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Animal groups are heterogeneous assemblages of individuals with differing fitness interests, which may lead to internal conflict over investment in group territorial defence. Differences between individuals may lead to different behavioural responses to intergroup conflict, particularly between the sexes. These potential impacts have been little studied. We used social network analysis to investigate the impact of simulated intergroup conflicts on social relationships in groups of wild banded mongooses Mungos mungo, in which intergroup fights are more costly for males than females. We predicted that social cohesion (specifically male-to-male and female-to-male grooming) would increase after conflict, and aggression would decrease, to minimize conflict between the sexes. Simulated intergroup conflicts were performed by exposing banded mongoose groups to scents, 'war cry' playbacks, and live intruders from a rival group. All grooming and aggression interactions between individuals were recorded, and grooming and aggression social networks were created for the 2 days preceding a simulated intergroup conflict (pre-conflict network) and the 2 days after (post-conflict network). We found no evidence of an increase in social cohesion after simulated conflicts, measured as grooming eigenvector centrality. Male-to-male, male-to-female and female-to-male grooming strength decreased after simulated intrusions compared to female-to-female grooming strength. However, male-female aggression decreased in intrusion trials compared to other interaction types, consistent with the hypothesis that intergroup encounters reduce the level of intragroup conflict between males and females. Males were more affected socially by intergroup encounters than females, which may be because they are investing in defence rather than internal relationships. Focusing on individual relationship changes, using social network analysis, can reveal changes in the directionality of behaviour in response to intergroup encounters, and highlight how individual responses to conflict may scale up to affect social networks and, potentially, group performance. This study highlights the importance of studying both group-level behaviours and individual relationships to more fully understand responses to intergroup encounters.

Behavioral mechanisms for avoiding inbreeding are common in the natural world and are believed to have evolved as a response to the negative consequences of inbreeding. However, despite a fundamental role in fitness, we have a limited understanding of the cues that individuals use to assess inbreeding risk, as well as the extent to which individual inbreeding behavior is repeatable. We used piecewise structural equation modeling of 24 years of data to investigate the causes and consequences of within- versus extra-group paternity in banded mongooses. This cooperatively breeding mammal lives in tight-knit social groups that often contain closely related opposite-sex breeders, so inbreeding can be avoided through extra-group mating. We used molecular parentage assignments to show that, despite extra-group paternity resulting in outbred offspring, within-group inbreeding occurs frequently, with around 16% litters being moderately or highly inbred. Additionally, extra-group paternity appears to be plastic, with females mating outside of their social group according to individual proxies (age and immigration status) and societal proxies (group size and age) of within-group inbreeding risk but not in direct response to levels of within-group relatedness. While individual repeatability in extra-group paternity was relatively low, female cobreeders showed high repeatability, suggesting a strong constraint arising from the opportunities for extra-group mating. The use of extra-group paternity as an inbreeding avoidance strategy is, therefore, limited by high costs, opportunity constraints, and the limited reliability of proxies of inbreeding risk.

Individual foraging specialisation has important ecological implications, but its causes in group‐living species are unclear. One of the major consequences of group living is increased intragroup competition for resources. Foraging theory predicts that with increased competition, individuals should add new prey items to their diet, widening their foraging niche (‘optimal foraging hypothesis’). However, classic competition theory suggests the opposite: that increased competition leads to niche partitioning and greater individual foraging specialisation (‘niche partitioning hypothesis’). We tested these opposing predictions in wild, group‐living banded mongooses (Mungos mungo), using stable isotope analysis of banded mongoose whiskers to quantify individual and group foraging niche. Individual foraging niche size declined with increasing group size, despite all groups having a similar overall niche size. Our findings support the prediction that competition promotes niche partitioning within social groups and suggest that individual foraging specialisation may play an important role in the formation of stable social groupings.

In animal societies, conflict within groups can result in eviction, where individuals are often permanently expelled from their group. To understand the evolution of eviction and its role in the resolution of within-group conflict requires information on the demographic consequences of eviction for individuals and groups. However, such information is usually difficult to obtain because of the difficulty in tracking and monitoring individuals after they are evicted from their natal groups. Here we used a 15-year data set on life history and demography to investigate the consequences of eviction in a tractable cooperatively breeding mammal, the banded mongoose, Mungos mungo . In this species, groups of individuals are periodically evicted en masse and eviction is a primary mechanism by which new groups form in the study population. Following eviction, we found sex differences in dispersal distance: some females established new groups on the study peninsula but males always dispersed away from the study peninsula. Evicted females suffered reduced reproductive success in the year after eviction. For the evicting group, eviction was associated with increased per capita reproductive success for females, suggesting that eviction is successful in reducing reproductive competition. However, eviction was also associated with increased intergroup conflict for the evicting group. Our results suggest that within-group conflict resolution strategies affect group productivity, group interactions and the structure of the population, and hence have fitness impacts that reach beyond the individual evictors and evictees involved in eviction.

Hette‐Tronquart (2019, Ecol. Lett.) raises three concerns about our interpretation of stable isotope data in Sheppard et al. (2018, Ecol. Lett., 21, 665). We feel that these concerns are based on comparisons that are unreasonable or ignore the ecological context from which the data were collected. Stable isotope ratios provide a quantitative indication of, rather than being exactly equivalent to, trophic niche.

When breeding females compete for limited resources, the intensity of this reproductive conflict can determine whether the fitness benefits of current reproductive effort exceed the potential costs to survival and future fertility. In group-living species, reproductive competition can occur through post-natal competition among the offspring of co-breeding females. Spontaneous abortion could be a response to such competition, allowing females to curtail reproductive expenditure on offspring that are unlikely to survive and to conserve resources for future breeding opportunities. We tested this hypothesis using long-term data on banded mongooses, Mungos mungo , in which multiple females within a group give birth synchronously to a communal litter that is cared for by other group members. As predicted, abortions were more likely during dry periods when food is scarce, and in breeding attempts with more intense reproductive competition. Within breeding events, younger, lighter females carrying smaller fetuses were more likely to abort, particularly those that were also of lower rank. Our results suggest that abortion may be a means by which disadvantaged females conserve resources for future breeding attempts in more benign conditions, and highlight that female reproductive competition may be resolved long before the production of offspring.

Among mammals, scent has long been known to encode oestrus; however, in many species, detecting pregnancy may also be important in terms of both competition and mate-choice. Here, we show, through odour presentation experiments, that pregnancy is discernible via scent by both sexes in the cooperatively breeding banded mongoose, Mungos mungo . Males spent more time investigating and were more likely to scent mark the odours of non-pregnant females, compared to pregnant females. Females showed increased levels of scent marking when odours were of the same reproductive state as themselves. These results present the first direct demonstration that pregnancy is detectable via scent in wild cooperative breeders. Detecting pregnancy may be particularly important in cooperative breeders as, in addition to the competition between males for receptive mates, there is also intense competition between females for access to alloparental care. Consequently, dominant females benefit from targeting reproductive suppression towards subordinates that represent direct threats, such as pregnant females.

Individual differences in contributions to cooperation can be strikingly consistent over time giving rise to alternative cooperative phenotypes within animal societies. Following the social niche specialization hypothesis, these consistent differences may be driven by social conflict over reproductive opportunities if individuals specializing as ‘breeders’ and ‘nonbreeders’ experience a beneficial reduction in social conflict and differential costs associated with cooperating. This hypothesis gives three testable predictions: (1) consistent individual differences in cooperative behaviour will be accompanied by consistent individual differences in reproductive behaviour, (2) individuals contributing heavily to reproduction will contribute relatively little to cooperative behaviours within the same breeding attempt and vice versa, and (3) individuals that consistently contribute heavily to reproduction over their lifetime will also consistently contribute less to cooperative behaviours and vice versa. We tested these predictions with a 15-year investigation into the lifetime patterns of mate guarding and two forms of cooperative offspring care (‘babysitting’ and ‘escorting’) in a wild population of banded mongooses. We found significant repeatability of individual contributions to both cooperative behaviours, as well as significant repeatability of individual levels of mate guarding. However, we found no evidence of negative covariance between contributions to cooperative and reproductive behaviours either within breeding attempts or across lifetimes. This suggests that the observed consistent individual differences in both cooperative behaviour and reproduction are not associated; there is no evidence of a trade-off between reproduction and cooperation. However, we found a significant positive covariance between babysitting and escorting when estimated both within breeding attempts and across lifetimes, which suggests that some group members are generally more cooperative than others, contributing more to both behaviours over their lifetimes. The drivers of this consistent individual variation in contributions to cooperation remain unknown.

Preference for uninfected mates is presumed beneficial as it minimizes one’s risk of contracting an infection and infecting one’s offspring. In avian systems, visual ornaments are often used to indicate parasite burdens and facilitate mate choice. However, in mammals, olfactory cues have been proposed to act as a mechanism allowing potential mates to be discriminated by infection status. The effect of infection upon mammalian mate choice is mainly studied in captive rodents where experimental trials support preference for the odors of uninfected mates and some data suggest scent marking is reduced in individuals with high infection burdens. Nevertheless, whether such effects occur in nonmodel and wild systems remains poorly understood. Here, we investigate the interplay between parasite load (estimated using fecal egg counts) and scent marking behavior in a wild population of banded mongooses Mungos mungo. Focusing on a costly protozoan parasite of the genus Isospora and the nematode worm Toxocara, we first show that banded mongooses that engage in frequent, intensive scent marking have lower Isospora loads, suggesting marking behavior may be an indicator trait regarding infection status. We then use odor presentations to demonstrate that banded mongooses mark less in response to odors of opposite sexed individuals with high Isospora and Toxocara loads. As both of these parasites are known to have detrimental effects upon the health of preweaned young in other species, they would appear key targets to avoid during mate choice. Results provide support for scent as an important ornament and mechanism for advertising parasitic infection within wild mammals.

Inbreeding, reproduction between relatives, often impinges on the health and survival of resulting offspring. Such inbreeding depression may manifest itself through immunological costs as inbred individuals suffer increased propensity to disease, infection and parasites compared to outbred conspecifics. Here, we assess how the intestinal parasite loads of wild banded mongooses (Mungos mungo) vary with pedigree inbreeding coefficient (f) and standardized multi-locus heterozygosity. We find a significant association between increased heterozygosity and lower parasite loads; however, this correlation does not stand when considering f. Such findings may be explained by local genetic effects, linkage between genetic markers and genes influencing parasite burdens. Indeed, we find heterozygosity at certain loci to correlate with parasite load. Although these tentative local effects are lost following multiple test correction, they warrant future investigation to determine their strength and impact. We also suggest frequent inbreeding within banded mongooses may mean heterozygosity is a better predictor of inbreeding than pedigree f. This is because inbreeding facilitates linkage disequilibrium, increasing the chances of neutral markers representing genome-wide heterozygosity. Finally, neither f nor heterozygosity had a significant influence on the loads of two specific gastrointestinal parasites. Nevertheless, more heterozygous individuals benefited from reduced overall parasitic infection and genetic diversity appears to explain some variation in parasite burdens in the banded mongoose. © 2016 The Zoological Society of London.

The ability to individually recognize conspecifics is acknowledged as one of the prerequisites for the development of sophisticated social relationships in group-living species. It has been hypothesized that the discrimination of individual identities is crucial for the maintenance of social relationships and cooperation based on repeated interactions, and for the evolution of many social behaviours. Previous studies have shown that the close calls of the cooperatively breeding banded mongoose Mungos mungo are individually distinct. For instance, banded mongoose pups are able to distinguish between close calls of their escort and of a non-escort. In this study, we used playbacks based on the recently proposed violation-of-expectation paradigm and a dominance/age class recognition setup to investigate whether adult banded mongooses use the individual signature of close calls to distinguish among adult group members. We found no evidence that the individual signature in close calls is used to discriminate identity in banded mongooses. Based on the previous work, we suggest that this is not because banded mongooses are incapable of using signatures as a means of individual discrimination, but because the benefits of such discrimination are low. The study highlights the importance of understanding the function of a signal (e.g. the expected response), timing and the biology of the species when designing and performing playback experiments.

In cooperatively breeding vertebrate societies, contributions to offspring care can vary greatly between group members. Kin selection theory predicts that cooperation will be favoured when directed towards relatives and when the cost to benefit ratio is low. The fitness costs of helping in turn depend on the impact of energetic investments in care on future reproductive success, which is likely to vary between helpers. For example, investments may impact more on a young helper, which needs to invest energy in growth and is an inexperienced forager. We investigated how a key predictor of cost, food availability (estimated using rainfall), influences helping behaviour in the banded mongoose, Mungos mungo. In this cooperative carnivore, a variable number of group members breed while almost all help to rear the communal litter. Nonbreeding females and juvenile males helped less when food was scarce, reflecting the potentially high costs of weight loss and reduced growth on survival and future reproductive success. In contrast, adult males maintained their investment in care as food supply decreased, probably because body condition has relatively little impact on male reproductive success in this species. Breeding females (with pups in the communal litter) also maintained their helping effort as food supply decreased. Although mothers invested highly in care, there was no evidence that they preferentially cared for their own pups, probably because synchronized birthing scrambles maternity cues. Patterns of care in the banded mongoose thus seem to reflect the benefits gained from helping and the long-term fitness costs to the helper.

Overmarking occurs when one individual places its scent mark directly on top of the scent mark of another individual. Although it is almost ubiquitous among terrestrial mammals, we know little about the function of overmarking and detailed field observations are rare. We investigated the chemical composition of scents and patterns of overmarking by wild banded mongooses, Mungos mungo. Chemical analyses of anal gland secretions showed that scents were sexually dimorphic. Both male and female adults were more likely to overmark the scents of same-sex individuals. An analysis of responses to two scents on the same site suggested that the sex of the top or most recent scent was more important than that of the bottom or original scent in determining overmarking response. Juveniles also overmarked scents at high rates, but did not respond to scents in a sex-specific way. Same-sex-specific patterns within groups have not been described in any other species, and may reflect a social system with intense intrasexual competition for reproduction within both sexes. Banded mongooses live in large mixed-sex groups, with intense competition between males for females, owing to the heavily male-biased adult sex ratio and highly synchronized oestrous cycles. Oestrous synchronization may also promote intrasexual competition for males within females, as females compete simultaneously for high-quality males. Female competition for males may also be enhanced by the rewards of male-biased parental care. This investigation highlights the need for detailed studies of overmarking in the natural context, to confirm and expand upon laboratory findings.

In contrast to numerous studies of scent marking in male mammals, studies of female scent marking are relatively rare. We have previously shown that communally breeding female banded mongooses, Mungos mungo, are more likely to overmark the scent of other females. Here we describe female overmarking patterns in more detail, and discuss these results in relation to hypotheses potentially explaining such ‘female intrasexual overmarking’. To our knowledge, this is the first study to investigate female overmarking in any wild mammal. First, although we found some evidence of individually distinctive scent marks in females, we found no evidence to suggest that female intrasexual overmarking was related to competition for food, as feeding competition was infrequent, and unrelated to overmarking scores. We also found no evidence to suggest that intrasexual overmarking in females was involved in reproductive suppression. Females with the highest and lowest overmarking scores in each group were mate-guarded by males for similar durations. Finally, we found little evidence to suggest that female intrasexual overmarking was involved in competition for males. Although the female with the highest overmarking score in each group tended to be mate-guarded by males in better condition than the female with the lowest overmarking score, a female’s overmarking score affected neither the amount of harassment she received from males nor the frequency of mating attempts received. These results are discussed in light of these and other untested hypotheses for female overmarking.

Of key importance for understanding cooperative societies is the way in which reproductive opportunities are distributed among group members. Traditionally, skew has been thought of as a product of intrasexual competition. However, cooperatively breeding species often live in mixed-sex groups, so the behaviour of one sex has the potential to influence skew in the other. We addressed the importance of inter- and intrasexual conflict in determining reproductive skew through a study of paternity sharing in the cooperatively breeding banded mongoose, Mungus mungo. Unlike banded mongoose females, where reproductive skew is low, males exhibited high skew, with 85% of paternities being assigned to the three oldest males in each group. Individual males appeared unable to monopolize reproduction because females come into oestrus in synchrony and mate multiply. Instead, older males increased their success by mate guarding the oldest, most fecund females. Our findings therefore highlight the importance of mate choice in males and reveal the behavioural differences between the sexes that generate reproductive skew. They also emphasize the considerable influence that female behaviour can have on male reproductive skew.

Scent marking is commonly described as a territorial behaviour, and scent marks might deter potential intruders from entering occupied areas. Conspecific neighbours present both a reproductive and a territorial threat, thus, determining which, if any, of these threats shapes scent-marking behaviour is difficult. Banded mongooses Mungos mungo provide a rare clear separation between reproductive rivals (found within groups) and territorial rivals (neighbouring groups), because immigration into social groups is extremely rare, and mating occurs almost exclusively within groups. This situation offers an opportunity to assess the relative importance of territorial defence and intra-group competition for mates in shaping scent-marking behaviour. We combined detailed behavioural observations of scent marking, chemical analyses of scent composition and discrimination experiments in the field, and found little evidence for higher rates of scent marking in overlapping areas, a lack of group specificity of scents and a failure of individuals to discriminate between the scents of different groups. Although scent may fulfill some role in territorial demarcation and defence, these results suggest that scent marks and scent-marking patterns are also involved in communicating within social groups.

Lay Summary Female-banded mongooses risk their lives to mate with rivals during pack “warfare.” Data from wild banded mongooses reveal that 18% of pups are fathered by males from rival packs. These pups are less likely to be inbred are heavier and have higher survival chances than their within-pack counterparts. However, their mothers risk a lot to mate with extra-pack males; aggressive encounters between packs account for 20% of pup deaths and 12% of adult deaths., Females of many animal species seek mating opportunities with multiple males, despite being able to obtain sufficient sperm to father their offspring from a single male. In animals that live in stable social groups, females often choose to mate outside their group resulting in extra-group paternity (EGP). One reason proposed to explain female choice for extra-group males is to obtain compatible genes, for example, in order to avoid inbreeding depression in offspring. The benefits of such extra-group paternities could be substantial if they result in fitter, outbred offspring. However, avoiding inbreeding in this way could be costly for females, for example, through retaliation by cuckolded males or through receiving aggression while prospecting for extra-group mating opportunities. We investigate the costs and benefits of EGP in the banded mongoose Mungos mungo, a cooperatively breeding mammal in which within-group mates are sometimes close relatives. We find that pups born to females that mate with extra-group males are more genetically heterozygous are heavier and are more likely to survive to independence than pups born to females that mate within their group. However, extra-group matings also involve substantial costs as they occur during violent encounters that sometimes result in injury and death. This appears to lead femalebanded mongooses to adaptively adjust EGP levels according to the current risk of inbreeding associated with mating within the group. For group-living animals, the costs of intergroup interactions may help to explain variation in both inbreeding rates and EGP within and between species.

In most co-operative breeding species, some individuals contribute much more to helping behaviour than others. The most well-established explanation of such variation is based on kin selection and suggests that, in the absence of detectable differences in relatedness, individuals who suffer lower costs for a given level of help should contribute more. Differences in helping effort between dominance/sex categories were investigated in co-operatively breeding banded mongooses Mungos mungo in Uganda. The most conspicuous form of help in this species is provided by individuals who babysit offspring at the den while the rest of the pack goes off to forage. Across eight groups, the survival rate of pups increased with the average number of babysitters guarding them, consistent with the hypothesis that helpers benefit the brood that they guard. There was no difference between dominant males, subordinate males and breeding females in total contributions to babysitting. Subordinate males, however, contributed more to babysitting in the mornings, which were the longest and presumably the most energetically expensive sessions of the day. In six litters in one well-studied pack, dominant males and breeding females reduced their contribution to babysitting for the period that females were in oestrus. By contrast, subordinate males increased their contribution to become the main babysitters during this time. These results are consistent with the hypothesis that, where helping conflicts with breeding, individuals with little chance of direct reproduction can help at a lower fitness cost than those with a high probability of successful reproduction.

The mode by which individuals disperse, and the cost of dispersal, are of great importance in attempts to understand variation in reproductive skew in animal societies. In this paper we report detailed information on dispersal and pack formation in banded mongooses Mungos mungo. Six pack fission events were recorded among 11 packs over 22 months. Pack fission occurred under two distinct circumstances. First, groups of individuals were evicted from their natal group as a result of intense aggression from other group members. A small fraction of group members was responsible for most of the aggression. Both sexes helped to attack and evict individuals from the group, and both males and females were driven out of their natal groups en masse. The second mode of pack fission occurred when groups of same-sex individuals left their natal group voluntarily to join dispersing individuals of the opposite sex, thereby forming new packs. Dispersing groups were more frequently involved in fights with rival packs of mongooses compared to established groups, and in one instance these fights seemed to be responsible for severe injury and increased mortality among members of a dispersing group. The observations of eviction provide one line of evidence that the presence of subordinates is sometimes detrimental to dominants, contrary to the assumptions of concession models of reproductive skew.

Recent theoretical work suggests that the distribution of reproduction, or degree of reproductive skew, in animal societies depends crucially on (1) whether dominant individuals can fully control subordinate reproduction, and (2) how subordinate reproduction affects the fitness of dominants. I investigated these two factors in cooperatively breeding banded mongooses, Mungos mungo. Female packmates entered oestrus together and were closely guarded by dominant males. These males were aggressive to subordinate males who attempted to mate, but females still managed to mate with males other than their mate guard. Older females were guarded and mated a few days before their younger packmates, yet all females usually gave birth on the same day, suggesting that older females may have a longer gestation period. Moreover, older females carried more fetuses. Overall, ca. 83% of adult females conceived in each breeding attempt and 71% carried to term. These results indicate that, among males, dominant individuals did not have full control over the mating attempts of subordinates (since they could not fully control the mating behaviour of the females they guarded), while among females there was little or no attempt to prevent subordinates from breeding (at least, prior to parturition). Two within-group infanticides by males suggested that some control over reproduction may be exercised postpartum. Per capita survivorship of young in the den increased with the number of females who gave birth. Thus, dominant females may benefit from subordinate reproduction, providing a possible explanation for the lack of reproductive suppression among females in this species. Copyright 2000 The Association for the Study of Animal Behaviour.

Long-term studies of cooperatively breeding vertebrates offer excellent opportunities to test theories about the evolution of cooperation and the demographic consequences of social behavior. Here we draw together over a decade of research on an unusually tractable cooperative mammal system, the banded mongoose (Mungos mungo) and compare our results against advances in social evolution theory that have occurred over the same period. We report recent data on the demographic and genetic structure of the population, and then focus on the main conflictual and cooperative features of the breeding system. Groups are founded by unrelated dispersal coalitions of males and females and consist of multiple male and female breeders. Genetic relatedness between breeding males and females increases with the number of years since group founding, but breeders nevertheless appear to avoid inbreeding. Reproductive competition between females is intense, but young females can escape infanticide by synchronizing birth to the same day as older, socially dominant females. Dominant females respond to reproductive competition by evicting subordinate females en masse. Helping behavior takes two main forms: “babysitting” offspring at the den in the early weeks of life, and “escorting” particular offspring after they emerge from the den. Males contribute most to both babysitting and escorting, particularly the low-ranking males that are excluded from breeding. The way that conflict over reproduction is resolved in this system has a strong influence on patterns of eviction and dispersal. Like many other cooperative vertebrates, each banded mongoose group represents a small, highly viscous population embedded within a larger “metapopulation”. Our research highlights the links between within-group conflict, demography, and the evolution of cooperative life histories.

Social animal species show considerable variation in the way in which reproduction is distributed among group members. Recent attempts to explain this variation have proposed that differences in reproductive skew are attributable to differences in the net benefits group members receive from suppressing each other’s breeding attempts. Despite receiving relatively little theoretical and empirical attention, the availability of resources required for successful breeding may have an important influence in determining the costs and benefits of suppressing reproduction and thus influence reproductive skew within social groups. Here, we test this possibility using a long-term study of female reproductive success in the banded mongoose Mungos mungo .W e find that females experience greater costs of co-breeding when resources are in short supply and that older, more dominant females respond to this cost by suppressing subordinate breeding. This results in differing patterns of reproductive success for females of different competitive abilities, with the oldest, most dominant females breeding regardless of resource availability and younger, subordinate females breeding only when resources are abundant. Our findings highlight the role of resource limitation in determining the distribution of reproductive opportunities within social groups. Key words: cooperative breeding, dominance, rainfall, reproductive conflict, resource availability, suppression. [Behav Ecol]

Reproductive events in animal societies often show a high degree of temporal clustering, but the evolutionary causes of this synchronization are poorly understood. Here, we suggest that selection to avoid the negative effects of competition with other females has given rise to a remarkable degree of birth synchrony in the communal-breeding banded mongoose ( Mungos mungo ). Within banded mongoose groups, births are highly synchronous, with 64 per cent of females giving birth on exactly the same night. Our results indicate that this extreme synchrony arises because offspring suffer an increased risk of infanticide if their mother gives birth before other females, but suffer in competition with older littermates if their mother gives birth after them. These findings highlight the important influence that reproductive competition can have for the evolution of reproductive synchrony.

Considerable research has focused on understanding variation in reproductive skew in cooperative animal societies, but the pace of theoretical development has far outstripped empirical testing of the models. One major class of model suggests that dominant individuals can use the threat of eviction to deter subordinate reproduction (the ‘restraint’ model), but this idea remains untested. Here, we use long-term behavioural and genetic data to test the assumptions of the restraint model in banded mongooses (Mungos mungo), a species in which subordinates breed regularly and evictions are common. We found that dominant females suffer reproductive costs when subordinates breed, and respond to these costs by evicting breeding subordinates from the groupen masse, in agreement with the assumptions of the model. We found no evidence, however, that subordinate females exercise reproductive restraint to avoid being evicted in the first place. This means that the pattern of reproduction is not the result of a reproductive ‘transaction’ to avert the threat of eviction. We present a simple game theoretical analysis that suggests that eviction threats may often be ineffective to induce pre-emptive restraint among multiple subordinates and predicts that threats of eviction (or departure) will be much more effective in dyadic relationships and linear hierarchies. Transactional models may be more applicable to these systems. Greater focus on testing the assumptions rather than predictions of skew models can lead to a better understanding of how animals control each other's reproduction, and the extent to which behaviour is shaped by overt acts versus hidden threats.