Reproductive skew, costs, and benefits of cooperative breeding in female wood mice (Apodemus sylvaticus)

Two current models seek to explain reproduction of subordinatesin social groups: incentives given by dominants for peacefullyremaining in the group (reproductive skew model) or incompletecontrol by dominants. These models make different predictionsconcerning genetic relatedness between individuals for thedistribution of reproduction and the stability of cooperativebreeding associations. To test these models and to furtherexplore the relationships between reproductive skew, geneticrelatedness, and investment of each participant, we performedbehavioral observations of female wood mice (Apodemus sylvaticus)raising pups communally. Our results do not support previousmodels. Differences in lifetime reproductive success were significantlygreater within mother—daughter pairs than within pairsof sisters or unrelated females. Subordinate females of eitherbreeding unit did not differ in their direct reproduction.Calculations of inclusive fitness based on our results leadto the following predictions: (1) Communal nests should occuronly when ecological circumstances prevent solitary breeding.(2) Subordinate females gain the highest inclusive fitnessjoining their mothers; they also show the highest nursing investment.(3) Mothers should accept daughters, who have no opportunityfor solitary breeding. (4) Dominant sisters and unrelated femalesshould reject subordinate females because cooperative breedingreduces their reproductive success. However, breeding unitsof dominant sisters and unrelated females nevertheless occurand can be explained by our finding that such females significantlyreduce nursing time, which may help them save energy for futurebreeding cycles. Our data demonstrate that both genetic relatednessand investment skew are important in the complex evolutionof reproductive skew in cooperative breeding.     

Transactional skew and assured fitness return models fail to predict patterns of cooperation in wasps

Cooperative breeders often exhibit reproductive skew, where dominant individuals reproduce more than subordinates. Two approaches derived from Hamilton's inclusive fitness model predict when subordinate behavior is favored over living solitarily. The assured fitness return (AFR) model predicts that subordinates help when they are highly likely to gain immediate indirect fitness. Transactional skew models predict dominants and subordinates "agree" on a level of reproductive skew that induces subordinates to join groups. We show the AFR model to be a special case of transactional skew models that assumes no direct reproduction by subordinates. We use data from 11 populations of four wasp species (Polistes, Liostenogaster) as a test of whether transactional frameworks suffice to predict when subordinate behavior should be observed in general and the specific level of skew observed in cooperative groups. The general prediction is supported; in 10 of 11 cases, transactional models correctly predict presence or absence of cooperation. In contrast, the specific prediction is not consistent with the data. Where cooperation occurs, the model accurately predicts highly biased reproductive skew between full sisters. However, the model also predicts that distantly related or unrelated females should cooperate with low skew. This prediction fails: cooperation with high skew is the observed norm. Neither the generalized transactional model nor the special-case AFR model can explain this significant feature of wasp sociobiology. Alternative, nontransactional hypotheses such as parental manipulation and kin recognition errors are discussed.     

Reproductive skew, concessions and limited control

Models of reproductive skew in cooperative and eusocial societies suggest that dominants allow subordinates to breed to induce them to remain peaceably in the group. However, it is not yet clear how widely the assumptions of these models apply to animal societies, and many of the trends that they predict are consistent with the simpler suggestion that there is a struggle for reproduction between dominants and subordinates, whose outcome depends on the potential costs and benefits of the contest to both parties. Models of reproductive skew that incorporate contests of this kind and empirical studies that can discriminate clearly between reproductive concessions and failures of control are now needed.     

Reproductive skew and group size: an N-person staying incentive model

Transactional models of social evolution emphasize that dominantbreeders may donate parcels of reproduction to subordinatesin return for peaceful cooperation. We develop a general transactionalmodel of reproductive partitioning and group size for N-persongroups when (1) expected group output is a concave (decelerating)functiong[N] of the number N of group members, and (2) thesubordinates may receive fractions of total group reproduction("staying incentives") just sufficient to induce them to stayand help the dominant instead of breeding solitarily. We focusespecially on "saturated" groups, that is, groups that havegrown in size just up to the point where subsequent joining by subordinates is no longer beneficial either to them (in parent-offspring groups) or to the dominant (in symmetric-relatedness groups).Decreased expected output for solitary breeding increases thesaturated group size and decreases the staying incentives.Increased relatedness decreases both the saturated group sizeand the staying incentives. However, in saturated groups withsymmetric relatedness, an individual subordinate's staying incentive converges to 1 — g[N^* — 1]/g[N^*]) regardless ofrelatedness, where N^* is the size of a saturated group, providedthat the g[N] function near the saturated group size N^* isapproximately linear. Thus, staying incentives can be insensitiveto relatedness in saturated groups, although the dominant's total fraction of reproduction (total skew) will be more sensitive.The predicted ordering for saturated group size is: Parent-fullsibling offspring = non-relatives > symmetrically relatedrelatives. Strikingly, stable groups of non-relatives can formfor concaveg[N] functions in our model but not in previousmodels of group size lacking skew manipulation by the dominant.Finally, symmetrical relatedness groups should tend to breakup by threatened ejections of subordinates by dominants, whereas parent-offspring groups should tend to breakup via unforceddepartures by subordinates.     

Sociality, mating system and reproductive skew in marmots: evidence and hypotheses

Marmot species exhibit a great diversity of social structure, mating systems and reproductive skew. In particular, among the social species (i.e. all except Marmota monax), the yellow-bellied marmot appears quite different from the others. The yellow-bellied marmot is primarily polygynous with an intermediate level of sociality and low reproductive skew among females. In contrast, all other social marmot species are mainly monogamous, highly social and with marked reproductive skew among females. To understand the evolution of this difference in reproductive skew, I examined four possible explanations identified from reproductive skew theory. From the literature, I then reviewed evidence to investigate if marmot species differ in: (1) the ability of dominants to control the reproduction of subordinates; (2) the degree of relatedness between group members; (3) the benefit for subordinates of remaining in the social group; and (4) the benefit for dominants of retaining subordinates. I found that the optimal skew hypothesis may apply for both sets of species. I suggest that yellow-bellied marmot females may benefit from retaining subordinate females and in return have to concede them reproduction. On the contrary, monogamous marmot species may gain by suppressing the reproduction of subordinate females to maximise the efficiency of social thermoregulation, even at the risk of departure of subordinate females from the family group. Finally, I discuss scenarios for the simultaneous evolution of sociality, monogamy and reproductive skew in marmots.     

The cost of dominance: suppressing subordinate reproduction affects the reproductive success of dominant female banded mongooses

Social species show considerable variation in the extent to which dominant females suppress subordinate reproduction. Much of this variation may be influenced by the cost of active suppression to dominants, who may be selected to balance the need to maximize the resources available for their own offspring against the costs of interfering with subordinate reproduction. To date, the cost of reproductive suppression has received little attention, despite its potential to influence the outcome of conflict over the distribution of reproduction in social species. Here, we investigate possible costs of reproductive suppression in banded mongooses, where dominant females evict subordinates from their groups, thereby inducing subordinate abortion. We show that evicting subordinate females is associated with substantial costs to dominant females: pups born to females who evicted subordinates while pregnant were lighter than those born after undisturbed gestations; pups whose dependent period was disrupted by an eviction attained a lower weight at independence; and the proportion of a litter that survived to independence was reduced if there was an eviction during the dependent period. To our knowledge, this is the first empirical study indicating a possible cost to dominants in attempting to suppress subordinate breeding, and we argue that much of the variation in reproductive skew both within and between social species may be influenced by adaptive variation in the effort invested in suppression by dominants.     

Reproductive skew and the threat of eviction: a new perspective

Most recent models of the partitioning of reproduction attempt to explain patterns of skew on the assumption that dominant individuals have complete control over breeding opportunities within the group, but may nevertheless concede a share of direct reproduction to subordinates as an incentive to remain peacefully in the association. Although these models may be applicable to some animal societies, we argue that they fail to provide a comprehensive theory of skew. Instead, we suggest that subordinates may often be able to claim unsanctioned reproduction for themselves, but will be forced to exercise a degree of reproductive restraint lest they incite ejection by the dominant. Reproductive skew, in other words, may reflect the threat of ejection (inducing subordinate restraint) rather than the threat of subordinate departure (inducing reproductive concessions by dominants). We present a simple ESS model of reproductive skew under these circumstances, which demonstrates that a shift in emphasis from reproductive concessions by dominants to reproductive restraint on the part of subordinates, radically alters the predictions of skew models. High group productivity, high relatedness and (when group members are related) strong ecological constraints are all expected to lead to reduced skew (the opposite conclusions to those of previous, concession-based analyses). The reason is that these factors reduce the benefits (or increase the costs) of ejection to the dominant, who therefore does best to tolerate more subordinate reproduction.     

Higher reproductive skew among birds than mammals in cooperatively breeding species

While competition for limited breeding positions is a common feature of group life, species vary widely in the extent to which reproduction is shared among females (‘reproductive skew’). In recent years, there has been considerable debate over the mechanisms that generate variation in reproductive skew, with most evidence suggesting that subordinates breed when dominants are unable to prevent them from doing so. Here, we suggest that viviparity reduces the ability of dominant females to control subordinate reproduction and that, as a result, dominant female birds are more able than their mammal counterparts to prevent subordinates from breeding. Empirical data support this assertion. This perspective may increase our understanding of how cooperative groups form and are stabilized in nature.     

Reproductive skew in female common marmosets: what can proximate mechanisms tell us about ultimate causes?

Common marmosets are cooperatively breeding monkeys that exhibit high reproductive skew: most subordinate females fail to reproduce, while others attempt to breed but produce very few surviving infants. An extensive dataset on the mechanisms limiting reproduction in laboratory-housed and free living subordinate females provides unique insights into the causes of reproductive skew. Non-breeding adult females undergo suppression of ovulation and inhibition of sexual behaviour; however, they receive little or no aggression or mating interference by dominants and do not exhibit behavioural or physiological signs of stress. Breeding subordinate females receive comparable amounts of aggression to non-breeding females but are able to conceive, gestate and lactate normally. In groups containing two breeding females,however, both dominant and subordinate breeders kill one another's infants. These findings suggest that preconception reproductive suppression is not imposed on subordinate females by dominants, at a proximate level, but is instead self-imposed by most subordinates, consistent with restraint models of reproductive skew. In contrast to restraint models, however, this self-suppression probably evolved not in response to the threat of eviction by dominant females but in response to the threat of infanticide. Thus,reproductive skew in this species appears to be generated predominantly by subordinate self-restraint, in a proximate sense, but ultimately by dominant control over subordinates' reproductive attempts.     

Dominance, reproduction and survival in banded mongooses: towards an egalitarian social system?

The banded mongoose, Mungos mungo, is a social species that forms multimale and multifemale family groups. Earlier studies suggest these family groups are relatively egalitarian with small differences in reproductive opportunities among individuals of different rank. In contrast, previous studies of other social mongooses have focused on species with more despotic control of reproduction (meerkats, Suricata suricatta, dwarf mongooses, Helogale parvula). In these species, the distribution of reproductive opportunities amongst individuals of different rank has met the predictions of reproductive skew theory: dominant individuals accrue greater reproductive benefits than subordinates, with subordinates breeding less often than dominants. In this paper we test how well two predictions of reproductive skew theory explain variance in measures of reproductive effort, and its correlates, in a wild population of banded mongooses in Queen Elizabeth National Park, Uganda. We measure dominance rank in males and females, and we investigate whether individuals of higher social rank accrue greater benefits than subordinates in terms of survival and reproduction. Banded mongoose dominance hierarchies showed linearity, but low reproductive skew. Rank was not significantly correlated with age. Furthermore, there were only small effects of dominance rank on nutritional levels, and no effects on reproduction and survival, suggesting that banded mongoose societies are indeed relatively egalitarian. No evidence of reproductive suppression was found and other forms of reproductive control were not observed. However, we do not exclude the possibility of increased reproductive competition in circumstances of higher ecological constraints. These findings show that reproductive skew theory is equally useful in explaining variation in reproduction in societies with low reproductive skew, as it is in explaining the allocation of reproductive effort in despotic social systems. Copyright 2001 The Association for the Study of Animal Behaviour.     

Reproductive sharing among queens in the ant Formica fusca

Reproductive sharing among cobreeders, in which reproductiveshares may vary from equal contribution (low reproductive skew)to reproductive dominance by one individual (high reproductiveskew), is a fundamental feature of animal societies. Recenttheoretical work, the reproductive skew models, has focusedon factors affecting the degree to which reproduction is skewedwithin a society. We used the parameters provided by skew modelsas a guideline to study determinants of reproductive sharingin polygyne ants. As a model system we used two-queen laboratorycolonies of the ant Formica fusca in which the reproductiveshares of each queen was assessed from offspring by using allozymesand DNA microsatellites. We tested how the different variablesincluded in reproductive skew models (queen-queen relatedness,potential fighting ability, productivity, and worker relatednessreflected by queen number in the colony of origin) affect reproductivesharing among queens. The results showed that the relatednessamong queens explained 26% of the variation in reproductiveskew. The size difference between queens (reflecting potentialfighting ability), colony productivity, and worker relatednessdid not have an effect on reproductive partitioning among cobreeders.To our knowledge, this is the first study to test for the effectsof various determinants of skew in an experimental setting.     

Sex ratios and skew models: the special case of evolution of cooperation in polistine wasps

Cooperative breeding often involves reproductive dominance hierarchies. Such hierarchies have been proposed to form and to be maintained through an equitable skew in reproduction for both dominants and subordinates. The general form of skew models also predicts that cooperation can be stable only if cooperation greatly increases group reproductive success or subordinates are greatly constrained in their reproductive prospects relative to dominants. Neither, however, seems to be generally present in the colony initiation phase of temperate polistine wasps, although the behaviors of individuals within such groups are often consistent with skew model predictions. This apparent contradiction can be resolved in the context of a special case of the skew models that incorporate mother-offspring conflicts over sex ratios. Data suggest that all the needed preconditions are present for cooperating foundresses to gain an added benefit through producing male-biased investment ratios. Therefore, the special case model predicts that cooperation can evolve in Hymenoptera with both the observed high skews and reduced per capita group productivity. Further predictions of the special case model (e.g., mixed populations of single and multifoundresses) are also supported. Because the special case model is applicable only to haplodiploids, this may explain why cooperation in vertebrates rarely occurs without significant ecological or physiological constraints. Finally, comparisons to other social Hymenoptera taxa suggest that factors stabilizing cooperation between colony-initiating females may simultaneously constrain the evolution of morphologically specialized worker castes.     

Testing a new version of the size-advantage hypothesis for sex change: sperm competition and size-skew effects in the bucktooth parrotfish, Sparisoma radians

A variety of field studies suggest that sex change in animalsmay be more complicated than originally depicted by the size-advantagehypothesis. A modification of the size-advantage hypothesis,the expected reproductive success threshold model, proposesthat sperm competition and size-fecundity skew can stronglyaffect reproductive pay-offs. Size-fecundity skew occurs ifa large female's fecundity is markedly higher than the aggregateof the other members of her social group and, together withpaternity dilution from sperm competition, can produce situationsin which large females benefit by deferring sex change to smallerfemales. Deferral by large females can create sex-size distributionscharacterized by the presence of large females and small sex-changedmales, and it is precisely these distributions that the traditionalsize-advantage model cannot explain. We tested the predictionsof the new model with the bucktooth parrotfish, Sparisoma radians,on coral reefs in St. Croix, U.S. Virgin Islands. Collectionsand spawning observations determined that the local environmentalregime of S. radians is characterized by pervasive sperm competition(accompanying 30% of spawns) and factors that can produce substantialsize-fecundity skew in social groups. Dominant male removalexperiments demonstrate that the largest females in social groupsoften do not change sex when provided an opportunity. Instead,smaller, lower-ranking females change sex when a harem vacancyarises. This pattern of sex change is in contrast to virtuallyall previous studies of social control of sex change in fishes,but provides strong support for the general predictions of theexpected reproductive success threshold model.     

Cichlids do not adjust reproductive skew to the availability of independent breeding options

Helpers in cooperatively breeding species forego all or partof their reproduction when remaining at home and assisting breedersto raise offspring. Different models of reproductive skew generatealternative predictions about the share of reproduction unrelatedsubordinates will get depending on the degree of ecologicalconstraints. Concession models predict a larger share when independentbreeding options are good, whereas restraint and tug-of-warmodels predict no effects on reproductive skew. We tested thesepredictions by determining the share of reproduction by unrelatedmale and female helpers in the Lake Tanganyika cichlid Neolamprologuspulcher depending on experimentally manipulated possibilitiesfor helper dispersal and independent breeding and dependingon helper size and sex. We created 32 breeding groups in thelaboratory, consisting of two breeders and two helpers each,where only the helpers had access to a nearby dispersal compartmentwith (treatment) or without (control) breeding substrate, usinga repeated measures design. We determined the paternity andmaternity of 1185 offspring from 47 broods using five to nineDNA microsatellite loci and found that: (1) helpers participatedin reproduction equally across the treatments, (2) large malehelpers were significantly more likely to reproduce than smallhelpers, and (3) male helpers engaged in significantly morereproduction than female helpers. Interestingly, in four broods,extragroup helper males had fertilized part of the brood. Nohelper evictions from the group after helper reproduction wereobserved. Our results suggest that tug-of-war models based oncompetition over reproduction within groups describe best thereproductive skew observed in our study system. Female breedersproduced larger clutches in the treatment compared to the controlsituation when the large helpers were males. This suggests thatmale breeder-male helper reproductive conflicts may be alleviatedby females producing larger clutches with helpers around.     

Colony genetic structure in a facultatively eusocial hover wasp

The degree of genetic heterogeneity among the individuals inan animal society depends on the society's genetic structure.Genetic heterogeneity, in turn, means that group members willdiffer in their reproductive objectives and conflicts over reproductionmay arise. The resolution of these conflicts may be reflectedin the way that reproduction is partitioned between potentialreproductives. We used 5 microsatellite loci to investigategenetic structure and reproductive skew in 17 nests of the Malaysianhover wasp, Parischnogaster alternata. Parischnogaster alternatacolonies are small (1–10 females), and all adult colonymembers are capable of mating and producing offspring. We foundthat colonies tended to consist of closely related individualsand that at any one time the production of both female and maleoffspring was nearly always monopolized by a single dominantfemale, despite considerable variation between nests in parameterspredicted to affect skew. Subordinate females that remainedin their natal colonies obtained indirect fitness benefits byhelping to raise offspring to which they were related. Subordinatefemales also appeared to be positioned within an age-based queuefor inheritance of the dominant egg-laying position. We suggestthat the high skew in P. alternata may result from strong ecologicalconstraints on solitary nesting, high relatedness, and a relativelyhigh probability that subordinates will eventually inherit theposition of dominance.     

Starving the competition: a proximate cause of reproductive skew in burying beetles (Nicrophorus vespilloides)

Proximate mechanisms underlying reproductive skew are obscure in many animals that breed communally. Here, we address causes of reproductive skew in brood-parasitic associations of burying beetles (Nicrophorus vespilloides). Male and female burying beetles feed and defend their larvae on buried carcasses. When several females locate the same small carcass, they engage in violent physical altercations. The subordinate then acts as an intraspecific brood parasite, laying eggs, but not providing care. The dominant female largely monopolizes access to the carcass; she alone provides parental care and her share of the brood is much larger than the subordinate's. On larger carcasses, subordinates have greater access to the carcass than on small ones, and reproductive skew is reduced. Differential fecundity, ovicide and larvicide have been suggested as causes of skew on small carcasses. Here, we report the results of the experiments pertaining to the first two of these potential mechanisms. Ovicide did not significantly contribute to reproductive skew on small carcasses, but differential fecundity did. Fecundity differences were due to dominance status, not body size per se. Fecundity differences disappeared when supplemental food was available, suggesting that reduced access to the carcass limits fecundity by causing nutritional deficiencies. Supplemental food prevented such nutritional deficiencies and allowed subordinates to produce as many eggs as dominants. Apparently, aggressive behaviour by dominants functions in the context of reproductive competition, limiting subordinate reproduction by preventing food intake on the carcass.     

Predicting the temporal dynamics of reproductive skew and group membership in communal breeders

Reproductive skew models attempt to predict the fraction ofreproduction contributed by each individual that participatesin a communal brood. One potential limitation of these modelsis that individuals make a single, fixed decision about groupmembership and reproductive allocation at the beginning of thebreeding period. While this is appropriate for animals thatreproduce in a synchronous bout, many cooperative breeders produceoffspring over a prolonged period of time. It is likely thatthese species adjust reproductive allocation and group membershipover time in response to temporal shifts in group productivityand ecological constraints. In this paper we adapt transactionalmodels of reproductive skew to a continuous form, generatingtime-dependent functions of reproductive allocation. We derivea general method for predicting temporal changes in group membershipas well as a general expression for reproductive skew acrossthe regions over which a group is stable. Using a linear approximationfor time-dependent reproduction, we derive new expressions forreproductive skew in cases where the subordinate departs duringthe breeding period. In this case we find that the traditionalmodel always overestimates the subordinate's share of reproductionwhen dominants are in control of both reproductive shares andgroup membership (i.e., concessions models). Conversely, wefind that the traditional model always underestimates the subordinate'sshare of reproduction when subordinates are in control of reproductiveshares (i.e., constraint models). We discuss the implicationsof these new calculations in relation to the traditional skewmodels and more recent empirical tests of reproductive skewin animal societies.     

Failures of reproductive suppression in dwarf mongooses (Helogale parvula): accident or adaptation?

In many communally breeding species, only the dominant femalenormally breeds, while subordinates tolerate reproductive suppression(these are "despotic" species, in the terminology of Brown,1987; Macdonald and Moehlman, 1983; Vehrencamp, 1983). Yet inmany species for which reproductive suppression is the norm(across a wide variety of taxa), subordinates do occasionallybreed. Because reproduction by subordinates is atypical forthese species, it is often regarded as simple failure of thenormal mechanisms of suppression. An alternate hypothesis isthat subordinate pregnancies represent an evolutionary compromisebetween dominant and subordinate, in which dominants concedetheir monopoly on reproduction in order to retain helpers. Weuse data from a long-term study of dwarf mongooses (Helogaleparvula) in Serengeti National Park, Tanzania, to test thishypothesis, using an inclusive fitness model adapted from oneby Vehrencamp (1983). We find that the incidence of subordinatepregnancy closely matches that predicted by the model, suggestingthat the mechanisms that underlie reproductive suppression indwarf mongooses are finely adjusted to the social and demographicenvironment. [Behav Ecol 1991; 2: 7–15]     

What do the indices of reproductive skew measure?

Several indices of reproductive skew that quantify the degree of unequal partitioning of reproductive output among individuals have been proposed without consensus on their merits and defects. We believe that the major reason for the disagreement is the lack of discussion on what the population parameter of skew (population skew or true skew) should measure. In our view, the skew index should be an unbiased estimate of a population skew, and the estimated skew needs to satisfy the following two conditions. First, if the group size is equal and the distribution of potential of reproductive output (pi), which is scaled by the proportion of the individual's to the total group reproductive output, is also fixed, skew remains constant even when the total number of offspring in the group changes. Second, if the group size is different, skew should have intuitive biological meaning. Our analyses revealed that, among various indices so far proposed, only the skews estimated by Kokko and Lindstr?m's lambda and Morisita's I(delta) satisfy the first condition. However, the two indices estimate different population parameters, thus implying different biological meanings. Morisita's I(delta) is a linear function of CV(2) (squared coefficient of variation) of pi, and lambda is a positive function of [Formula: see text] when offspring number follows a multinomial distribution. In the special cases where a group consists of discrete classes of breeders and nonbreeders, lambda behaves roughly inversely parallel to the absolute number of breeders, while I(delta) moves almost parallel to the proportion of nonbreeders. Furthermore, lambda is sensitive to the total proportion of reproductive output possessed by the dominants but is relatively less sensitive to the number of subordinates. We discussed the possible situations where either of the two indices will be useful.     

Concessions of an alpha male? Cooperative defence and shared reproduction in multi-male primate groups

By living in social groups with potential competitors, animals forgo monopolizing access to resources. Consequently, debate continues over how selection might favour sociality among competitors. For example, several models exist to account for the evolution of shared reproduction in groups. The 'concession model' hypothesizes that dominant reproducers benefit from the presence of subordinates, and hence tolerate some reproduction by subordinates. This mutual benefit to both dominants and subordinates may provide a foundation for the formation of social groups in which multiple members reproduce-a necessary step in the evolution of cooperation. To date, however, the concession model has received virtually no support in vertebrates. Instead, the vast majority of vertebrate data support 'limited control models', which posit that dominant reproducers are simply unable to prevent subordinates from reproducing. Here we present the most comprehensive evidence to date in support of the concession model in a vertebrate. We examined natural variation in the number of adult males in gelada (Theropithecus gelada) reproductive units to assess the extent of reproductive skew in multi-male units. Dominant ('leader') males in units that also had subordinate ('follower') males had a 30 per cent longer tenure than leaders in units that did not have followers, mainly because followers actively defended the group against potential immigrants. Follower males also obtained a small amount of reproduction in the unit, which may have functioned as a concession in return for defending the unit. These results suggest that dominants and subordinates may engage in mutually beneficial reproductive transactions, thus favouring male-male tolerance and cooperation.