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Dominance hierarchy (DH) is never bi-(inter-)sexual. Rare ostensible female dominance (over males) is not even amenable to modelling, and actually is male non-engagement in dominance terms (deference, not submission). Any female hierarchy is not DH, as it is not contest-based, but from either mutually signalled differential fecundity or maternal rank inheritance. Otherwise, apparent female hierarchy is either non-dominance-based first-among-equals-winner-takes-all-sociality of a female sole reproducer, or a loose artefact of ad hoc resource competition. DH entails neuro-hormonally processing winner/loser effects, for which there is evidence only in males, and requires the Y chromosome's SRY gene. Male-specificity is anticipated from the male root function of genetic filtration / mutational cleansing, necessitating male ranking in terms of genomic integrity.
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Photograph by https: //
Steve Moxon
Dominance hierarchy (DH) is never bi-(inter-)sexual. Rare ostensible female dominance
(over males) is not even amenable to modelling, and actually is male non-engagement in
dominance terms (deference, not submission). Any female hierarchy is not DH, as it is not contest-
based, but from either mutually signalled differential fecundity or maternal rank inheritance.
Otherwise, apparent female hierarchy is either non-dominance-based first-among-equals -- winner-
takes-all -- sociality of a female sole reproducer, or a loose artefact of ad hoc resource competition.
DH entails neuro-hormonally processing winner/loser effects, for which there is evidence only in
males, and requires the Y chromosome’s SRY gene. Male-specificity is anticipated from the male
root function of genetic filtration / mutational cleansing, necessitating male ranking in terms of
genomic integrity.
Keywords: dominance hierarchy, male-specific, contest-based, male deference, winner effect
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A full century on (1922-2022) from the inception of the concept of dominance hierarchy
(DH), remarkably it is still beset with serious misconceptions, none of which even were broached
when marking the centenary in the special themed issue on DH of Philosophical Transactions of
the Royal Society (compiled and edited by Strauss, Curley, Shizuka & Hobson, 2022).
Generally understood as a significantly stable, linear and transitive rank ordering of the
(same-sex, usually male) individuals of a group, created by a partial permutation of mutual
contests, DH has the putative function of differentially allocating resources generically (and, in a
rationalised and ritualised way, to reduce agonistic interaction). This longstanding operational
(working) definition of DH is undermined empirically when attempts are made to determine a
DH within any group of conspecific individuals. This suggests the nature of DH as usually
assumed is merely a derived or subsidiary function hiding a narrower, core one; and/or unrelated
phenomena are being included within an umbrella too loosely labelled DH. Funkhouser et al.
(2018) complain that “dominance ranks depend on the behavioral context, and many studies
have derived different rank orders for the same group across different measured behaviors
(Norscia & Palagi, 2015; Vervaecke, De Vries & Van Elsacker, 2000; Bayly, Evans & Taylor, 2006;
Paoli, Palagi & Tarli, 2006)”. Context-dependent hierarchy is a well-recognised phenomenon and
thus termed (e.g Nagy et al., 2013; Hewitt, Macdonald & Dugdale, 2009). Finding similarly that
DH is “situation- or resource-specific”, Lanctot & Best (2000) point out that “… dominance
measurements that were conducted in similar environmental contexts, regardless of the
response variable recorded, ranked animals similarly”. The problem, then, is not modes of
measurement, but of the unacknowledged differences in what is being measured, with
competition scenarios mistakenly regarded as being generic.
This pervading issue with DH has so long been recognised that there is a now thirty-year-
old review of pertinent studies by Fedigan (1992), who neatly summed up the problem of taking
as generic observation of conflict over a particular type of resource (the most obvious one, of
In fact, it has been found that food tests do not generalize to other conflict situations in any
consistent way. Not only does the test-situation only exist in the artificial laboratory-test
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setting, it has been found that priority to food does not necessarily correlate with priority to
other incentives, and that dominance determined through dyadic tests does not generalize
to dominance relationships for the same individuals within the group as a whole. … Rather
than peeling away the layers of the behavioral onion, to arrive at the core of an underlying
‘real’ dominance rank or dominance relationship, it can be argued that the experimenter has
in fact created the dominance relationship.
Taking all observations of competitive (or seemingly competitive) behaviour as
ubiquitously pertaining to the formation or maintenance of DH is likely to encompass far more
than the phenomenon at issue. It’s a superficial, top-down view, when examination bottom-up
from biology is required to ensure a clear, deeper understanding; if not, indeed, to uncover the
actual function that otherwise may have been largely or wholly hidden. Given that reproduction
is central to biology, its differential allocation a priori is the chief candidate for the function of
DH, rather than reproduction somehow being considered just one type of resource in a range of
them, with all being allocated differentially by DH. Reproduction is, after all, the supreme goal,
not a resource. Resources are means to ends, and all ultimately are instrumental to reproduction.
This basis of DH was put forward by the present author in 2009 as a formal hypothesis, including
that differential reproductive self-suppression mechanism appears to be integral to DH, which
last, if accurate, would strongly support the core hypothesis. [Note the present updated review of
DH is to better specify its nature (as male-only), not to revisit integral reproductive self-
suppression (which would be for another paper).]
The function of DH pertaining to reproduction rather than resources entails DH being
same-sex, given this is necessarily the nature of competition over reproduction, with individuals
attempting to better assert or display their mate value relative exclusively to same-sex others, so
as to be preferentially chosen as a sexual partner by those of the opposite sex deemed to possess
sufficient mate value. Given that the female is the limiting factor in reproduction, it would be
expected that competition over reproduction would be very different according to sex.
Unsurprisingly, to cite the title of a 2015(a) review by the present author, competitiveness is
profoundly sex-differential, consistent with being biologically based and within-, not between-sex.
No sense can be made of inter-sexual competition, including that which supposedly is indirect. If
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a strategy employed by one sex interferes with that of the other (sexual conflict), this always
resolves to intra-sexual competition. [This is revealed in considering even an extreme form in
male-perpetrated infanticide (to render the female sexually receptive); among lions, for example.
The mother as well as the male achieve fitness gains in terms of offspring quality (essential given
an extremely high offspring mortality rate; more important than number) relative to same-sex
rivals.] The reality of DH as being same-sex (and usually if not overwhelmingly specifically male)
is attested by this being so well and long recognised as hardly to require comment or even
acknowledgement, so although it is explicitly stated in the literature, such is uncommon.
That DH is a same-sex phenomenon is confirmed by the discovery of a mechanism
whereby sexual behaviour is default, to be overridden, in males, by behaviour relating to DH
only if encountered other individuals are not of the same sex. The research of Stowers et al.
(2002), echoed by Leypold et al. (2002) and Kimchi, Xu & Dulac (2007), shows that in a mammal
model there is a profound (so almost certainly evolutionarily highly conserved) neural algorithm
whereby sexing precedes any consideration of how to engage an encountered conspecific other.
In male individuals there is then applied a decision rule whereby if the other individual is female,
courtship behaviour is employed. Should the other individual instead be male, engagement is in
DH (dominance-submission) terms. This root decision-making architecture precludes any
possibility of bi-(inter-)sexual DH. Elaborating the mechanism of this mating/aggression switch,
very recently there has been the outlining of what would appear to be both its basis neurally
(Yamaguchi et al., 2020) and in terms of the pertinent hormonal receptors (Olivier & Olivier,
By this analysis, DH can be properly understood as a ramification of sex and the sexes,
stemming from what has been dubbed the genetic filter (Atmar, 1991) or mutational cleanser
(West-Eberhard, 2005) root function of the male, in turn from the function of sex and the sexes
to maintain genomic integrity (Moxon, 2019, after Gorelick & Heng, 2011). [Sex and the sexes do
not function to create variation as assumed. That notion arose from a vast over-estimation of the
variation produced by sexual reproduction, in a false analysis of meiosis by Weismann back in
1891 (Gorelick & Heng, 2011; Gorelick & Villablanca, 2018). An historical mistake has been missed
and forgotten, not least through its seeming utility in providing a major source of variation to
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drive evolution, thereby underpinning the theory. Entrenchment ensued with the great volume
of research on the major question as to what can be the utility of sex given its twofold extra cost
in comparison to the asexual mode. The paradigm persists despite (the above authors review)
overwhelming evidence from many disciplines and fields that it is asexual reproduction that
creates the greater variation.]
This male core function entails male same-sex assortment according to relative absence of
deleterious genetic material (rather than relative presence of good genes, which is the obverse,
lesser consideration; a corollary and less accurate criterion -- a proxy), to allocate differential
sexual access and facilitate corresponding female mate choice. Male same-sex DH is necessary to
facilitate this.
Note that with its concerning reproduction and not resources, DH is not only same-sex but
likely also male-specific, in that males compete primarily for reproduction, whereas females
compete primarily for resources. See Benenson & Abadzi (2020), specifically in regard to the
human case. Williamson et al. (2018) point out that in contrast to females:
males typically compete first for mates and only compete for resources insomuch
as they provide further access to mates. Recent work in house mice suggests that
this pattern exists in mice as well, with females not engaging in any more
agonistic behaviors when given the opportunity to compete for males (Weidt et
al., 2018) (p122-123). Females have distinct evolutionary mechanisms from those
of males that drive intra-female conflict, as they compete primarily for resources
to increase chances of survival for themselves and their offspring, where males
compete primarily for mates (Stockley & Bro-Jørgensen, 2011)” (p332-333).
It becomes apparent that in an important sense a superficial view of DH is so badly
mistaken as to be an inversion of a proper understanding of the phenomenon, in that rather
than it being damped down, conflict instead needs actually to be generated in order that relative
mate value -- male relative mate value, that is -- can be tested and a rank order established in
these terms. This is evident in a profound sex dichotomy in stress response, as reviewed in
2015(b) by the present author: Stress mechanism is sex-specific: female amelioration or escape
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from stress to avoid compromising reproduction contrasts with male utilisation or in effect
manufacture of stress to fulfil male genetic filter function. Once established, a hierarchy itself
further drives conflict between individuals of near-adjacent ranking, and does not reduce
conflict between individuals ranked far apart. The latter have little or no reason to engage in
contest, given the chances of a rank reversal are so very low. The disparities underpinning the
rank differential would be so apparent that thus cementing them in DH ranking hardly would be
a requirement to preclude the low-ranked individual from mounting a challenge. This is not so
in the case of more nearly adjacent rankings, however; the very opposite occurs. Here it is the
rank ordering itself that might prompt renewed contest, with a rank being such a reliable overall
summation of an individual’s strengths. It is by virtue of the closeness in ranking that a lower
ranked individual might well consider challenging another individual of higher albeit not too
dissimilar rank, to try to switch their respective rank order, given the potential gain outweighs
any costs of losing, making it worth the risk. Note, given transitivity, that this one contest may
do for the other potential contests in a permutation of those dyads within the compass of the
rank differential between the two individuals. By contrast with the significant potential gain,
there would be no direct cost of losing in terms of ranking, albeit its subordinate aspect is
underlined in a loser effect (see below), corresponding to the winner effect (again, see below)
that would be yet another pay-off if the contest were successful.
A usual assumption, then, that DH functions to ameliorate agonistic interaction, does not
make sense, so appears to be false. With DH in place, however, it is then available for co-option
secondarily to differentially allocate resources of whatever type. As resources can be
instrumental to reproduction, this would have utility in reproductive terms as a derived,
subsidiary function, and here there would be a service of rendering unnecessary any fighting that
otherwise may occur. However, resource categories differ in how relevant they may be to
reproduction, so it would be anticipated that there may be contest entirely outside of DH,
and/or the strength or reliability of any employment of DH might vary correspondingly. Thereby
are accounted the problems in determining any consistent rank order and linearity outlined
herein at the outset: of their varying according to context.
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Given that the genetic filter function is purely male -- in effect quarantined on the male
side of the lineage away from the female (necessarily so, owing to the female being the limiting
factor in reproduction, through gestation, lactation, etc; making it important to avoid any
further hindrance) -- then there is no basis here for female same-sex DH. Neither would it have
any basis in corresponding competition over female mate value, as this is by the narrower
criteria of fecundity, which is not indicated by fighting ability or any general measure of quality.
Instead, a combination of signs of developmental stability encapsulated in bodily symmetry and
of physiological fertility in youthfulness (what in human terms we would call beauty) would
sufficiently indicate fecundity; with this being immediately all too apparent in physical
appearance. Any facet of fecundity that is not visible could be mutually signalled. No
behavioural, attitudinal or ability testing is required. Consequently, there would be no utility in a
contest-based female hierarchy in respect of reproduction.
Neither regarding resource allocation: with the female necessarily having evolved to be
risk-averse, thereby to avoid any compromise of fecundity, females hardly would be adapted to
engage in agonistic contest if its purpose is merely its own subsequent reduction. Not only may
injury risk not be sufficiently reduced with ritualisation of combat, but initially determining
relative rank would entail a higher level of agonism necessary to reveal salient individual
differences in fighting ability, requiring a great deal of physiological effort, not unlikely to
significantly threaten fecundity. Even if there were a benefit to the female of ameliorating
agonistic contest, the determination of rank itself would risk driving an escalation of agonistic
interaction to a level well above that from which the female subsequently would be spared by the
The only contest-based agonism for which it would be worth females taking the risk of
significant injury is the winner-takes-all scenario of determining sole breeder status. But rather
than a hierarchy, this would be a sociality of what could be dubbed first-among-equals (winner-
takes-all): one apex individual with everyone else equally non-eminent also-rans. A DH would be
an alpha plus a beta, a gamma, and a delta, etc. -- a multi-ranked order. Albeit a DH may not be
perfectly linear, in featuring shared ranks, especially further down the hierarchy (it may vary in
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steepness along an egalitarian-despotic dimension), this is entirely different to no hierarchy at all
and a single apex individual, which indicates a very different mode of formation. Either way,
then, in the female case, proper agonistic contest and DH seem not to go together. Any apparent
female DH is likely to be either contest-less hierarchy or ad hoc low-level competition.
A contest-less ordering of reproductive privilege in terms of relative fecundity, inasmuch
as it isn’t self-evident, could be achieved simply through mutual signalling. Alternatively, it could
be the outcome of ad hoc competition over resources, where females avoid the risk of
compromising their fecundity by contesting other than agonistically, or with no more than mild
agonism. Yet this would provide no utility in outcomes of contests being registered in a way that
impacts on future encounters. No value would be conferred by DH to females, unlike for males.
Theoretically, there appears to be no conceivable utility to females of the contest-based
determination of consistent hierarchy that would be adaptive for males.
Just such a sex dichotomy in reality for humans is recognised by Benenson & Abadzi (2020)
in the title of their paper, Contest versus scramble competition: sex differences in the quest for
status. Zilkha et al. (2023) outline something similar: they find in mice “sex-specific behavioral
strategies to establish a stable hierarchy ... wild females are characterized by more pro-social
behavioral traits, whereas wild males seem to be characterized by agonistic behaviors, social
abstinence, and hiding (p9) .... the social organization formed in male mice is mostly dependent
on their degree of aggression. In contrast, social organization formed in female mice seems to
depend more on social factors such as familiarity and physiological factors” (p10). “Specifically,
males engaged in significantly more agonistic and less pro-social behaviors compared with
females, in both lab and wild mice …” (p2). Zilkha et al. point out (citing multiple studies re each
species) that their findings are widely applicable: “Similarly, extensive sex differences in the
dynamics and behavioral strategies undertaken to form hierarchies were also found in other
mammalian species, including hyenas, foxes, rhesus monkeys, baboons, and even humans” (p10).
Stockley & Bro‐Jørgensen (2011) concur with Zilkha et al. that females employ far less aggression
and instead use alliances competitively. And in a major set of studies of hierarchy in mice,
Williamson et al. (2018) find that “females indeed exhibited significantly less aggression than
males, as is demonstrated by their scores on the various measures examined”. Even more
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profoundly, in a social insect species (in this case a paper wasp), Brown (2019) finds no
aggression at all in the replacing of the dominant individual by a successor.
Social insects are the great bulk of species featuring ostensible female DH. In most cases
there is a sole breeder (a queen), in an afore-mentioned first-among-equals (winner-takes-all)
sociality (a pre-eminent individual with all others un-ranked also-rans, not an alpha + beta +
gamma + delta, etc., as in DH). There also may be a deputy: a potential replacement queen. That
in some species there appear to be a handful of apex individuals in a seeming short hierarchy, is
likely an over-interpretation of a queen plus a deputy and one or two others who fell not far
short of them in fecundity. Communication of social information is via a set of complex organic
chemicals on the outer surface cuticle -- consequently known collectively as cuticular
hydrocarbons (CHCs). These very recently have been shown in several species to encode
fecundity indicators, that in their mutual comparison between individuals decides who becomes
the sole breeder, without any agonistic contest. “There were no dominance hierarchies”,
conclude Abril & Gomez (2020), instead “intrinsic physiological differences among queens”.
Cuticular hydrocarbons correlate with fertility, not dominance is the title of Izzo, Wells, Huang &
Tibbetts’ (2010) paper on a wasp species. Leonhardt et al. (2016) outline that “In communal or
facultative eusocial species where several females compete for reproductive dominance, the
relationship between CHC profile and ovarian activity informs both competitors and potential
helpers about the reproductive potential of each female and can be used to establish dominance
and induce helping behavior.” In confirmation, Honorio, Châline & Chameron (2019) find there
are “pre-existing differences in putative fertility signals”, and that “idiosyncratic differences in a
putative fertility signal (and therefore presumably in ovarian activity) between workers in the
queen's presence reliably predict the outcome of reproductive conflict after queen loss”. Yagound
et al. (2014) elaborate: “status discrimination abilities were in fact sufficient for the establishment
and stabilization of linear hierarchies. The observed level of accuracy allowed fine-scale
discrimination of all top rankers' hierarchical status. Low-ranking workers did not exhibit such
fine-scale status discrimination. We moreover showed that a putative signal of fertility, 13-
methylpentacosane, precisely labelled the workers' position in the hierarchy”. Building on this,
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Yagound et al. (2015) state: “Furthermore, this compound might play a key role in the
establishment of the reproductive hierarchy, since workers with low fertility at the onset of
hierarchy formation already have relatively high amounts individuals with experimentally
increased amounts of 13-MeC25 triggered less agonistic interactions from top rankers, in accord
with them ‘advertising’ higher status”.
More revealing still, cuticular hydrocarbon profiles differ between ant body parts, to cite the
title of a 2021 paper by Sprenger, Gerbes, Sahm & Menzel, building on the discovery of location-
specific cuticular hydrocarbon signals in a social insect by Wang, Goodger, Woodrow & Elgar in
2016. This would appear to account for the various forms of particular physical interactions
between individuals that have been assumed to be agonistic contest but, without imputing
function, more objectively are rubbing or grasping manoeuvres involving different, specific body
parts. For a listing and description of these, see Monnin & Peeters (1999). Albeit their
classification is specifically in respect of an ant species and its relatives, similar pertains across
social insects. The key body parts involved are the antennae and the mandibles, both sets of
which are known to feature sensory organs (sensillae) to detect CHCs (e.g Ozaki & Wada-
Katsumata, 2010), and mandibular glands themselves produce oderants (Ferguson, 2021), as do
antennae hence antennal so-called duelling or boxing.
Sasaki et al. (2016) find in an ant species that the great bulk (86%) of interaction features
antennal duelling, with most of the rest (10%) involving the mandibles. Honorio, Châline &
Chameron (2019) describe what is usually denoted biting as being “when the individual uses its
mandibles to grip a part of another individual’s body. In most instances, biting was prolonged ...
with no apparent damage or cuts, and we consider this behaviour ritualized biting”. This is, as
the authors thus label, a grip rather than a bite, and with no more obvious ostensible agonistic
behaviour in social insect repertoire, it is a more than merely plausible hypothesis that social
insect ostensible agonistic interaction in general actually is to facilitate the tactile contact
required to accurately detect honest fecundity signals.
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A class of modes of determining the female sole (or parallel multiple) breeder(s) is dubbed
convention-based dominance, as termed and neatly summarised by Tibbetts, Pardo-Sanchez &
Weise (2022), (though again, note, the absence of either hierarchy or contest means it is
misleading and unwarranted to employ dominance terminology):
In societies with convention-based dominance, individuals have unique attributes that single
them out as the next dominant, (e.g., age, tenure in a group, or maternal rank) without
reflecting intrinsic characteristics that allow individuals to win contests (Lewis, 2008;
Horrocks & Hunte, 1983). For example, some social insects determine dominance based on
the seniority convention. The oldest worker is the most dominant and will take over if the
queen disappears (Strassmann & Meyer, 1983; Taylor et al., 2020). Nepotistic hierarchies are
common dominance conventions where dominance rank is inherited from the mother.
Juveniles acquire status immediately below their mother, with younger offspring outranking
older siblings (Holekamp & Smale, 1991). … Although nepotistic hierarchies are considered
convention-based, rank inheritance depends on support from mother, kin and coalition
members to ensure offspring acquire the appropriate rank (Strauss & Holekamp, 2019;
Chapais, 1992).
Tellingly, while inheritance of rank by females from their mothers is known for many
species, there is no corresponding case of inheritance of rank by males from their fathers
(Mattison et al., 2019). Inheritance from the mother, like succession by age, occurs only for
females, not for males, in line with neither inheritance nor age concerning contest, hierarchy or
Convention appears to apply in particular when individuals are closely related, and,
therefore, there are inclusive fitness considerations. A coalition of close relatives is familial
sociality, and the pre-eminent female of a family hardly could be considered to be at the apex of
a DH. Familial relations are in terms of age-based seniority and the simple scramble of sibling
rivalry, not dominance, for which there is no use, as reproduction (sexual access) is not at issue,
with mating being extra-familial, as it has to be to avoid in-breeding depression. Even a
derivative DH function of ameliorating aggression is not needed, as close relatedness
(consanguinity) would preclude fierce rivalry, and when, in some species, it is necessary to cull
weak offspring, amelioration of ferocity would be the very antithesis of what is required.
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A famous case of falsely imputing female DH when sociality instead is familial is that of the
wolf. L David Mech, the pre-eminent authority on the animal, had previously described the wolf
in DH terms, but completely revised his position upon discovering all groups in the wild are
established by mating and having offspring, so that rather than an alpha female (and male) the
supposed top-ranked individual should be designated breeder or mother/father (Mech, 1999).
Long before, Mech (1970) had realised that what had been taken to be submission actually is
simply a food-begging gesture or a food-gathering motivator by offspring to the parents (who
monopolise food in order to allocate it, making sure younger offspring get their share). Agonistic
behaviour is more or less confined to driving offspring away from the group upon their reaching
maturity. Instead of a DH there is simply seniority (age-based ranking). Issue is taken with
Mech’s position by Cafazzo, Lazzaroni & Marshall-Pescini (2016), who claim there is DH, yet
agree there is age-based determination, which being convention-, not contest-based hierarchy, is
a hallmark of familial sociality, not DH. In any case, Cafazzo et al.’s work is in respect of captive
wolves, and it’s the contrast with wild populations that is Mech’s point. It is well recognised that
in general across fauna, the close confines of captivity engenders repeat conflict that would not
occur in the wild, to the point that natural sociality can break down, with some individuals
becoming withdrawn through the experience of multiple negative encounters. There is no reason
to assume this is sub-dominance in a DH if the species does not display it outside of captivity. In
the wild, Mech’s conclusions appear likely to apply across the Canidae family of species, with
even domestic dogs (dogs long bred in captivity) also being known to have an age-based tolerant
hierarchy when feral (Bonanni, 2017). As already mentioned, some social insect species also have
age-based female hierarchy (e.g., Bridge & Field, 2007).
The meerkat is another case in point: the sole-reproducer female is determined by
seniority, primarily, and secondarily, weight (Thavarajah, Fenkes & Clutton-Brock, 2014),
without contest. It would seem in effect that differential weight given optimum reproductive age
and experience is a ready-reckoned proxy for fecundity, settling the matter as to which
individual is to take the reproductive crown without need of agonistic interaction. Weight would
be an excellent indicator of overall condition and stored reserves available to be utilised in
feeding and protecting offspring. Meerkat communities consist of the single breeding female and
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her mate, with everyone else non-breeding alloparents, most if not all individuals (of both sexes)
being the pair’s offspring. This is essentially familial sociality. [Note the males don’t form a DH,
as they have been well tested (after dispersal from the natal group upon reaching sexual
maturity), in having to survive and thrive either individually or among fellow male rovers to the
point of out-competing other males in achieving sufficient weight to show a pre-eminent genetic
fitness (genomic integrity), thereby to join a group as the sole male reproducer.]
Finding “a sex-specific mechanism underlying social hierarchies in mice”, Van den Berg,
Lamballais & Kushner (2015) conclude that this is “whereby males are strongly influenced by
prior experience, whereas females are reliant upon intrinsic attributes”. By prior experience the
authors mean previous contest, of course. Furthermore, the male-only facility to make use of
these they find arises from the Y chromosome’s SRY gene, so the mechanism cannot be female,
as anyway is clear from its being known to be mediated by testosterone. This facility is known as
the winner effect, when a prior win provides a physiological (and, in higher animals, a
psychological) boost to future winning, and/or the loser effect when a prior loss provides a
physiological (and, in higher animals, a psychological) brake, predisposing to future losing. By
contrast, for females there are no such shifts, and instead each encounter entails mutual
assessment anew, irrespective of past dealings. Without carrying over of an impact from one
encounter to another, there is no basis for ranking in females. Hierarchy may be aparent and it
may be fairly linear, but it would be simply an artefact of differences between individuals
according to basic criteria that are so obvious as not to require the sort of testing to uncover
underlying difference in which males engage.
Winner-loser effects are shown to produce a DH in modelling using cyber interactants in a
computer simulation known as DomWorld. With no predication other than the ability of
interactants individually to process winner-loser effects, then (as long as there is no constraint on
the option to escape) the result is a self-organising hierarchy of similar linearity to DH in reality
(Van Haeringen & Hemelrijk, 2022). As winner-loser effects are also the basis of hawk-dove game
theory, unsurprisingly, similar results are obtained in thus modelling DH (Huang & Wu, 2022).
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In real living interactants, a winner effect has to be “evolutionarily wired” (Morgulev & Avugos,
2020), with specific neuro-hormonal circuitry required to process it (Robertson, 2012), now more
fully outlined (Choi et al., 2023), necessitating a genetic basis, of which there are clear
indications (e.g Franz et al., 2015), and which van den Berg’s team confirm. That the mechanism
is implicit and not through rational acting is shown in human males by Kubitz, Page & Wan
(2023). It cannot be cognitively sophisticated, because the winner effect has been found in
invertebrate as well as vertebrate species (Hashikawa, Hashikawa, Lischinsky & Lin, 2018). It’s
long been well-documented across fauna (e.g Fuxjager, Oyegbile, & Marler, 2011; Mesterton-
Gibbons, 1999). All of these findings are exclusively in respect of males, and in human males a
number of studies find a winner effect across a range of contexts, from sports to board games and
elections, as summarised by Gorelik (2023), revealing it to be ubiquitous in male-male
competitive scenarios.
The contrasting absence of a female winner effect is newly evidenced in mammals. Dai et
al. (2022) point out that “… while in male mice repeated attacks lead to an increase in aggression
(winner effect) and preference to the winning-associated context, such behavioral changes are
not observed in female mice (Aubry et al., 2022; Hashikawa et al., 2018)”. So profoundly lacking is
a winner effect in females that Hashikawa et al. posit it is through attack failing to provide
reward to females. There has accumulated a sufficient body of research into this phenomenon in
humans to conduct a meta-analysis. Over the past 35 years, research into the impact of
competition outcome on testosterone concentrations in humans reveals an effect only in men
and not in women (Geniole, Bird, Ruddick & Carré, 2017; Geniole & Carré, 2019).
This extends to loser as well as winner effects, albeit there is a study where although there
was no female winner effect instead a female loser effect was found (Casto et al., 2020), but
finding even a loser effect in women makes this an outlier among studies. Not even a female loser
effect is found by Abad-Tortosa et al. (2019), whereas it is well evidenced by numerous studies in
men, complementing the winner effect. Page & Coates (2017) (and Gauriot & Page, 2019) looked
at both winner and loser effects in tennis players of both sexes and found “these effects did not
exist among women, a finding consistent with the hypothesis that androgens mediate winner
and loser effects”. Cohen-Zada, Krumer & Shtudiner (2017) found likewise with judo. The same
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sex dichotomy pertains in considering prestige dominance (Cheng, Kornienko & Granger, 2018).
That there is both a winner and a loser effect specifically in male mammals is shown by Chase,
Bartolomeo & Dugatkin (1994). The few rodent studies there have been show no female loser
effect (Hashikawa, Hashikawa, Lischinsky & Lin, 2018).
Authors increasingly are denoting the winner/loser effect as momentum. That for humans
this is psychological (rather than merely physiological) is outlined by Clark & Nilssen (2021),
citing in particular Iso-Ahola & Dotson (2014), who conclude “the influence of psychological
momentum seems to be greater for male than female performers”.
Even considering not testosterone but instead what is normally considered the female sex
hormone, estradiol, Scaia et al. (2018) find that whereas in males there are higher circulating
levels of estradiol after winning, there is no such change in females. So, with estradiol too there
is no basis of a winner effect in women; again, only in men. Likewise in considering not
testosterone on its own but in conjunction with cortisol the dual hormone hypothesis (Knight,
Sarkar, Prasad & Mehta, 2020). A meta-analysis of studies investigating this by Dekkers et al.
(2019) is supportive in the case of men but not women (for whom there is an almost negligible
correlation in the data). Knight et al concur that there is “stronger support for the dual-hormone
hypothesis in males than females”. The only support for a female winner effect here is in terms of
an exception proving the rule: in the case of the “special population” of the minuscule proportion
of the female population who compete at Olympic level (Casto, Hamilton & Edwards, 2009),
which would be explained by a necessarily highly unusual extent of masculinisation of such
women (hyperandrogenism), possibly mild congenital adrenal hyperplasia (CAH), producing
profoundly aberrational hormonal profiles.
Strikingly, then, there is no evidence of a female winner effect little if any for a loser effect
either: any form of momentum despite the enormous ideological motivation for it to be
demonstrated. Certainly, there is a dearth of published studies, and this is likely through null
findings failing to interest journals (as is generally the case, and specifically in respect of this
topic, given the ideological implications). Countless investigations will have gone unpublished,
and countless potential investigations will not have commenced given researchers’ anticipation
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of null findings. If a female winner effect indeed were there for the finding, this would be
prominent in the published literature starting many years ago. Albeit impossible to prove a
negative, it is possible now at least provisionally to conclude that there is no winner effect in the
female; that is, the winner effect is male-specific; likewise, regarding even a loser effect.
The profundity of the sex dichotomy in respect of the winner/loser effect is revealed by
Gorelik (2022) in pointing out that “it is not winning per se that precedes a testosterone boost;
rather, showcasing dominance over potential reproductive rivals seems to be the main cause”,
and that “… research on the effect of winning on men’s sexual interest suggests that testosterone
may likewise be upregulating men’s sex drive in anticipation of greater reproductive
opportunities (Gorelik & Bjorklund, 2015; Markey & Markey, 2011; Markey & Markey, 2010)”.
What is known about winner/loser effects further reveals that not resources but reproduction is
what is at issue with DH.
A useful test of DH as being not bi-(inter-)sexual, but instead same-sex (indeed, male-
specific), and requiring contest -- and to be a hierarchy, not simply first-among-equals (winner-
takes-all) sociality -- is to examine the ultimate manifestation of the still standard view of DH in
the unusual putative phenomenon of female dominance. This is where, in a few species, females
supposedly are dominant to males either wholly (all females dominant to all males) or partially
(a minority of females dominant to a minority of males). If even the proverbial exception to
prove the rule is absent here, then the rule would be strong indeed. Should the new
understanding of DH herein be accurate, then imputation of female dominance would be the
clearest inappropriate application of the standard view.
That the notion of female dominance (over males) not only is a priori theoretically
misguided in terms of an understanding of sex and the sexes but makes no logical sense, is clear
from the corollary that in the usual case (in the vast majority of species: all those not supposedly
female dominant), there would be correspondingly male dominance (that is, over females). Even
assuming there could be such a thing as bi-(inter-)sexual DH, there would not be 100%
dominance in one direction, given that in most species there is a degree of overlap across the
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sexes in measures of key factors (e.g. size, weight, and associated strength) that in males greatly
contribute to gaining dominance. If dominance were inter-sexual it would be anticipated that in
all species, other than the most extremely sexually dimorphic, a small proportion of males (the
most sub-dominant) would be sub-dominant not just to most males but also to a minority of
females. A coterie of females would be sufficiently large, heavy and (especially if unusually
masculinised) strong to outdo a minority of males in these male very terms, thereby to be
dominant over them. That across species this is very much not the case, and certainly not
generically, demonstrates that a relationship other than dominance pertains between the sexes.
If male dominance does not exist, then neither can female dominance.
To try to account for rare female dominance, Lewis (2020) puts forward a wider
conceptualisation, of “female power”, in terms of “leverage”, in particular through a male-biased
sex ratio. Lewis is quite vague, though intimates this power involves sexual favours, yet this is
implausible. The female already has leverage in being the limiting factor in reproduction through
being the gestating and lactating sex. Inclusive fitness considerations of local reproductive
output require evolved mutualism (accounted for by any of several empirically equivalent
theoretical outlines without entailing group selection) to drive male protection of and/or
giving precedence to the female. This could be a facility that is initiated or upped according to
fecundity. Its absence would be to the detriment of local reproduction generally, thereby
negatively impacting individuals within the local group generically, either indirectly and/or
The most well-known species said to sport female dominance is the spotted hyena, yet
according to McCormick et al. (2022) “without any observed provocation” males “submit” to
females, begging the question of why males would signal subdominance in the absence of
signalled dominance. It would seem the male is simply deferring to the female, as males would
be expected to do. It’s possible males employ a sub-dominance signal to acknowledge registering
a female’s presence and will not compete with her for food or other resources, in a co-option for
want of a more specific signal to indicate non-engagement in dominance terms. McCormick et
al. state their study was to try to decide between “intrinsic attributes” (aggression) and social
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support for the basis of male-female interaction, and “were unable to assess the leverage
hypothesis”. In any case, hyena females do not contest for rank and instead acquire it by
convention, in this case by inheritance from the mother (e.g Engh, Esch, Smale & Holekamp,
2000), so if there is hierarchy it is not DH, and hence there could not be a dominance-
submission interaction even in principle.
A male act of deference is explicit non-engagement. It is nothing to do with female
leverage. Not initiation of courtship or signalling of male mate value, if anything it is a signal of
non-courtship; a retiring gesture indicating insufficient own mate value to warrant initiating
courtship. Male deference is fairly obviously evolved male consideration for or protectiveness
towards the female. That male deference has been assumed to be sub-dominance (submission) is
likely through there being little or no difference in the form of the behaviour from that observed
between males where the function indeed is sub-dominance. A staple of the evolutionary process
is that forms, whether morphological structures or genetically based behaviours, are forever
being co-opted for different function; exaptation, to use the term devised by Gould & Vrba.
Selection forces can work only on what is already present, utilising different aspects or
combinations, possibly aided by a new or hitherto non-utilised mutation. All evolution is co-
option (or exaptation) of some kind. McLennan (2008) sums this up as “… traits that had evolved
under one set of conditions were co-opted to serve a different function under a second set of
conditions. … Behavior, like morphology, has both structure (what it looks like) and function
(what it does). Julian Huxley, a founding father of ethology, had the fundamental insight that
behavioral displays often evolve in one context, then change function later in evolution”. What is
here seemingly male sub-dominance is a co-option of the same or similar form to become
functionally a quite different signal more appropriate for the context: to reassure the female that
the male is not going to behave agonistically or in some other obstructive or deleterious manner
towards her.
Most female dominant species are lemurs (that for brevity are here considered together,
though papers cited may be in respect of one particular species). Depending on the species, they
display a range of putative female dominance, from complete to merely partial. Yet the behaviour
is anything but that of dominance-submission. Males are very rarely aggressive toward females,
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even if provoked, according to Radespiel & Zimmerman (2001), who also find that the males
simply avoid females, so that invariably the female prevails (in 99.9% of all encounters). Film of
lemurs in TV documentaries (e.g BBC Wildlife on One, 2002) show males placid and non-
resisting when females simply take food out of their hands, in what is no sort of agonistic or any
kind of interaction, with neither active yielding nor request. “Dominance within sexes differs
from that between sexes”, Pochron et al. (2003) argue in respect of a sexually monomorphic
(same-sized) lemur species, where “If females win a disproportionate number of interactions, it
is likely that males allow this to happen”; this being in feeding contexts, where “males may trade
off the immediate caloric reward available to winners for increased reproductive success” [p182].
If inter-sexual outcomes were in terms of dominance, size would be a major factor in who wins
contests, but maturity, not size, is determinant (Voyt, Sandel, Ortiz & Lewis, 2019).
It has long been concluded that the phenomenon indeed is simply male deference,
through female feeding priority (e.g Kappeler, 1993), or more widely “cost asymmetry” (Dunham,
2008), or both feeding priority and “reproductive strategy” (White et al., 2007). White et al.
found that the top-ranked male deferred the most: the inverse of what would be expected if the
sexes were in a dominance relationship. The conclusion of male deference is reaffirmed in the
conclusions of the most recent studies/reviews, by Grebe, Sheikh & Drea (2022), and also by
Kappeler, Fichtel & Radespiel (2022), who state that there is “spontaneous male submission in
the absence of female aggression and linked to female sexual maturation”. In other words,
specifically to fertile females, males are deferent; that is, they signal non-engagement in
dominance terms. Males hardly could be submissive (sub-dominant) anyway, given the apparent
absence of a male DH: Bauer (2004) describes the male interactions as “unstable over time due
to male migration and extreme male-male competition during the brief mating season (Budnitz
and Dainis, 1975; Taylor, 1986; Sauther, 1992; Sauther and Sussman, 1993; Pereira, 1993; Gould,
1994; 1997”. Males are not sufficiently together to establish male DH as claimed. Neither would
the female hierarchy appear to be a DH: Bauer finds that establishment is according to age and
weight, adding that it is also according to aggressiveness, but it needs to be considered if any
agonism is the exercise of rank already established through age and weight criteria, in which case
ranking would not be established by contest, and there would be no female DH.
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White et al. argue that females have evolved to be more masculinised in order to better
compete intra-sexually for unusually restricted food availability, given the high energetic
demands of lemur species (though both this restriction and the demands are disputed in the
literature). On this view, female-female competition gives rise to female dominance as a by-
product. However, that the phenomenon here indeed is the male yielding rather than the female
actively prevailing, is shown by the hormonal profiles of male lemurs: “unusually high estrogen
concentrations year-round potentially facilitating male deference via male-initiated affiliation”
(Grebe, Sheikh & Drea, 2022). Male estradiol levels are described as not significantly lower than
those in females, which is highly unusual, indeed likely unique to some lemur species.
Verreaux’s sifaka, actually another species of lemur, is supposedly completely female
dominant, but this is contradicted in reports of its behaviour. Lewis, Bueno & Di Fiore (2022)
point out that “While 96% of the acts of submission were directed from males toward females,
females only won a third of their conflicts with males. … evoking submission does not translate
into winning a resource. Indeed, intersexual power is dynamic, contextual, and dependent on the
individuals in the dyad”. Voyt, Sandel, Ortiz & Lewis (2019) write:
Power relationships between mature females and males were unambiguously female biased.
Female reproductive maturity, and not body mass, predicted intersexual conflict outcomes.
Once reproductively mature, females almost never lost to males, except when females had
not yet had an infant survive past weaning. Our results are thus consistent with the
hypothesis that female leverage characterizes social structures of adult Verreaux’s sifaka
more than female dominance.
Again, a result of 100% unidirectionality is also consistent with a phenomenon other than
dominance-submission being at play (as explained above). Lewis (2019) reveals the supposed
subordination signals given out by males are other than they seem: “… males respond to
aggression by females with ‘chatter’ indicating ‘subordination’, but the pattern of ‘chatter’ is
quite different to that amongst females”. If male supposed submission signals are very different
from any signals females use to each other, then the basis of submission signalling cannot
pertain to any form of inter-sexual hierarchy, DH or otherwise. In any case, there first has to be
established if either sex actually has a DH, and also if males and females have sufficient social
proximity for there to be a putative inter-sexual DH. With males in this species dispersing upon
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sexual maturity to join other males as a floating group, then the sexes are too separated to be
able to form a putative inter-sexual hierarchy in any case. Again, no sense can be made here of a
notion of female dominance.
Modelling that purports to show primate female dominance (Hemelrijk, Wantia & Isler,
2008) actually is a failure, being circular and anyway based in assumptions so implausible as to
have nil ecological validity (that is, no application to the real world). Reviewing work using the
DomWorld simulation of interacting cyber agents, Bryson, Ando & Lehmann (2012) conclude:
“Contrary to Hemelrijk et al. (2008), DomWorld could never account for complete female
dominance, unless the starting condition had the females already in a dominant position” [p19].
The strange inbuilt assumptions of the model (perhaps the least of which is that the level or
intensity of aggression is set only slightly higher for males than for females) lead to the highly
implausible outcome that “the female would have a one-in-three chance of defeating an agent
very much her superior. This high number of ‘improbable’ outcomes is what creates the
dynamicism of the ranking system in DomWorld [p19]. Furthermore, DomWorld actually
produces nothing more than “a perfectly random dominance hierarchy … The apparent rise in
female dominance is only a trend towards complete randomness” [p19]. Bryson, Ando &
Lehmann further outline in detail other key problems with the DomWorld simulation that
together irretrievably undermine any claim for its reflecting a basis in reality for female
dominance ever to emerge. They explain why the model produces more female dominance as the
proportion of males increases:
Adding males to the system increases the rate at which random order is achieved because
males have a higher StepDom [the level or intensity of aggression set for a cyber agent in
DomWorld] than females and thus the average amount of ‘violence’ per interaction
increases with the proportion of males. Since the experimental run times are truncated at an
arbitrary but fixed period of time before the ordering of agents is perfectly random, the rate
of change determines the proportion of dominant females. [p19]
With no attempt to address, reply to or acknowledge Bryson et al.’s outline of DomWorld’s
fatal flaws by those continuing to use it as the basis of research, a recent flurry of papers
employing it make no useful contribution to the literature on supposed female dominance,
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serving only to perpetuate the notion despite its being shown to be chimeric. Even if were
accepted the falsehoods that there can be a bi-(inter-)sexual DH and that the sexes essentially
are interchangeable with only small differences between them, there is nonetheless no feasible
model that produces female dominance; only one employing outlandish assumptions designed to
generate that outcome, if not embodying key facets of the outcome within the model itself.
Recent research provides further evidence to bolster the analysis of the phenomenon of
DH as being invariably intra-sexual (never inter-sexual not either rare female dominance or
presumed usual male dominance), and, further, that DH is male-specific. That is, DH proper a
hierarchy that is formed through contest, not first-among-equals (winner-takes-all) sociality,
and that is linear and transitive is male-only, with any female hierarchy, when not more
apparent than real, a quite different phenomenon: merely signalled relative fecundity, or by
maternal inheritance (or some other convention), or ad hoc resource competition. This is in
keeping with the function of DH to allocate reproduction rather than resources, in the service of
the male genetic filter function, leaving the hitherto assumed function of DH to reduce agonistic
interaction pertaining not to male actual DH but to female sociality, that anyway is other than
This analysis hardly is of mere theoretical interest, as it relates directly to profound new
understanding of sex and the basis of the dichotomy of the sexes, without which sense cannot be
made of sociality and behaviour (human or animal). Current hegemonic extreme ideological
notions have led to nonsense notions about human social behaviour, that will come to be seen as
outlandish historical mistake. Meantime, scientific investigation and its dissemination may not
survive growing totalitarian censure preventing informed debate.
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Steve Moxon is an English independent (non-affiliated) cross-disciplinary
researcher/writer of science review papers and books outlining original theory on the
biological roots of human sociality, behaviour and psychology, with a special interest in
the sexessex-difference/dichotomy. Regularly journal-published for the past decade,
his topics include dominance hierarchy (and associated reproductive suppression), pair-
bonding, partner violence, competitiveness, stress response mechanism, the origin of the sexual divide,
and why culture is biology. Throughout is a necessary bottom-up approach, excluding all ideology: an
avowed stance against ‘PC’ (‘identity politics’), especially its core of feminism; all being non-, indeed anti-
science. Steve also researches/writes about mythologies (ancient and contemporary), these being another
window on understanding humanity; and is a songwriter, singer & guitarist. He resides in the Pennine hills
north of Sheffield, Yorkshire, where he grew up, feels at home, and can walk or cycle through the stunning
countryside of steep-sided wooded valleys and gritstone edges.
Contact details:
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Introduction Dominance relationships in which females dominate males are rare among mammals. Mechanistic hypotheses explaining the occurrence of female dominance suggest that females dominate males because (1) they are intrinsically more aggressive or less submissive than males, and/or (2) they have access to more social support than males. Methods Here, we examine the determinants of female dominance across ontogenetic development in spotted hyenas ( Crocuta crocuta ) using 30 years of detailed behavioral observations from the Mara Hyena Project to evaluate these two hypotheses. Results Among adult hyenas, we find that females spontaneously aggress at higher rates than males, whereas males spontaneously submit at higher rates than females. Once an aggressive interaction has been initiated, adult females are more likely than immigrant males to elicit submission from members of the opposite sex, and both adult natal and immigrant males are more likely than adult females to offer submission in response to an aggressive act. We also find that adult male aggressors are more likely to receive social support than are adult female aggressors, and that both adult natal and immigrant males are 2–3 times more likely to receive support when attacking a female than when attacking another male. Across all age classes, females are more likely than males to be targets of aggressive acts that occur with support. Further, receiving social support does slightly help immigrant males elicit submission from adult females compared to immigrant males acting alone, and it also helps females elicit submission from other females. However, adult females can dominate immigrant males with or without support far more often than immigrant males can dominate females, even when the immigrants are supported against females. Discussion Overall, we find evidence for both mechanisms hypothesized to mediate female dominance in this species: (1) male and female hyenas clearly differ in their aggressive and submissive tendencies, and (2) realized social support plays an important role in shaping dominance relationships within a clan. Nevertheless, our results suggest that social support alone cannot explain sex-biased dominance in spotted hyenas. Although realized social support can certainly influence fight outcomes among females, adult females can easily dominate immigrant males without any support at all.
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Social behaviors are among the most important motivated behaviors. How dopamine (DA), a “reward” signal, releases during social behaviors has been a topic of interest for decades. Here, we use a genetically encoded DA sensor, GRABDA2m, to record DA activity in the nucleus accumbens (NAc) core during various social behaviors in male and female mice. We find that DA releases during approach, investigation and consummation phases of social behaviors signal animals’ motivation, familiarity of the social target, and valence of the experience, respectively. Positive and negative social experiences evoke opposite DA patterns. Furthermore, DA releases during mating and fighting are sexually dimorphic with a higher level in males than in females. At the functional level, increasing DA in NAc enhances social interest toward a familiar conspecific and alleviates defeat-induced social avoidance. Altogether, our results reveal complex information encoded by NAc DA activity during social behaviors and their multistage functional roles.
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Female mammals employ reproductive strategies (e.g., internal gestation) that result in power asymmetries specific to intersexual dyads. Because the number of eggs available for fertilization at any given time for most mammals is quite limited, having a fertilizable egg is potentially an important source of economic power for females. Control over mating opportunities is a source of intersexual leverage for female Verreaux’s sifaka ( Propithecus verreauxi ). We examined economic factors thought to influence the value of mating opportunities, and, thus, the extent of female leverage: kinship and market effects. Using a longitudinal dataset of agonistic interactions collected during focal animal sampling of all adult individuals in 10 social groups from 2008 to 2019, we tested the effects of relatedness, female parity, reproductive season, and adult sex ratio (population and group) on (1) the direction of submissive signaling and (2) which sex won a contested resource. While 96% of the acts of submission were directed from males toward females, females only won a third of their conflicts with males. Thus, our study has implications for evolutionary explanations of female-biased power. If female power evolved due to their greater need for food and other resources, then intersexual conflicts would be expected to result in males more consistently relinquishing control of resources. As expected, males were more likely to chatter submissively toward successful mothers, during the mating season, and when the sex ratio was male-biased. Although females generally had less power to win a conflict when their fertilizable egg was less valuable (when they were nulliparous or unsuccessful mothers or when interacting with male kin) and with an increasing female-bias in the sex ratio, this ability to win additionally was influenced by which sex initiated the conflict. Our study demonstrates that female leverage can be influenced by the supply and demand for mating opportunities, but evoking submission does not translate into winning a resource. Indeed, intersexual power is dynamic, contextual, and dependent on the individuals in the dyad.
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Distinct brain mechanisms for male aggressive and sexual behavior are present in mammalian species, including man. However, recent evidence suggests a strong connection and even overlap in the central nervous system (CNS) circuitry involved in aggressive and sexual behavior. The serotonergic system in the CNS is strongly involved in male aggressive and sexual behavior. In particular, 5-HT1A and 5-HT1B receptors seem to play a critical role in the modulation of these behaviors. The present chapter focuses on the effects of 5-HT1A- and 5-HT1B-receptor ligands in male rodent aggression and sexual behavior. Results indicate that 5-HT1B-heteroreceptors play a critical role in the modulation of male offensive behavior, although a definite role of 5-HT1A-auto- or heteroreceptors cannot be ruled out. 5-HT1A receptors are clearly involved in male sexual behavior, although it has to be yet unraveled whether 5-HT1A-auto- or heteroreceptors are important. Although several key nodes in the complex circuitry of aggression and sexual behavior are known, in particular in the medial hypothalamus, a clear link or connection to these critical structures and the serotonergic key receptors is yet to be determined. This information is urgently needed to detect and develop new selective anti-aggressive (serenic) and pro-sexual drugs for human applications.
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The extant primates of Madagascar (Lemuriformes) represent the endpoints of an adaptive radiation following a single colonization event more than 50 million years ago. They have since evolved a diversity of life history traits, ecological adaptations and social systems that rivals that of all other living primates combined. Their social systems are characterized by a unique combination of traits, including the ability of adult females to dominate adult males. In fact, there is no other group of mammals in which female dominance is so widespread. Yet, recent research has indicated that there is more interspecific variation in lemur intersexual relationships than previously acknowledged. Here, we therefore review and summarize the relevant literature, quantifying the extent of sex-bias in intersexual dominance relations documented in observational and experimental studies in captivity and the wild. Female dominance is often, but not always, implemented by spontaneous male submission in the absence of female aggression and linked to female sexual maturation. We connect the available evidence to the hypotheses that have been proposed to explain the evolution of female dominance among lemurs. The occurrence of female dominance in all lemur families and the interspecific variation in its extent indicate that it has evolved soon after lemurs colonized Madagascar – presumably in response to particular ecological challenges – and that it has since been reduced in magnitude independently in some taxa. Our study contributes important comparative information on sex roles from an independent primate radiation and provides general insights into the conditions, opportunities and obstacles in the evolution of female-biased power.
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In many groups of animals the dominance hierarchy is linear. What mechanisms underlie this linearity of the dominance hierarchy is under debate. Linearity is often attributed to cognitively sophisticated processes, such as transitive inference and eavesdropping. An alternative explanation is that it develops via the winner-loser effect. This effect implies that after a fight has been decided the winner is more likely to win again, and the loser is more likely to lose again. Although it has been shown that dominance hierarchies may develop via the winner-loser effect, the degree of linearity of such hierarchies is unknown. The aim of the present study is to investigate whether a similar degree of linearity, like in real animals, may emerge as a consequence of the winner-loser effect and the socio-spatial structure of group members. For this purpose, we use the model DomWorld, in which agents group and compete and the outcome of conflicts is self-reinforcing. Here dominance hierarchies are shown to emerge. We analyse the dominance hierarchy, behavioural dynamics and network triad motifs in the model using analytical methods from a previous study on dominance in real hens. We show that when one parameter, representing the intensity of aggression, was set high in the model DomWorld, it reproduced many patterns of hierarchical development typical of groups of hens, such as its high linearity. When omitting from the model the winner-loser effect or spatial location of individuals, this resemblance decreased markedly. We conclude that the combination of the spatial structure and the winner-loser effect provide a plausible alternative for hierarchical linearity to processes that are cognitively more sophisticated. Further research should determine whether the winner-loser effect and spatial structure of group members also explains the characteristics of hierarchical development in other species with a different dominance style than hens.
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In the decades since female social dominance was first described in strepsirrhine primates, researchers have sought to uncover the proximate and ultimate explanations for its development. In the females of various female-dominant species, androgens have been implicated as regulators of behavior and/or predictors of seasonal fluctuations in aggression (the ‘Female Masculinization Hypothesis’). Males, more generally, respond to changing social demands via seasonal fluctuations in androgen-mediated behavior (the ‘Challenge Hypothesis’), that may also entail changes in activation of the hypothalamic-pituitary-adrenal axis. Here, we explore if androgens, glucocorticoids, and intersexual behavior fluctuate seasonally in the female-dominant, blue-eyed black lemur (Eulemur flavifrons), with potential consequences for understanding female aggression and male deference. Across two studies conducted during the breeding and nonbreeding seasons, we assessed rates of mixed-sex, dyadic social behavior (aggression and affiliation) and concentrations of fecal glucocorticoid metabolites (Study 1) and serum sex hormones (androstenedione, testosterone, and estradiol; Study 2). Our results align with several predictions inspired by the Female Masculinization and Challenge Hypotheses for intersexual relations: During the breeding season, specifically, both aggression and androstenedione peaked in females, while female-initiated affiliation decreased, potentially to facilitate female resource access and reproductive control. By comparison, all target hormones (androgens, estrogen, and glucocorticoids) peaked in males, with glucocorticoid concentrations potentially increasing in response to the surge in female aggression, and unusually high estrogen concentrations year-round potentially facilitating male deference via male-initiated affiliation. These results suggest complex, seasonally and hormonally mediated behavior in Eulemur flavifrons.
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Animal groups are often organized hierarchically, with dominant individuals gaining priority access to resources and reproduction over subordinate individuals. Initial dominance hierarchy formation may be influenced by multiple interacting factors, including an animal's individual attributes, conventions and self-organizing social dynamics. After establishment, hierarchies are typically maintained over the long-term because individuals save time, energy and reduce the risk of injury by recognizing and abiding by established dominance relationships. A separate set of behaviours are used to maintain dominance relationships within groups, including behaviours that stabilize ranks (punishment, threats, behavioural asymmetry), as well as signals that provide information about dominance rank (individual identity signals, signals of dominance). In this review, we describe the behaviours used to establish and maintain dominance hierarchies across different taxa and types of societies. We also review opportunities for future research including: testing how self-organizing behavioural dynamics interact with other factors to mediate dominance hierarchy formation, measuring the long-term stability of social hierarchies and the factors that disrupt hierarchy stability, incorporating phenotypic plasticity into our understanding of the behavioural dynamics of hierarchies and considering how cognition coevolves with the behaviours used to establish and maintain hierarchies. This article is part of the theme issue ‘The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies’.
Dominance hierarchy is a fundamental social phenomenon in a wide range of mammalian species, critically affecting fitness and health. Here, we investigate the role of pheromone signals in the control of social hierarchies and individual personalities within groups of wild mice. For this purpose, we combine high-throughput behavioral phenotyping with computational tools in freely interacting groups of wild house mice, males and females, in an automated, semi-natural system. We show that wild mice form dominance hierarchies in both sexes but use sex-specific strategies, displaying distinct male-typical and female-typical behavioral personalities that were also associated with social ranking. Genetic disabling of VNO-mediated pheromone detection generated opposite behavioral effects within groups, enhancing social interactions in males and reducing them in females. Behavioral personalities in the mutated mice displayed mixtures of male-typical and female-typical behaviors, thus blurring sex differences. In addition, rank-associated personalities were abolished despite the fact that both sexes of mutant mice formed stable hierarchies. These findings suggest that group organization is governed by pheromone-mediated sex-specific neural circuits and pave the way to investigate the mechanisms underlying sexual dimorphism in dominance hierarchies under naturalistic settings.
We propose a novel model to explain the mechanisms underlying dominance hierarchical structures. Guided by a predetermined social convention, agents with limited cognitive abilities optimize their strategies in a Hawk-Dove game. We find that several commonly observed hierarchical structures in nature such as linear hierarchy and despotism, emerge as the total fitness-maximizing social structures given different levels of cognitive abilities.