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Second, we fitted the number of sessions an individual was observed escorting as the binomial response variable with the total number of observation sessions as the denominator, for individuals that had been escorting at least once in that litter. Splitting the analyses in this way also accounted for problems with zero-inflation in the data. Predictor variables in both analyses were rainfall during the previous month, weight and age of the potential escort and their interaction, number of pups in the litter, number of same-sex adults in the group, parentage whether or not the focal individual was parent to any pups in that litter , the focal individual's average relatedness to pups in the litter and the focal individual's average relatedness to adults in the group.

Both analyses included individual, litter and social group as random effects. We then looked at pairwise interactions between pups and escorts within-litter. Similar to analysis 1, this analysis was done in two parts, as different factors may predict which pups an adult associates with, and how much care is given when they do. For this all potential pairs of pups and escorts were constructed, so that all individuals that were observed escorting at least once in a given litter were included as potential escorts for all the pups in that litter.

Predictor variables were sex of the pup, pup weight, parentage and relatedness between the adult and the pup, and to investigate whether escorting might be contingent on within-sex competition, we also included as a predictor the interaction between the number of same-sex adults and the sex of the pup.

Second, among observed pup—escort pairs, we used a binomial GLMM with the proportion of observation sessions the focal pup was being escorted by the focal adult as the binomial response variable, using the same set of predictor variables as above. While most of the relationships are dyadic in nature, an adult can sometimes escort multiple pups in a litter and a pup may have multiple escorts.

To account for this, we included both pup and escort identity, as well as litter and social group, as random effects in these analyses. Finally, we looked at escorting relationships from the pup perspective, with the analysis split as above. Second, we looked at predictors of the amount of care those pups received that had an escort proportion of observation sessions they were escorted with pup weight, pup sex, litter size, sex of the escort, parentage and relatedness between pup and the escort as covariates.

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In cases where the pup had multiple escorts, we included the characteristics of the adult that provided most care. In this analysis, escort identity, litter and social group were included as random effects. Neither relatedness to the litter after controlling for the effect of parentage; see electronic supplementary material nor relatedness to other group members predicted the probability that a female escorted in a given litter relatedness to the litter: Figure 2. Effects of group size and parentage on patterns of escorting in banded mongooses.

For males, neither parentage nor relatedness predicted the probability that a male escorted in a given litter parentage: As in females, the effect of weight on escorting depended on age, with both the likelihood of escorting and escorting effort decreasing with age in heavy but not in light individuals electronic supplementary material, table S1 and figure S1 b. The escorting effort of males was not predicted by rainfall. Parentage did not predict the amount of care escorts gave to a particular pup female escorts: Other factors did not predict dyadic association or the amount of care provided by females, see electronic supplementary material, table S2, for full results.

Figure 3. Sex differences in patterns of care.

Acknowledgements

Numbers are counts of observed escorting relationships. Compared with female pups, male pups received more care from helpers proportion of observation sessions being escorted for all escorted and non-escorted pups: Male pups were also slightly heavier at emergence than females The effect of pup weight on its probability of being escorted depended on sex of the pup: Neither the relatedness between the pup and the escort providing most care, parentage, nor sex of the escort predicted the total amount of care that the escorted pups received relatedness: Much research has focused on the influence of relatedness on selection for helping behaviour [ 47 ], but why individuals might conceal identity or avoid discrimination within groups is a much less explored topic, particularly in vertebrates.


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The naive prediction from Hamilton's rule is that care should be directed at closer relatives, but this ignores the problems that being able to discriminate creates, both at the individual and at the group level. Nepotism can be disruptive to the group and lead to selection against the very recognition cues that form the basis of such discrimination [ 7 , 9 ]. More pressingly, for the recipient to identify themself as highly related to some group members also comes at the cost of revealing themself as less related to others, exposing them to negative discrimination and spite [ 14 , 48 ].

Where the average relatedness between helpers and helpees is high anyway, such costs may override any benefit, making returns from discrimination marginal at best [ 7 , 12 ]. In such systems, individuals may be better off adjusting their behaviour according to other predictors of costs and benefits of cooperation, without the need for possibly costly within-group kin discrimination, and this is indeed what we find in the banded mongoose. Earlier studies have suggested that pups have an active role in establishing relationships with particular escorts [ 26 , 27 , 49 ], implying that the escort—pup relationship is not solely the result of choices made by the helper.

However, previously, we have been unable to exclude the possibility that escorts were typically the parents of the pups they care for, bringing into question whether adults that engage in this behaviour should be termed helpers at all [ 50 ]. This study shows that escorts do indeed care for pups that are not their own offspring, and that, despite the presence of high-relatedness offspring within the communal litter, neither males nor females preferentially form pairwise associations with pups that are more related to them.

Although females are more likely to escort when the current litter contains some of their own young, they do not preferentially pair with their own offspring, supporting previous claims that mothers do not or cannot discriminate their own young in the communal litter [ 21 , 38 , 51 ].

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Moreover, neither pairwise relatedness nor parentage predicts the amount of care females allocated to an individual pup. The lack of kin discrimination by females is perhaps surprising given that escorting boosts the survival and growth of pups [ 26 , 28 ]. However, in banded mongooses, the potential costs of nepotistic discrimination may be particularly high because within-group infanticide is common [ 52 ].

Any pup advertising its close relatedness to a particular female or, potentially, male could be targeted by others and could also lose out on allosuckling by other females, even if not directly aggressed [ 53 ]. In males, neither paternity nor relatedness to the pups predicted patterns of assortment in escort—pup relationships.

Nor did relatedness predict male escorting effort across litters. However, we did find two correlations between relatedness and patterns of male helping. First, across litters, males increased the time spent escorting when they were more closely related on average to the rest of the group. Second, within pup—escort pairs, more related dyads spent more time together. These results might suggest kin discrimination by males. However, these patterns could also arise as a result of other factors that are correlated with relatedness.

For example, there may be subtle similarities in genetically heritable foraging preferences or character traits, such as preference of closed versus open habitat, or boldness and shyness, that could explain why more related partners spend more time together. There may also be subtle effects of group size on the observed relationships between escorts and pups.

In small groups, in which relatedness is high, pups are particularly valuable in terms of group recruitment, and all adults may be more attentive escorts. Without cross-fostering experiments to manipulate which pups pair with which escorts, or experimental manipulation of group size, we are currently unable to fully understand the causality of the relationship between relatedness and helping effort in males. We did find strong discrimination based on sex of the recipient. Both males and females were more likely to pair with a pup of their own sex and reduced their overall helping effort in response to increasing number of same-sex adults in the group.

As group size was highly correlated with numbers of both adult males and females, individuals may simply reduce their contribution to care as there are more helpers present. However, females also provided more care to female pups when adult female numbers were low, which implies that within-sex cooperation and competition may be driving the preferential direction of help to the same sex. For female banded mongooses, there appears to be an optimal group size that maximizes their reproductive success [ 52 ]. Females are evicted in same-sex cohorts when the number of breeding females grows large [ 53 — 55 ], and patterns of dispersal and eviction may therefore create incentives for female adults to adjust care given towards female pups depending on the competitive environment.

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Males may also have an incentive to target care towards other males, since males may be particularly important in defending the territory against neighbouring groups and evicted cohorts of males that attempt to take over and supplant existing males [ 29 ]. Sex bias in care has been observed in many biparental birds, as well as other cooperatively breeding mammals, with varying direction of bias and consequences for the offspring.

For example, in the toc-toc Foudia sechellarum , the brood is divided by sex post-fledging between the mother and the father [ 56 ] with no overall differences between the sexes in the amount of care. In zebra finches, mothers preferentially provision sons over daughters, while fathers show no bias, and sons receive more food than daughters overall [ 57 ]. For example, in the cooperatively breeding arabian babblers Turdoides squamiceps , helpers invest in offspring of the opposite sex in order to avoid competition [ 58 ], as do spotted hyaenas Crocuta crocuta , where males associate more with daughters, than with sons [ 59 ].

Preferential helping of the same sex has been previously observed in the cooperatively breeding meerkat Suricatta suricata. Similar to the banded mongoose, meerkat female helpers preferentially feed female pups, but males show no bias [ 60 ]. Females also provide more help than males. These patterns of care may be explained by sex differences in dispersal and the benefits of philopatry.

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In meerkats, males are the dispersing sex, and hence benefit less from any group augmentation benefits of helping compared with females. This explanation fits with our findings in the banded mongoose, where both sexes remain in their natal group, and are also more likely to pair up with a pup of the same sex.

Another explanation for the sex bias in caring relationships observed here is that the competitive ability of the pups may be driving the association. Male banded mongooses are more likely to be escorts than females, and they also provide more care. Larger pups were more likely to be escorted and received more care, despite the caring effort of individual helpers not being correlated with pup size.

As male pups were on average slightly larger than female pups, they also received more care overall, with the total amount of escorting care received increasing more steeply with size in female than in male pups. This result suggests that bigger pups may be able to secure the best helpers, which often are young males.