R. |last2=Jaeggi |first2=Adrian V. |year=2016 |title=Men's status and reproductive success in 33 nonindustrial societies: Effects of subsistence, marriage system, and reproductive strategy |journal=Proceedings of the National Academy of Sciences |volume=113 |issue=39 |pages=10824–10829 |doi=10.1073/pnas.1606800113 |pmid=27601650 |pmc=5047206 |bibcode=2016PNAS..11310824V |doi-access=free }}</ref>
High-ranking [[bonnet macaque]] 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 females; when their rank decreases, they get less time.<ref>{{cite journal |last1=Samuels |first1=A. |last2=Silk |first2=J. B. |last3=Rodman |first3=P. |s2cid=53186523 |year=1984 |title=Changes in the dominance rank and reproductive behavior of male bonnet macaques (''Macaca radiate'') |journal=Anim. Behav. |volume=32 |issue=4 |pages=994–1003 |doi=10.1016/s0003-3472(84)80212-2}}</ref> In many primates, including bonnet macaques and [[rhesus monkey]]s, 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.<ref name="Huntingford"/> 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<!--i.e. wait until they arrive at the top, the older males having died or left--> for dominance, rather than fighting for it, meaning that the alpha male is not necessarily the strongest or most attractive male.<ref>{{cite journal |last1=Widdig |first1=Anja |last2=Kessler |first2=Matthew J. |last3=Bercovitch |first3=Fred B. |last4=Berard |first4=John D. |last5=Duggleby |first5=Christine |last6=Nürnberg |first6=Peter |last7=Rawlins |first7=Richard G. |display-authors=3 |year=2016 |title=Genetic studies on the Cayo Santiago rhesus macaques: a review of 40 years of research |journal=American Journal of Primatology |volume=78 |issue=1 |pages=44–62 |doi=10.1002/ajp.22424 |pmid=26031601 |s2cid=32784846 }}</ref><ref>{{cite journal |last1=Dubuc |first1=Constance |last2=Muniz |first2=Laura |last3=Heistermann |first3=Michael |last4=Engelhardt |first4=Antje |last5=Widdig |first5=Anja |year=2011 |title=Testing the priority-of-access model in a seasonally breeding primate species |journal=[[Behavioral Ecology and Sociobiology]] |volume=65 |issue=8 |pages=1615–1627 |doi=10.1007/s00265-011-1172-8 |pmid=21874084 |pmc=3134767 |doi-access=free |bibcode=2011BEcoS..65.1615D |quote=Whereas in several primate species, males contest for rank, leading to high-ranking males being usually strong young prime males, in rhesus macaques, males mainly reach dominance through succession or queuing (Berard 1999): males slowly increase in rank as the males who dominate them die or leave the group. }}</ref>
In rodents, the highest-ranking male frequently sires the most offspring. The same pattern is found in most carnivores, such as the [[dwarf mongoose]]. 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.<ref name="Huntingford"/>
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=== Interpersonal complementarity hypothesis ===
{{main|Interpersonal complementarity hypothesis}}
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Alpha male [[baboon|savanna baboons]] have high levels of [[testosterone]] 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.<ref name= "Gesquiere"/> The mating tactics of savanna baboons are correlated with their age. Older, subordinate males form alliances to combat higher-ranking males and get access to females.<ref>{{cite journal |last1=Noe |first1=R. |last2= Sluijter |first2=A. A. |year=1990 |title=Reproductive Tactics of Male Savanna Baboons |journal=Behaviour |volume=113 |issue=1–2 |pages=117–170 |doi=10.1163/156853990x00455 |s2cid=14258881 }}</ref>
Fighting with dominant males is a risky behavior that may result in defeat, injury or even death. In [[bighorn sheep]], 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.<ref>{{cite journal |last1=Hogg |first1=J. T. |last2=Forbes |first2=S. H. |year=1997 |title=Mating in bighorn sheep: Frequent male reproduction via a high-risk ''unconventional'' tactic |journal=Behavioral Ecology and Sociobiology |volume=41 |issue=1 |pages=33–48 |doi=
[[Burying beetle]]s, 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 [[pheromone]]s and attempt to copulate before the dominant male can drive them forcefully away.<ref>{{cite journal |last1=Pettinger |first1=Adam M. |last2=Steiger |first2=Sandra |last3= Mueller |first3=Josef K. |last4=Sakaluk |first4= Scott K. |last5=Eggert |first5= Anne-Katrin |display-authors=3 |year= 2011 |title=Dominance status and carcass availability affect the outcome of sperm competition in burying beetles |journal=Behavioral Ecology |volume= 22 |issue=5 |pages=1079–1087 |doi= 10.1093/beheco/arr093 |doi-access= free }}</ref> In [[flat lizard]]s, 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.<ref>{{cite journal |last1= Whiting |first1=Martin J. |last2=Webb |first2=Jonathan K. |last3=Keogh |first3= J. Scott |year=2009 |title=Flat lizard female mimics use sexual deception in visual but not chemical signals |journal=Proceedings of the Royal Society B: Biological Sciences |volume=276 |issue=1662 |pages= 1585–1591 |doi= 10.1098/rspb.2008.1822 |pmid= 19324828 |pmc=2660994 }}</ref>
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=== Costs to subordinates ===
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. [[Brown hyena]]s, which display defined linear dominance in both sexes, allow subordinate males and females decreased time of feeding at a carcass.<ref name="Owens">{{cite journal |last1=Owens |first1=D. |last2=Owens |first2=M. |year=1996 |title=Social dominance and reproductive patterns in brown hyaenas, Hyaena brunnea, of the central Kalahari desert |journal=Animal Behaviour |volume=51 |issue=3 |pages= 535–551 |doi= 10.1006/anbe.1996.0058 |s2cid= 53163212 }}</ref> In [[toque monkey]]s 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.<ref>{{cite journal |last1=Dittus |first1=W. P. J. |title=The Social Regulation of Population Density and Age-Sex Distribution in the Toque Monkey |year= 1977 |journal=[[Behaviour (journal)|Behaviour]] |volume=63 |issue=3 |pages=281–322 |doi= 10.1163/156853977x00450 }}</ref>
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 far less with females than do the high-ranking males.<ref name="Owens"/> In [[Lycaon pictus|African wild dogs]] 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.<ref>{{cite journal |last1=Creel |first1=S. |year= 1997 |title=Handling of African wild dogs and chronic stress: Reply |journal= Conservation Biology |volume=11 |issue= 6 |pages=1454–1456 |doi= 10.1046/j.1523-1739.1997.0110061454.x |s2cid= 85088576 }}</ref>
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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 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. [[Red deer|Red stag]]s, for example, engage in exhausting roaring contests to exhibit their strength.<ref name="Huntingford 1984"/> 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.<ref name="Huntingford 1984"/>
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, [[white-crowned sparrow]]s display a unique white plumage; the higher the percentage of the crown that consists of white feathers, the higher the status of the individual.<ref>{{cite thesis |last=Laubach |first=Zachary |title=Functional Importance of Plumage Badges as Intraspecific Signals in White-Crowned Sparrows (zonotrichia Leucophrys Oriantha) |publisher=Deep Blue at the University of Michigan |date=27 November 2012 |hdl=2027.42/77948 |url=http://hdl.handle.net/2027.42/77948 |type=Thesis }}</ref> 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 [[rhesus monkey]]s, 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 [[Canada goose]] 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.<ref>{{cite book |last=Yahner |first=Richard H. |title=Wildlife Behavior and Conservation |___location=New York |publisher=Springer |year=2012 |pages=95–100}}</ref>
== Regulation mechanisms ==
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