Signature whistles of free-ranging bottlenose dolphins Tursiops truncatus: stability and mother-offspring comparisons

Summary Mother-calf whistle exchanges were recorded from temporarily captured free-ranging bottlenose dolphins from 1975 to 1989. This is part of a long-term research project studying social structure and behavior of a community of approximately 100 dolphins in waters near Sarasota, Florida. Analysis of whistle exchanges from 12 mothercalf pairs shows that signature whistles can remain stable for periods up to at least 12 years. We looked for effects of vocal learning on the development of the signature whistle by comparing whistles of calves to those of their mothers. Eight female calves produced whistles distinct from those of their mothers, while four male calves produced whistles similar to those of their mothers. Male calves appeared to produce a greater proportion of whistles other than the signature whistle (termed variants). We hypothesize that these sex differences in whistle vocalizations may reflect differences in the roles males and females play in the social structure of the community.     

Early maternal investment in male and female African elephant calves

Summary The suckling behaviour of 130 freeranging elephant calves aged between birth and 4.5 years old was examined in Amboseli National Park, Kenya. Analyses of frequencies of suckling and durations of suckling bouts showed that males attempted to suckle more often, were more successful at their attempts, and as a result were estimated to have a higher milk intake than did female calves. Mothers were equally tolerant of their sons' and daughters' demands to suckle at young ages, but were less tolerant of their older sons' demands. The growth rates of males based on hind footprint length were faster than those of females from birth onwards. During drought years with low food availability, male calf survivorship in the first year was lower than that of female calves. During wet years, there was little difference between sexes in survivorship. It appeared that during dry years mothers were unable to sustain milk production at a level that met the metabolic requirements of their sons, and as result male calves were more likely to die. Females with a surviving son tended to have a longer interbirth interval than did females with a surviving daughter. We suggest that greater early maternal investment in male calves occurs because, in the highly-competitive polygynous mating system of elephants, size in adult male elephants is an important factor in mating success.     

Philopatry,reproductive success of females,and maternal investment in the red-necked wallaby

Summary Female red-necked wallabies settle within their mothers' home ranges, apparently for life, while males disperse at about two years of age. However, sons spend much more time with their mothers before dispersing than do daughters of similar ages. Females who associate regularly with their subadult offspring are less likely to reproduce successfully at their next breeding attempt than are females who spend little time with their subadults, and sons therefore impose greater short-term reproductive costs on their mothers than do daughters. Females who are generally gregarious also suffer reduced reproductive success, even though reproductive success is independent of local density. It is suggested that the reproductive costs to females of associating with their subadult offspring, and other relatives, are incurred through tolerance of ecological competition from those kin, and therefore reflect a form of prolonged maternal investment, which is initially heaviest in sons but is sustained for longer periods in daughters. Females produce equal numbers of male and female offspring, and spend equal amounts of time suckling them in infancy.     

Group histories and offspring sex ratios in ringtailed lemurs (Lemur catta)

Birth sex ratios were examined for ringtailed lemurs (Lemur catta) at the Duke University Primate Center. This population provides a long-term database of births under a variety of demographic and management conditions, including two semi-freeranging groups between which males transfer freely and females defend stable territorial boundaries. We examined three hypotheses usually considered in studies of primate sex ratio bias. The Trivers-Willard hypothesis predicts that dominant females produce males, local resource competition at the population level (LRC-population) predicts that the dispersing sex (males) will be overproduced in dense populations, and local resource competition among individuals (LRC-individual) predicts that dominant females overproduce the philopatric sex (females). We also examined a fourth hypothesis, local resource enhancement (LRE), which is usually subsumed under LRC-individual in studies of primate sex ratio evolution. LRE predicts that under certain conditions, females will produce the sex that provides later cooperative benefits, such as alliance support for within- or between-group competition. Our data provide support for LRE: females overproduce daughters given prospects of new group formation, either through group fission or threatened expulsion of young mothers. Behavioral data from Duke and also wild populations show that daughters serve mothers as important allies in this context and LRE effects also have been documented in other mammals that experience similar group histories. Nonsignificant trends in the data supported the LRC-population hypothesis, and we suggest that LRC interacts with LRE to explain offspring sex ratios in ringtailed lemurs. Received: 27 August 1999 / Received in revised form: 6 March 2000 / Accepted: 18 March 2000     

Whistle sharing in paired male bottlenose dolphins,<Emphasis Type="Italic"> Tursiops truncatus</Emphasis>

The signature whistle hypothesis states that dolphins produce highly stereotyped, individually distinctive whistles when in isolation. The presence of signature whistles has been called into question by recent studies proposing that dolphins produce a shared, simple upsweep whistle when in isolation, and that whistles produced by socializing dolphins are shared across individuals and social groups. This shared repertoire hypothesis suggests that when two animals produce the same whistle type, it is due to sharing the same common repertoire rather than one animal learning to produce the whistle of another. One difference between studies supporting or denying the existence of signature whistles is the method used to classify whistle types. We examined whistle production by 17 free-ranging bottlenose dolphins while temporarily restrained. We used both a quantitative comparison technique similar to that used to support the shared repertoire hypothesis and human judges to classify whistle types and quantify similarity between types. Contrary to recent studies that emphasize shared whistles, overall whistle sharing between isolated individuals was low (25%) and a simple upsweep did not account for the most common whistle type in half of the animals. Some species of birds, bats, and primates with stable social groups use vocal learning to converge over time to one common group distinctive call type. We examined whistle similarity between adult male dolphins that are partners in a close social alliance in order to test whether vocal learning may enable a similar vocal convergence. Whistle similarity was rated very high between partners and low between non-partners by both the quantitative technique and human observers. This suggests that as in songbirds and some other mammals, adult male bottlenose dolphins may use vocal learning to converge on similar whistles as they develop affiliative social relationships.Communicated by G. Wilkinson     

Maternal quality and differences in milk production and composition for male and female Iberian red deer calves (Cervus elaphus hispanicus)

Several theories predict a sex-biased investment either through unbalanced sex ratios in offspring or through differences in provisioning. According to them, one would expect an optimisation in indirect fitness, or else a compensation for increased mortality of one sex. In addition, biases in provisioning may also arise as a consequence of weight-dependent non-adaptive nutrient demands by offspring. This study examines milk provisioning and sex biases in offspring sex ratio together with maternal quality variables. Mothers of higher quality (weight and age) showed greater milk provisioning ability (in terms of production) resulting in greater calf weight gain. Mothers of sons produced greater yields of milk, milk protein, fat and lactose than mothers of daughters, and increased percentage of protein after controlling for higher male birth weight. In contrast, mothers of males did not differ from mothers of females in age or any body weight variables related to maternal quality. These results suggest that differences in milk production and composition for sons and daughters are rather a mechanism to optimise indirect fitness than a mechanism to compensate for increased mortality in male calves, or a consequence of greater weight-dependent nutrient demands by heavier male calves. Results also suggest that biases in milk provisioning may occur without biases in offspring sex ratio, and furthermore, in contrast to the prediction that biases should be relative to the mean investment of the population, that milk provisioning biases might not be relative.Communicated by F. Trillmich     

Reproductive isolation between populations from Northern and Central Europe of the leaf beetle Chrysomela lapponica L.

Summary. Allopatric populations of the leaf beetle Chrysomela lapponica are known to feed upon either willow (Salicaceae) or birch (Betulaceae). This study aimed to elucidate the differentiation process of these allopatric populations. We investigated whether these allopatric populations specialized on different host plants are still able to produce fertile offspring when interbreeding. Individuals from a population in Finland (willow specialists) and one in the Czech Republic (birch specialists) were crossed in laboratory. Hybrid formation succeeded only between females from the Czech, birch specialized population and males from the Finnish, willow specialized population, while no eggs were produced by females of the willow specialists having mated males of the birch specialists. Behavioral, morphological, physiological, and chemical features of the F₁ hybrids were studied. The chemical composition of larval defensive secretion and feeding preferences of the resulting F₁ hybrids mainly showed similarities with the paternal phenotype, while the area of black coloring on the offspring’s elytra was intermediate between those of the parental elytra. F₁ hybrids did not accept the host plant (birch) of their mothers for feeding and only survived on willow. Thus, since mothers only lay eggs on birch, we found evidence for a postzygotic isolation mechanism between the individuals of the two investigated populations: when having been mated with willow specialized Finnish males, the birch specialized Czech mothers place the hybrid eggs on a plant species (birch), on which the hatching larvae cannot survive.     

Use of signature whistles during separations and reunions by wild bottlenose dolphin mothers and infants

We examine the contexts and patterns of signature whistle production by wild bottlenose dolphin mother-infant pairs (Tursiops spp.) to gain insight into the functional significance of whistles. Results are based on focal observations and simultaneous recordings of underwater vocalizations. Whistles occur primarily when mother-infant pairs are separated, and the probability of whistles increases with distance of separation. The timing of whistles during separations varies, but whistles tend to be produced in repetitive series and are generally concentrated toward the later stages of the separation, i.e., during the process of reunion. Although we focused on infants, mothers do not appear to whistle during separations as frequently as infants. Infant whistles may function to facilitate reunions by conveying information to the mother concerning the infant's motivation to reunite and/or its location. Infant whistles could induce a cooperative response from the mother including approach, slowing to allow the infant to catch up or whistling. Highly individualized signature whistles may be particularly useful in a fission-fusion society in which individuals (mothers and infants as well as adults) join and leave temporary parties in a fluid manner, yet maintain consistent, long-term associations with particular individuals. Correspondence to: R.A. Smolker     

The relationship between maternal phenotype and offspring quality: do older mothers really produce the best offspring?

Maternal effects are increasingly recognized as important drivers of population dynamics and determinants of evolutionary trajectories. Recently, there has been a proliferation of studies finding or citing a positive relationship between maternal size/age and offspring size or offspring quality. The relationship between maternal phenotype and offspring size is intriguing in that it is unclear why young mothers should produce offspring of inferior quality or fitness. Here we evaluate the underlying evolutionary pressures that may lead to a maternal size/age-offspring size correlation and consider the likelihood that such a correlation results in a positive relationship between the age or size of mothers and the fitness of their offspring. We find that, while there are a number of reasons why selection may favor the production of larger offspring by larger mothers, this change in size is more likely due to associated changes in the maternal phenotype that affect the offspring size-performance relationship. We did not find evidence that the offspring of older females should have intrinsically higher fitness. When we explored this issue theoretically, the only instance in which smaller mothers produce suboptimal offspring sizes is when a (largely unsupported) constraint on maximum offspring size is introduced into the model. It is clear that larger offspring fare better than smaller offspring when reared in the same environment, but this misses a critical point: different environments elicit selection for different optimal sizes of young. We suggest that caution should be exercised when interpreting the outcome of offspring-size experiments when offspring from different mothers are reared in a common environment, because this approach may remove the source of selection (e.g., reproducing in different context) that induced a shift in offspring size in the first place. It has been suggested that fish stocks should be managed to preserve these older age classes because larger mothers produce offspring with a greater chance of survival and subsequent recruitment. Overall, we suggest that, while there are clear and compelling reasons for preserving older females in exploited populations, there is little theoretical justification or evidence that older mothers produce offspring with higher per capita fitness than do younger mothers.     

Pathological and toxicological investigations of chronic nitrate poisoning in cattle

In this study, three stillborn and five premature and blind calves from cows suffering from chronic nitrate poisoning were examined pathologically. The live calves were euthanized, due to a poor prognosis. Complete necropsy was performed on all the eight affected calves. Hydroperitoneum and ascites were the most predominant lesions in the stillborn calves. Edema and hemorrhage were observed in the lungs of all the calves. Hemorrhages were seen in the digestive systems of four calves. Blood samples were taken from five live calves and their mothers, and from two cows that had delivered stillborn or blind calves, for biochemical observations and measurement of methemoglobin levels. Blood values of five healthy cattle from a herd with no history of stillborn or blind calves were used as controls. Water and feed samples were taken from five different farms with a history of stillborn or blind calves and examined for nitrates. Nitrate and nitrite levels of feeds ranged from 154 to 480 and 1.5 to 20?ppm respectively; these levels can cause chronic poisoning. Levels of nitrates and nitrites in blood samples from cows and calves were found to be high. While hemoglobin values were found to be low, methemoglobin values were very high. No differences were observed between liver enzyme levels of the affected and control cattle. There were no bacteria or virus isolated from these animals, and histopathological examination revealed no indication of any other diseases.     

Sex bias or equal opportunity? Patterns of maternal investment in bison

Summary In polygynous mammals, it may be adaptive for mothers to invest more in sons and/or to adjust the sex ratio of offspring in relation to body condition. Calving patterns were examined over an 8-year period (1982–1989) for a population of Bison bison in which barren females are not selectively culled. From these data, we tested predictions of the sex ratio adjustment hypothesis as well as two assumptions: (1) that offspring weight at the end of the period of parental investment (PI) is correlated with later condition, and (2) that maternal and offspring condition during the period of PI are correlated. In contrast to predictions, there was little evidence that mothers in better condition bear more sons. Short- and long-term measures of maternal condition (previous reproductive status, age, dominance status, pre-pubertal body weight, age at first reproduction, birth date, and the duration of the mother's own suckling period) were little related to offspring sex ratio, although the last calves of old females were nearly always female. Similarly, there was little evidence for sex-biased PI. Weights at about 7 months of age were greater for males than females; males also had somewhat later birth dates, suggesting either longer gestation or later conception. However, maternal reproductive costs, as measured by subsequent fecundity, weight loss, and interbirth intervals, did not vary with calf sex. Both assumptions of the model received some support. However, while maternal condition was correlated with offspring condition, there may be sex differences in investment patterns. Mothers appear better able to influence the condition of daughters than of sons. This sex difference may negate any benefit from male-biased investment.     

Delayed breeding in the cooperatively breeding Florida scrub-jay (Aphelocoma coerulescens): inhibition or the absence of stimulation?

To determine whether fundamental differences exist in the reproductive physiology of breeder and nonbreeder Florida scrub-jays (Aphelocoma coerulescens), we compared plasma levels of testosterone (T) and luteinizing hormone (LH) in males, and estradiol (E₂) and LH in females. Although male breeders had higher overall T and larger testes, nonbreeders’ T paralleled that of breeders, and their testes were more than an order of magnitude larger than regressed testes. Breeder and nonbreeder males had equivalent baseline LH, and equivalent changes in LH following a gonadotropin-releasing hormone (cGnRH-I) challenge. The T, LH and GnRH challenge data indicate that nonbreeder males have functional hypothalamo-pituitary-gonadal (HPG) axes. We found no hormonal evidence of inbreeding suppression in males: nonbreeders that did not live with their mothers and those that did had similar T. Male nonbreeders that were exposed to E₂-implanted females had higher T than did controls, suggesting that the lack of within-pair stimulation is a key factor in whether an individual delays breeding. Female nonbreeders had E₂ titres equal to or higher than breeders and neither basal LH nor LH following GnRH challenge differed by breeding status. Nonbreeders’ ovarian follicles were smaller than breeders’, but were larger than they would be during the non-breeding season. These data suggest that nonbreeders were primed for breeding and were simply waiting for an opportunity or a required stimulus. Female nonbreeders that lived in a territory with an unrelated male breeder had significantly higher E₂ than those that remained with their fathers. Similarly, nonbreeders that were captured away from their home territories had elevated E₂. However, nonbreeders that lived with their fathers had E₂ that was equivalent to breeding females, suggesting that inbreeding avoidance may not be the primary factor leading to delayed breeding in females. Received: 13 June 1995 /Accepted after revision: 27 April 1996     

The association of female kin in the arctic ground squirrel Spermophilus parryii

Summary The association of kin in arctic ground squirrels (Spermophilus parryii) was studied near Haines Junction, Yukon Territory, during the summers of 1977, 1978, and 1979. Males dispersed in this population, thus only females were likely to live near relatives.Close female kin (sisters, mothers/daughters) had greater overlap of home ranges and interacted more amicably and less agonistically than did less closely related females. Closely related females clumped their young at emergence, whereas more distantly related females did not; there was little indication of clumping of pre-emerged young. The overlap of home ranges of distant relatives (known genetic relatives that had not associated in a natal burrow) was intermediate to that of close relatives and non-relatives. The types of interactions between distant relatives were more similar to those between non-relatives than between close relatives.I conclude that female arctic ground squirrels exhibit nepotism. Females may benefit from associations with relatives by sharing watching for predators once juveniles become conspicuous. Male arctic ground squirrels commit infanticide and several females may be more effective at protecting their young from infanticidal males than females living alone. I suggest that clumping of young by close relatives may provide a mechanism allowing distantly related females to identify each other.Address for offprint requests     

Sexual dimorphism in sea lion pups: differential maternal investment,or sex-specific differences in energy allocation?

Proximal mechanisms underlying a faster growth rate in male compared to female California sea lion pups were investigated. Males are significantly larger at birth than females. Specifically, we asked if differential maternal investment contributed to enhanced male growth via: (1) larger mothers having disproportionately more male pups, (2) more time and energy put into foraging by mothers of male pups, and (3) greater milk production in mothers of male pups. We also considered four aspects of differential energy utilization and acquisition by male and female pups: (1) male pups attempting to save energy for growth by changes in behavior, (2) longer suckling bouts with mother and more sneak suckling of non-filial females by male pups, (3) lower maintenance costs in males via a lowered resting metabolic rate, and (4) increased assimilation efficiency in males. Our study showed that there are no differences in the size of females or length of foraging trips for mothers of male and female pups. Male pups received more milk from their mothers, but the difference was no longer significant when the larger body size of males was considered. There were no differences in either the activity budgets or suckling behavior of male and female pups. Male pups, however, did have lower resting metabolic rates than females. We conclude that enhanced male perinatal growth is a consequence of a larger size at birth, proportionally more milk from mothers to support the greater demands of larger body size, and lower maintenance costs due to a lower resting metabolic rate. Received: 28 April 1995/Accepted after revision: 25 July 1995     

Age-related foetal sex ratio bias in Iberian red deer (<Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis>): are male calves too expensive for growing mothers?

Several theories predict offspring biases towards males or females with increasing reproductive resources of the mother to maximize reproductive returns by offspring, or as a result of prohibitive cost of the most expensive sex for young mothers or those in poor condition. This study examines foetus sex of 221 harvested hinds in a food-supplemented game estate for 10 years, according to hind age class (yearlings, subadults or adults), precise age, body mass and condition, and jaw length. A logistic model showed that hinds had a greater probability of bearing a male foetus with increasing age class, but not with any other variable. The greatest bias was found in yearling hinds. After controlling for age class and mass, jaw length was smaller in pregnant compared to non-pregnant yearlings and subadults, which suggests a trade-off between reproduction and growth. The bias towards females in yearlings increased as gestation proceeded, which suggests that the bias might be a result of selective abortion of male foetuses. Although results do not exclude an investment in males to increase number of grand-offspring, they suggest that young hinds may produce daughters as a trade-off between low energetic-cost offspring and their need to grow.Communicated by S. Krackow     

Mate choice among sympatric fur seals: female preference for conphenotypic males

When closely related species breed in sympatry, and where hybrids have lower fitness, reinforcement theory predicts that selection should favour mechanisms that reduce the probability of interspecific matings. If this situation arises among species that exhibit resource defence polygyny where males and females of different species reside in the same territories, there may be some conflict between mate choice based on territory-holding ability (sexual selection) and mate choice for correct species. We investigated this in a population of fur seals where three species are sympatric and where some females breed in the territories of heterospecific males, and where interspecific matings and hybrid pups are observed. The territorial status of males and the birthing sites of females were determined during daily observations, as were the movements of males and females, the location of matings and mating partners. DNA extracted from skin samples was used to determine paternities using DNA fingerprinting and the mtDNA genotype of individuals. Individuals were also classed on the basis of species-typical phenotype. We found that extra-territory inseminations (ETIs) were significantly more prevalent (67%) when territorial males and resident females were of different phenotype than when of similar phenotype (27%), but mtDNA genotype had no effect on the rate of ETIs. ETIs were probably by males with the same phenotype, as pups born to these females in the following season had the same phenotype as their mothers, suggesting they were not hybrids. These results suggest that within the resource defence polygynous mating system of these sympatric fur seals, female mate choice is more influenced by male phenotype than genotype. Contrary to our predictions, our study indicates that potential conflict between mate choice based on sexual selection and species recognition is unlikely, because females have some capacity to discriminate between males both within and between species on phenotypic traits additional to those under sexual selection. Although at least 25% of the pups born in this study were hybrid, this study can only support reinforcement theory if hybrids have reduced fitness. The fitness of hybrids among the species studied is currently unknown. Received: 19 January 1998 / Accepted after revision: 12 September 1998     

Brood-provisioning strategies in Dawson's burrowing bee, Amegilla dawsoni (Hymenoptera: Anthophorini)

Males of Dawson's burrowing bee are dimorphic in size. Although large (major) males defeat smaller ones in competition for emerging females and therefore are more likely to mate, majors are greatly outnumbered by half-sized (minor) males. Nesting females might produce many minor males, despite their low reproductive value, because female behaviour is governed by a mixed evolutionarily stable strategy (ESS), in which case the ratio of majors to minors should not be affected by changes in female condition. In contrast, a conditional-strategy hypothesis predicts that older, wing-worn or stressed females unable to forage efficiently should be especially likely to produce minor offspring, which require less brood provisions. To test these alternative hypotheses, we manipulated the condition of nesting female bees by the addition of weights and the removal of their wing margins. These manipulations, done early in the flight season, failed to increase the production of minor males, a result consistent with the mixed-ESS hypothesis. However, unmanipulated females were far more likely to produce minor males if they were small or if they were nesting late in the season, when foraging conditions had deteriorated, findings that are consistent with a conditional provisioning strategy. Thus it appears that the abundance of minor males is the result of a conditional provisioning strategy of nesting females, which may be superimposed on a fixed tendency to produce large offspring early in the season and small ones later.     

Social behavior of free-ranging juvenile sooty mangabeys (Cercocebus torquatus atys)

Sooty mangabeys are terrestrial monkeys exhibiting female philopatry and male dispersal. Studies in captivity as well as in the wild have found that adult females form linear dominance hierarchies. However, while captive studies found no evidence for a matrilineal social system, a previous study in Taï National Park, Ivory Coast, suggested that relatedness could influence both dominance rank and affiliation pattern among adult females. Here I test whether the dominance rank, coalitionary behavior, and affiliative behavior of juveniles in a group of free-ranging mangabeys in the Taï National Park are in accordance with a matrilineal, individual, or age-related dominance system. I found that juvenile females' dominance ranks remained stable over time and were highly correlated with the dominance ranks of their mothers, whereas juvenile males' dominance ranks were initially correlated with the ranks of their mothers, but showed greater instability with increasing age. Moreover, coalitions occurred mainly between juveniles and animals that were close in rank, including their mothers and siblings. Finally, juvenile females associated and groomed preferentially with close-ranking juvenile and adult females. Juvenile males showed similar preferences in affiliation with adult females, but when associating with juvenile males, they preferred peers. The observed social behavior of free-ranging juvenile sooty mangabeys resembled the social behavior described for juveniles of many matrilineal primate species.     

High-ranking females bias their investment in favour of male calves in captive Ammotragus lervia

Calf suckling behaviour is a valid measure of maternal investment in the Saharan arrui, Ammotragus lervia sahariensis, since this variable is strongly correlated with the inter-birth interval. High-ranking females allocate their resources preferentially towards their sons, as the average suckling rate is significantly higher in male calves than in female calves during their 1st month of life, when maternal investment reaches the highest values of the whole lactation period. However, average suckling-bout duration shows no sex differences. Some maternal behaviours, such as sniffing and licking, are strongly correlated with suckling events. Only during the calves’ 1st week is the mother responsible for maintaining proximity; but from the following week on the calves are mainly responsible for maintaining it. In addition, when the calves are 1 month old, high-ranking females tend to maintain a stronger link with their male calves. Female calves spend more time with their mothers than male calves during their 1st month of life, if the mother’s rank is lower than 60%; otherwise, the opposite occurs, male calves being close to their mothers for longer, even from their 1st week of life. Finally, the higher the maternal rank the higher the proportion of male calves delivered. Received: 18 April 1995/Accepted after revision: 11 February 1996     

“Friendships” between new mothers and adult males: adaptive benefits and determinants in wild baboons (<Emphasis Type="Italic">Papio cynocephalus</Emphasis>)

Close associations between adult males and lactating females and their dependent infants are not commonly described in non-monogamous mammals. However, such associations [sometimes called “friendships” (Smuts 1985)] are regularly observed in several primate species in which females mate with multiple males during the fertile period. The absence of mating exclusivity among “friends” suggests that males should invest little in infant care, raising questions about the adaptive significance of friendship bonds. Using data from genetic paternity analyses, patterns of behavior, and long-term demographic and reproductive records, we evaluated the extent to which friendships in four multi-male, multi-female yellow baboon (Papio cynocephalus) groups in Amboseli, Kenya represent joint parental care of offspring or male mating effort. We found evidence that mothers and infants benefited directly from friendships; friendships provided mother–infant dyads protection from harassment from other adult and immature females. In addition, nearly half of all male friends were the genetic fathers of offspring and had been observed mating with mothers during the days of most likely conception for those offspring. In contrast, nearly all friends who were not fathers were also not observed to consort with the mother during the days of most likely conception, suggesting that friendships between mothers and non-fathers did not result from paternity confusion. Finally, we found no evidence that prior friendship increased a male’s chances of mating with a female in future reproductive cycles. Our results suggest that, for many male–female pairs at Amboseli, friendships represented a form of biparental care of offspring. Males in the remaining friendship dyads may be trading protection of infants in exchange for some resources or services not yet identified. Our study is the first to find evidence that female primates gain social benefits from their early associations with adult males. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.