Low-ranking Mothers Research Articles

Social risk to infant: The role of kin for maternal visual monitoring in Tibetan macaques.

Maternal monitoring of conspecifics is a crucial anti-predator strategy that also protects infants against risks within the social group. This study examines how maternal characteristics, infant characteristics, mother-infant distance, and the social environment affect maternal monitoring behaviors in free-ranging Tibetan macaques (Macaca thibetana). We observed 12 females with infants and analyzed their visual monitoring patterns. Our findings indicate that maternal rank significantly influences the time allocated to maternal visual monitoring, higher-ranking mothers spending less time than lower-ranking mothers. Maternal experience also played a role in monitoring strategies. Differences in monitoring strategies were observed based on maternal experience: first-time mothers (primiparity) engaged in longer but less frequent monitoring sessions compared to experienced mothers (multiparity). The time and frequency of maternal monitoring decreased as infants aged, and mothers with male infants showed higher levels of monitoring than those with female infants. The distance between mother and infant also affected visual monitoring behavior, with mothers increasing their monitoring levels when infants were nearby (1-5 m), rather than within reach (0-1 m) or beyond nearby (>5 m). Additionally, the presence of kin and non-kin influenced monitoring: as the number of nearby kin increased, monitoring levels decreased, while the presence of more non-kin males led to an increase in monitoring time, and higher-ranking non-kin neighbors increased the frequency of monitoring. These results suggest that Tibetan macaque mothers can adapt their visual monitoring to the social risks faced by their infants, adjusting their strategies to their status and the needs of their offspring.

Mothers' Social Status and Children's Health: Evidence From Joint Households in Rural India.

The premise that a woman's social status has intergenerational effects on her children's health has featured prominently in population science research and in development policy. This study focuses on an important case in which social hierarchy has such an effect. In joint patrilocal households in rural India, women married to the younger brother are assigned lower social rank than women married to the older brother in the same household. Almost 8% of rural Indian children under 5 years old-more than 6 million children-live in such households. We show that children of lower-ranking mothers are less likely to survive and have worse health outcomes, reflected in higher neonatal mortality and shorter height, compared with children of higher-ranking mothers in the same household. That the variation in mothers' social status that we study is not subject to reporting bias is an advantage relative to studies using self-reported measures. We present evidence that one mechanism for this effect is maternal nutrition: although they are not shorter, lower-ranking mothers weigh less than higher-ranking mothers. These results suggest that programs that merely make transfers to households without attention to intrahousehold distribution may not improve child outcomes.

Stable isotopes reveal the effects of maternal rank and infant age on weaning dynamics in wild chacma baboons

Maternal strategies reflect the trade-off between offspring needs and maternal ability to invest, a concept described by the evolutionary theory of parent–offspring conflict. In mammals this conflict has often been investigated by studying weaning, the transition from maternal milk consumption to dietary independence. An investigation of individual variation in weaning can provide information on the adaptive significance of maternal strategies in relation to social and biological variables. We analysed stable nitrogen isotopes of hair samples collected from 22 mother–infant dyads in a wild population of chacma baboons, Papio ursinus, in conjunction with behavioural data on suckling, to explore the temporal dynamics of weaning, as well as the extent and determinants of individual variation in these dynamics. The weaning pattern suggested by isotope values and behavioural data were congruent. The difference between infant and mother stable nitrogen isotope values decreased faster with age in infants of low-ranking mothers, which suggests a faster progression towards weaning, perhaps due to subordinate females experiencing lower resource availability and so being less able to bear the costs of lactation over prolonged periods. Additionally, within-infant variation in stable nitrogen isotope values showed an increase with age (which was not detectable between infants), potentially highlighting the nutritional costs that weaning imposes on offspring. Our combination of isotope analysis and behavioural data from a wild population provides insight into the evolution of maternal strategies. In particular, it suggests that the quantity of care a mother can provide is affected by her rank, with subordinate females possibly not able to lactate for as long and perhaps benefiting from weaning earlier.

Infant adoptions in wild bonnet macaques (Macaca radiata)

Though uncommon, adoption of orphaned infants has been observed in both wild and captive non-human primates. In two groups of wild bonnet macaques (Macaca radiata), we observed five instances of infants being cared for after they lost their mothers at a pre-weaning age (< 6 months). Orphaned infants had one or more caregivers (juvenile, subadult, and adult female or male) involved in carrying, grooming, hugging, and protecting them. Adoption did not appear to be related to the age/sex class of the infant, or directly to the mother’s rank. Although the dominance rank of the mother of an orphaned infant did not have a direct effect on orphan survivorship, it determined the number of caregivers available to the orphaned infant, and infant survivorship was positively related to the number of caregivers of the orphaned infant. Thus, survivorship was likely a function of the mother’s sociality. Two other infants born to high-ranking mothers were also adopted by more individuals and survived longer than the infants of low-ranking mothers.

Effects of Mother's Dominance Hierarchy on the Development of Social Relationships among Immature Tibetan Macaques.

Simple SummaryIn this study, we explored the role of females’ social ranks on social behaviors among immature Tibetan macaques (Macaca thibetana). The results suggest that females’ social ranks affected their offspring’s social play and grooming during the juvenile and adolescent periods, but not the infancy period. The present study provides new insight into understanding the effects of the female dominance hierarchy on the development of social relationships among immature offspring in nonhuman primates.During a relatively long period of growth, immature individuals rely on their mothers to obtain nutrition, and a good environment for learning social skills needed to cope with complex environments in adulthood. In this study, we collected the behavioral data of Tibetan macaques (Macaca thibetana) to investigate the effects of females’ social rank on the development of social relationships among their immature offspring from November to June 2021. The results show that there was no difference in the rate/type of social play and grooming among infants. However, among juveniles and adolescents, the higher their mother’s social rank, the higher the rate of social play they participated in, and the more aggressive play they engaged in. Immatures with high-ranking mothers initiated more social play among each other. A similar pattern of playmates was found among juveniles/adolescents with middle-ranking and low-ranking mothers. We also found that immatures preferred immatures with higher-ranking mothers as grooming mates and initiated more grooming with immatures with higher-ranking mothers than with those with lower-ranking mothers. Our study suggests that females’ social ranks affect the development of social relationships among their immature offspring. In despotic nonhuman primates, this indicates that the mother’s dominance hierarchy would directly or indirectly influence the processes of participating in social interactions and choosing partnerships among immature individuals with age (i.e., infancy, juvenile, and adolescent periods).

Male care in a free-ranging group of Japanese macaques (Macaca fuscata).

Among the Macaca species, adult male Japanese macaques are the least likely to perform male care (i.e., affiliative interactions between adult males and immatures, including holding, carrying, and grooming); however, they perform male care for infants, albeit infrequently. We examined 17 cases of male care observed for the first time when the immature was younger than 1year of age. Eleven of the 31 adult males who remained as central males during the 30-year observation period performed male care. Their age and dominance rank did not influence the occurrence of male care. Most cases were first recorded between the last part of the mating season and the first part of the birth season (January-March), whereas male care was rarely observed during the mating season (October-December). In 12 of the 17 cases, male care ceased within 6months after the first observation, whereas in the remaining cases, it continued for at least 1year. In 15 of the 17 cases, males tended to perform male care for matrilineally unrelated female infants of low-ranking mothers. In some cases, the male and infant mother showed grooming interactions for 6months both before and after the start of male care, whereas such grooming interactions were never recorded either before or after the start of male care in other cases. We also examined some hypotheses on male-immature associations and the probable benefits that males and infants might acquire through male care.

Sources of variation in weaned age among wild chimpanzees in Gombe National Park, Tanzania.

A key feature of human life history evolution is that modern humans wean their infants 2-4 years earlier on average than African apes. However, our understanding of weaning variation in apes remains limited. Here we provide the first such report in chimpanzees by examining weaned age variation using long-term data from Gombe National Park, Tanzania. We analyzed 41 years of observational behavioral data from 65 offspring of 29 mothers to examine the relationships between weaned age (defined as cessation of suckling) in wild chimpanzees and maternal age, dominance rank and parity, and offspring sex. We used Cox proportional hazards regression with mixed effects to model time to weaning and to examine potential sources of variation in offspring weaned age. We found that male offspring were less likely than female offspring to wean by a given age and that weaned age of males varied more than weaned age of females. In addition, maternal dominance rank interacted with offspring age, such that low-ranking mothers were less likely to wean offspring early, but this effect decreased with offspring age. We found that male offspring and offspring of low-ranking females were less likely to wean early, but did not find evidence for variable weaning according to maternal age or parity. As more data accumulate, we will be better able to disentangle the effects of maternal dominance rank, age and parity. Such studies will not only provide a richer understanding of living ape life history characteristics, but will also provide an important framework for understanding the evolution of early weaning in humans.

Early life social and ecological determinants of global DNA methylation in wild spotted hyenas.

Environmental factors early in life can have lasting influence on the development and phenotypes of animals, but the underlying molecular modifications remain poorly understood. We examined cross-sectional associations among early life socioecological factors and global DNA methylation in 293 wild spotted hyenas (Crocuta crocuta) in the Masai Mara National Reserve, Kenya, grouped according to three age classes (cub, subadult and adult). Explanatory variables of interest included annual maternal rank based on outcomes of dyadic agonistic interactions, litter size, wild ungulate prey density and anthropogenic disturbance in the year each hyena was born based on counts of illegal livestock in the Reserve. The dependent variable of interest was global DNA methylation, assessed via the LUminometric Methylation Assay, which provides a percentage methylation value calculated at CCGG sites across the genome. Among cubs, we observed approximately 2.75% higher CCGG methylation in offspring born to high- than low-ranking mothers. Among cubs and subadults, higher anthropogenic disturbance corresponded with greater %CCGG methylation. In both cubs and adults, we found an inverse association between prey density measured before a hyena was 3months old and %CCGG methylation. Our results suggest that maternal rank, anthropogenic disturbance and prey availability early in life are associated with later life global DNA methylation. Future studies are required to understand the extent to which these DNA methylation patterns relate to adult phenotypes and fitness outcomes.

Maternal Care in Free-Ranging Arboreal Grey-Cheeked Mangabeys (Lophocebus albigena johnstoni) in Kibale National Park, Uganda

Maternal styles have been intensively studied in a variety of terrestrial species of primates, but far less in arboreal species. However, to have a balanced view of the evolution of maternal care, it is necessary to investigate this behaviour in the context of habitat. Here, we investigate whether the mother’s parity, age and dominance rank, as well as the infant’s age and sex, influence maternal care and mother-infant proximity in arboreal grey-cheeked mangabeys (Lophocebus albigena). We observed 13 mother-infant dyads in four free-ranging groups for 6 months. Our main finding is that maternal care is a dynamic process affected by a mix of mothers’ and infants’ characteristics. We found that first-time mothers spent more time watching their infants than multiparous mothers, who carried, groomed and protected their infants more often. We also found that low-ranking mothers prevented their infants from leaving them more often than did high-ranking mothers. Moreover, mothers adjusted their care as infants became older. They groomed and protected female infants more than male infants, behaviours common in female-bonded species. Our study shows the ever-changing dynamics of maternal care related to infant age and highlights the role of the mother’s parity and rank in this process.

Maturation is prolonged and variable in female chimpanzees

Chimpanzees are important referential models for the study of life history in hominin evolution. Age at sexual maturity and first reproduction are key life history milestones that mark the diversion of energy from growth to reproduction and are essential in comparing life history trajectories between chimpanzees and humans. Yet, accurate information on ages at these milestones in wild chimpanzees is difficult to obtain because most females transfer before breeding. Precise age at first birth is only known from a relatively small number of non-dispersing individuals. Moreover, due to small sample sizes, the degree to which age at maturation milestones varies is unknown. Here we report maturation milestones and explore sources of variance for 36 wild female chimpanzees of known age, including eight dispersing females born in Gombe National Park, Tanzania. Using Kaplan-Meier survival analysis, including censored intervals, we find an average age of 11.5 years (range 8.5-13.9) at sexual maturity and 14.9 years (range 11.1-22.1) at first birth. These values exceed previously published averages for wild chimpanzees by one or more years. Even in this larger sample, age at first birth is likely underestimated due to the disproportionate number of non-dispersing females, which, on average, give birth two years earlier than dispersing females. Model selection using Cox Proportional Hazards models shows that age at sexual maturity is delayed in females orphaned before age eight years and those born to low-ranking mothers. Age at first birth is most delayed in dispersing females and those orphaned before age eight years. These data provide improved estimates of maturation milestones in a population of wild female chimpanzees and indicate the importance of maternal factors in development.

Maternal influences on primate social development

Primate mothers have the potential to influence the development of species-typical aspects of their offspring’s behavior as well as their individual sociosexual and reproductive strategies. If mothers experience psychosocial stress during pregnancy or lactation, their own stress hormones can have a long-term impact on their offspring’s physiology. The mother’s own behavior, especially weaning-related rejection, can be stressful to infants and have long-lasting effects on their infants’ neuroendocrine reactivity to stress. Exposure to variable maternal style early in life has long-term effects on the development of offspring behavior, including exploration and play, affiliation and aggression, and parenting. Primate mothers influence the kinship- and rank-related social preferences of their offspring, typically by providing opportunities to interact with some individuals more than others. Although mothers can contribute to the development of sex-typical behavior in the offspring (along with other environmental and genetic factors), sex-typical behavior is generally not the product of maternal socialization. Mothers provide their offspring with opportunities for social learning but rarely teach their infants new skills. In primate species with despotic dominance hierarchies, maternal transmission of rank through agonistic aid to their offspring makes a crucial contribution to the offspring’s fitness (as the daughters of high-ranking mothers reproduce more successfully than the daughters of low-ranking mothers). Future studies of maternal influences on social development could benefit from a deeper theoretical and experimental investigation of the evolutionary significance of these effects as well as of their underlying proximate mechanisms.

Sex and rank affect how infant rhesus macaques look at faces.

We investigated how differences in infant sex and mothers' dominance status affect infant rhesus macaques' (Macaca mulatta) interest in visually exploring emotional facial expressions. Thirty-eight infants were presented with animated avatars of macaque facial expressions during the first month of life. Sons of high-ranking mothers looked more at faces, especially the eye region, than sons of low-ranking mothers, but no difference in looking duration was found for daughters. Males looked significantly more at eyes than females, but this effect was reversed in infants who were reared without mothers in a primate nursery facility. In addition, in mother-infant interactions, mothers of sons were more likely to gaze at their infant's face compared to mothers of daughters. Combined with previous research indicating that rhesus macaque mothers interact differently with infants based on their own rank and infant's sex, these results support the view that social experiences shape early face preferences in rhesus macaques.

Maturation is prolonged and variable in female chimpanzees

Chimpanzees are important referential models for the study of life history in hominin evolution. Age at sexual maturity and first reproduction are key life history milestones that mark the diversion of energy from growth to reproduction and are essential in comparing life history trajectories between chimpanzees and humans. Yet, accurate information on ages at these milestones in wild chimpanzees is difficult to obtain because most females transfer before breeding. Precise age at first birth is only known from a relatively small number of non-dispersing individuals. Moreover, due to small sample sizes, the degree to which age at maturation milestones varies is unknown. Here we report maturation milestones and explore sources of variance for 36 wild female chimpanzees of known age, including eight dispersing females born in Gombe National Park, Tanzania. Using Kaplan-Meier survival analysis, including censored intervals, we find an average age of 11.5 years (range 8.5–13.9) at sexual maturity and 14.9 years (range 11.1–22.1) at first birth. These values exceed previously published averages for wild chimpanzees by one or more years. Even in this larger sample, age at first birth is likely underestimated due to the disproportionate number of non-dispersing females, which, on average, give birth two years earlier than dispersing females. Model selection using Cox Proportional Hazards models shows that age at sexual maturity is delayed in females orphaned before age eight years and those born to low-ranking mothers. Age at first birth is most delayed in dispersing females and those orphaned before age eight years. These data provide improved estimates of maturation milestones in a population of wild female chimpanzees and indicate the importance of maternal factors in development.

Bioactive factors in milk across lactation: Maternal effects and influence on infant growth in rhesus macaques (Macaca mulatta).

Among mammals, numerous bioactive factors in milk vary across mothers and influence offspring outcomes. This emerging area of research has primarily investigated such dynamics within rodent biomedical models, domesticated dairy breeds, and among humans in clinical contexts. Less understood are signaling factors in the milk of non-human primates. Here, we report on multiple bioactive components in rhesus macaque (Macaca mulatta) milk and their associations with maternal and infant characteristics. Milk samples were collected from 59 macaques at multiple time points across lactation in conjunction with maternal and infant morphometrics and life-history animal records. Milk was assayed for adiponectin (APN), epidermal growth factor (EGF) and its receptor (EGF-R), and transforming growth factor beta 2 (TGF-β2 ). Regression models were constructed to assess the contributions of maternal factors on variation in milk bioactives, and on the relationship of this variation to infant body mass and growth. Maternal body mass, parity, social rank, and infant sex were all predictive of concentrations of milk bioactives. Primiparous mothers produced milk with higher adiponectin, but lower EGF, than multiparous mothers. Heavier mothers produced milk with lower EGF and EGF-R, but higher TGF-β2 . Mothers of daughters produced milk with higher TGF-β2 . Mid-ranking mothers produced milk with higher mean EGF and adiponectin concentrations than low-ranking mothers. Milk EGF and EGF-R were positively associated with infant body mass and growth rate. Importantly, these signaling bioactives (APN, EGF, EGF-R, and TGF-β2 ) were significantly correlated with nutritional values of milk. The effects of milk signals remained after controlling for the available energy in milk revealing the added physiological role of non-nutritive milk bioactives in the developing infant. Integrating analyses of energetic and other bioactive components of milk yields an important perspective for interpreting the magnitude, sources, and consequences of inter-individual variation in milk synthesis. Am. J. Primatol. 78:838-850, 2016. © 2016 Wiley Periodicals, Inc.

Maternal effects on offspring stress physiology in wild chimpanzees.

Early life experiences are known to influence hypothalamic-pituitary-adrenal (HPA) axis development, which can impact health outcomes through the individual's ability to mount appropriate physiological reactions to stressors. In primates, these early experiences are most often mediated through the mother and can include the physiological environment experienced during gestation. Here, we investigate stress physiology of dependent offspring in wild chimpanzees for the first time and examine whether differences in maternal stress physiology are related to differences in offspring stress physiology. Specifically, we explore the relationship between maternal rank and maternal fecal glucocorticoid metabolite (FGM) concentration during pregnancy and early lactation (first 6 months post-partum) and examine whether differences based on maternal rank are associated with dependent offspring FGM concentrations. We found that low-ranking females exhibited significantly higher FGM concentrations during pregnancy than during the first 6 months of lactation. Furthermore, during pregnancy, low-ranking females experienced significantly higher FGM concentrations than high-ranking females. As for dependent offspring, we found that male offspring of low-ranking mothers experienced stronger decreases in FGM concentrations as they aged compared to males with high-ranking mothers or their dependent female counterparts. Together, these results suggest that maternal rank and FGM concentrations experienced during gestation are related to offspring stress physiology and that this relationship is particularly pronounced in males compared to females. Importantly, this study provides the first evidence for maternal effects on the development of offspring HPA function in wild chimpanzees, which likely relates to subsequent health and fitness outcomes. Am. J. Primatol. 80:e22525, 2018. © 2016 Wiley Periodicals, Inc.

Effects of reproductive condition and dominance rank on cortisol responsiveness to stress in free‐ranging female rhesus macaques

The hypothalamic-pituitary-adrenal (HPA) axis modulates individuals' physiological responses to social stress, which is an inevitable aspect of the daily lives of group-living animals. Previous nonhuman primate studies have reported that sex, age, rank, and reproductive condition influence cortisol levels under stressful conditions. In this study we investigated cortisol responses to stress among 70 multiparous, free-ranging female rhesus macaques (Macaca mulatta) on the island of Cayo Santiago, PR. Plasma cortisol samples were collected in two consecutive years under similar conditions. Twenty-two females were sampled both years, and most of those females were lactating in only one of the years. Individual differences in cortisol levels were stable across years, even though reproductive condition changed for most individuals. No relationship was found between age or social rank and cortisol levels. Of the females that changed reproductive conditions, cortisol levels were higher when they were lactating than when they were cycling, and the amount of change in cortisol from cycling to lactating was greatest for low-ranking individuals. Heightened reactivity to stress during lactation may be the result of concerns about infant safety, and such concerns may be higher among low-ranking mothers than among higher ranking mothers. Psychosocial stress and hyperactivation of the HPA axis during lactation can suppress immune function and increase vulnerability to infectious diseases, thus explaining why adult females in the free-ranging rhesus macaque population on Cayo Santiago have a higher probability of mortality during the birth season than during the mating season.

Abdominal odours of young, low-ranking European rabbit mothers are less attractive to pups: an experiment with animals living under natural breeding conditions

Lactating rabbit females (Oryctolagus cuniculus) emit abdominal, ventral odour cues that are involved in guiding the pups to the nipples. We tested the impact of the females’ social rank and age on the emission of this odour signal; both factors are highly collinear in wild rabbits and we did not aim to disentangle them in our study. We predicted that the abdominal odours of young, low-ranking females, which are usually more stressed and have a comparatively lower body condition, should be less attractive to pups than those of older, high-ranking females. For the experiment, we caught females from a population of wild-type European rabbits living in a large field enclosure. Pairs of adult females were placed in boxes with wire-mesh floor fixed over an arena with three rabbit pups aged 4–7 days old. First, we evaluated the effectiveness of our two-choice design by testing lactating females (LF) against non-lactating (NLF) controls (n = 17 pairs). In accordance with recent studies, the pups showed a clear preference for lactating females by means of a higher exploration time and more local searching under the ventral region of these females. In a second experiment, the pups chose between lactating 1-year-old, low-ranking females (Y/LRF) and older, high-ranking females (O/HRF) (n = 10 pairs). These latter assays suggested a comparatively lower attractiveness of Y/LRF females to pups searching for milk. Although the pups showed no significant preference in exploration behaviour, they preferably directed local searching events to the abdominal region of O/HRF females. In conclusion, our study suggests that mother–offspring chemosensory communication is modified by the mother’s age and social rank, i.e., by a complex of factors that are strongly linked in rabbit females in the wild.

Persistence of maternal effects in baboons: Mother's dominance rank at son's conception predicts stress hormone levels in subadult males

Dominance status and reproductive experience are maternal characteristics that affect offspring traits in diverse taxa, including some cercopithecine primates. Maternal effects of this sort are widespread and are sources of variability in offspring fitness. We tested the hypothesis that maternal dominance rank and reproductive experience as well as a male's own age and dominance rank predicted chronic fecal glucocorticoid (fGC) concentrations in 17 subadult wild male baboons, Papio cynocephalus (median age 6.5 years), in the Amboseli basin, Kenya. Among these variables, maternal dominance rank at a subadult male's conception was the sole significant predictor of the male's fGC and accounted for 42% of fGC variance; sons of lower ranking mothers had higher fGC than did those of high-ranking mothers. This result is striking because subadult male baboons are approximately 4–6 years past the period of infant dependence on their mothers, and are larger than and dominant to all adult females. In addition, many males of this age have survived their mothers' death. Consequently, the influence of maternal dominance rank persisted well beyond the stage at which direct maternal influence on sons is likely. Persistence of these major maternal influences from the perinatal period may signal organizational effects of mothers on sons' HPA axis. Although short-term, acute, elevations in GC are part of adaptive responses to challenges such as predators and other emergencies, chronically elevated GC are often associated with stress-related pathologies and, thereby, adverse effects on fitness components.

Maternal care and foal social relationships in a herd of Sorraia horses: Influence of maternal rank and experience

The influence of maternal rank and experience on patterns of maternal care and social relationships of foals were investigated in a managed herd of Sorraia horses, Equus caballus. Social interactions and spatial relationships of 13 foals (seven females and six males) born to seven mares were examined from birth to 10 months of life, within the three major periods of foal development. Conflict over suckling between dam and foal was not generally affected by rank and experience, but higher-ranking mothers allowed more suckling during late lactation than lower-ranking mothers. Foals of higher-ranking mares spent more time in proximity to the mother during socialization. Maternal rank and experience did not significantly affect maternal protectiveness, foal independence from the mother or the development of affiliative relationships between foals and group members. Foals of higher-ranking mares received lower frequencies of aggression from other horses only in the first month of life. Dominance relationships among foals depended mainly on aggressiveness and were not associated with maternal rank. The large variability in maternal behaviour, the absence of a significant association between maternal rank and body condition at parturition and the stable social environment within this herd may partly account for the reported results.

Maternal response to neonatal sibling conflict in the spotted hyena, Crocuta crocuta

Among spotted hyenas, Crocuta crocuta, neonatal aggression in twins is a well-known phenomenon and serves to establish intra-litter dominance soon after birth. As the stronger more aggressive cub presumably attains dominance over its twin, intra-litter dominance presents mothers with an ideal opportunity to assess individual cub fitness and, thereafter, to selectively favor one cub over the other. This study quantified maternal response to sibling conflict in 26 sets of twins born to 16 wild-living females to determine whether mothers of different social ranks exhibited favoritism towards sons or daughters, or in the case of same-sex twins, the dominant or subordinate cub. Maternal response to sibling conflict did not vary with litter sex composition, suggesting that mothers do not favor offspring of one sex over the other. All mothers intervened when their cubs fought and sometimes punished their cubs. Higher-ranking mothers more often punished both cubs, while lower-ranking mothers were more selective and punished the dominant cub. Where sibling aggression was most extreme, rather than favor the dominant sibling, mothers of all ranks made concessions to the subordinate cub that included procuring private nursing bouts for the subordinate and temporarily housing twins in separate dens, presumably to decrease sibling conflict. These findings represent a complex example of parent–offspring conflict and support both the insurance cub hypothesis and resource tracking hypothesis that mothers endeavor to keep all offspring alive for as long as possible in the event that the dominant sibling dies or that resources provide for the rearing of twins.