Reproductive success increases with degree of kinship in cooperative coalitions of female red howler monkeys (Alouatta seniculus)

Evaluation of evolutionary mechanisms proposed to promote cooperative behavior depends on the relative influence of the behavior on the reproductive success of individuals, the reproductive success of the group in which they interact behaviorally, and the degree of gene correlation among cooperators. The genetic relationship within cooperative coalitions of female red howler monkeys was examined for three populations with different densities and growth rates. Patterns of gene correlation change within coalitions is documented using data from the mitochondrial and nuclear genomes, and long-term census monitoring. Differences in fecundity and infant survivorship within and between groups of unrelated (rˉ=0) and related (rˉ≥ 0.25) females are compared. Females that emigrate from their natal groups form coalitions with other migrant females. These coalitions attempt to establish a territory and, once successful in producing offspring, exclude other females from feeding resources. Females in these coalitions had different mtDNA haplotypes and a genetically estimated mean r of 0, supporting demographic data on emigration patterns indicating that these females rarely have the opportunity to form coalitions with kin. Patterns of recruitment and rate of matriline development within social groups supported behavioral data indicating that females actively attempt to promote their own matriline as breeders over that of other females, and that some matrilines are more successful at this than others. Mean r among females was significantly higher in coalitions established as social groups for several generations (rˉ=0.44). In these groups, females all shared the same mtDNA haplotype, and mtDNA haplotype divergence was significantly higher between than within groups. Females in coalitions with kin had significantly higher reproductive success than females in unrelated coalitions in all populations. This difference was not a function of coalition size, number of males, socionomic sex ratio, or primiparity, although anecdotal evidence suggests that allomothering may compensate for inept new mothers in related coalitions more often than in unrelated ones. Differences in territory quality could not be ruled out as a potential causal factor in the saturated populations, but were unlikely in the low-density, growing population. There were substantial differences among long-established coalitions in overall reproductive output in all three populations, and this was significantly correlated with the number of breeding females. Increase in coalition size was a function of both group age and the behavioral tolerance among females. Regardless of the underlying reasons for the patterns observed, reproductive success clearly increases with degree of gene correlation among females within cooperative coalitions, and coalitions that recruit more daughters produce more offspring. The nature of the cooperative relationship among group females directly influences both of these outcomes. This is associated with substantial genetic differentiation among social groups within populations, creating conditions in which genetic tendencies towards cooperative behavior can become tightly associated with group reproductive success. Received: 15 September 1999 / Revised: 27 April 2000 / Accepted: 27 May 2000     

Infant handling by female Barbary macaques (Macaca sylvanus) at Affenberg Salem: testing functional and evolutionary hypotheses

Assisting the genetic parents in the rearing of young, a widespread phenomenon in many birds and mammals, is usually regarded as an altruistic or mutualistic behavior. Infant handling by females other than the mother is also common in many primates, but due to high within- and between-species variation and limited knowledge about fitness consequences there is no consensus about its evolutionary and functional significance. Analysis of female infant-handling patterns and its reproductive consequences in three groups of semifree-ranging Barbary macaques revealed that nulliparous females significantly more often handled infants than parous females, but infant handling experience did not affect survival of their own first live-born offspring. Females interacted preferentially with closely related infants, but infant handling frequency improved neither infant survival nor maternal fecundity. Reciprocation of infant handling by mothers was rare. Although “aunting to death” occurred in the population, the hypothesis that infant handling serves to reduce the fitness of competitors was not supported. Limited evidence suggests that females at least sometimes use infants as strategic tools in the course of alliance formation. In concert with this poor evidence for a functional basis of the behavior, several lines of evidence support the hypothesis that infant handling evolved as a non-adaptive by-product of a strong selection for mother-offspring bonding. (1) Rates of infant handling were highest among females that experienced early infant loss. (2) Females caring for infants or yearlings of their own handled other infants significantly less often than females without dependent offspring. (3) Infant handling by females was most prevalent during the infants’ first month of life. (4) Both “aunting to death” and a successful adoption occurred irrespective of kinship relations. Although the by-product hypothesis appears to be the only one able to explain all results of this study, the apparent rarity of infant handling in non-female-bonded species suggest that kin selection is a possible alternative explanation for the evolution of female infant-handling in primates. Received: 30 December 1995/Accepted after revision: 10 May 1996     

Association patterns of spider monkeys: the influence of ecology and sex on social organization

Summary In this paper I consider how the costs and benefits of group living in spider monkeys (Ateles geoffroyi) vary between troop members. The results suggest that ecological factors set an upper limit to the number of spider monkeys that can associate and still efficiently exploit the available resources. In addition, the needs of the individual appears to influence the type and size of the subgroup it chooses. Adult males band together, travel over a large area, and are frequently sighted near the community's boundary. In contrast, adult females spend more time solitary than males and have association patterns that are strongly influenced by the presence of a dependent infant. Females with dependent infants tend to travel in small subgroups or alone, avoid the boundaries of the community's home range, and exhibit a restricted pattern of use of their range. The results suggest that males may be attempting to locate females with which they can breed, while mothers attempt to protect their infants by avoiding conspecifics and potentially dangerous situations near territorial boundaries.     

Communal nesting, kinship, and maternal success in a social primate

Communal nesting, where several mothers regularly pool and cooperatively rear offspring, is unusual in mammals. This type of crèching behavior is especially rare among primates, with the notable exceptions of humans, some nocturnal strepsirrhines, and—as we show in this study—black-and-white ruffed lemurs (Varecia variegata). Here, we combine data on nesting behavior, genetic relatedness, and infant survivorship to describe variation in ruffed lemur infant care and to examine the potential benefits of ruffed lemur communal breeding. Reproductive events were rare, and females produced litters (synchronously) only once in 6 years of observation. We show that not all mothers participate in communal crèches, but those that did had greater maternal success; communal breeders spent more time feeding and their offspring were more likely to survive. Although cooperating mothers were often related, females also cooperated with non-kin, and those who shared infant care responsibilities had greater maternal success than mothers who did not participate. If there is indeed a causal link between maternal cooperation and reproductive success, this unusual behavior, like that of human communal rearing, may have evolved via some combination of kin selection and mutualism.     

Activity budgets in two species of bee flies (Lordotus: Bombyliidae,Diptera): a comparison of species and sexes

Summary This study compares time budgets of males and females of two sympatric species of bee flies, Lordotus pulchrissimus and L. miscellus. These species occur synchronously in sand dune habitat on the north shore of Mono Lake, Mono Co., California, and feed almost exclusively on the flowers of a composite shrub, Chrysothamnus nauseosus. The two closely related species, with similar ecological requirements and in an identical environment, allocate time and energy in very different ways, even when accomplishing the same ecological goals. L. pulchrissimus males and females engage respectively in aggressive interactions and feeding primarily in the morning. In contrast, L. miscellus males and females intersperse brief periods of resting with flying when defending territories and feeding, and they keep up these activities until later in the day. Similarly within species, males are involved in aggressive interactions for a shorter period each day, and females feed over a longer period. Flies in all sex-species classes but male pulchrissimus allocate their time energetically in similar ways. Male pulchrissimus spend more time each day in energetically costly activities; they engage in continuous hovering flight and intense interactions in aerial aggregations. While male miscellus feed little, male pulchrissimus spend a large portion of their time feeding, approximately as much as females, contrary to the expectation that males should be foraging-time minimizers. This study corroborates the conclusions of previous studies on bee-fly communities by showing that nectar and pollen are important resources for adult bee flies, at least for some species: Individuals of these two species spend a large proportion of their adult lifetimes feeding.     

Influence of male harassment and female competition on female feeding behaviour in a sexual–asexual mating complex of mollies (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, <Emphasis Type="Italic">P. formosa</Emphasis>)

Differential interests between the sexes regarding the number of copulations can result in sexual harassment. Hence, females may have less time available for foraging. Male sexual harassment often leads to fitness reduction in females. We used the mating complex of the bisexual fish Poecilia mexicana and the co-occurring all-female Poecilia formosa to study sexual harassment and its incurred cost on female feeding efficiency. P. formosa is a sperm-dependent parthenogen that requires mating with host males to induce embryogenesis, but the male genes are not used. We therefore predicted P. mexicana males to prefer conspecific females. Hence, costs of male sexual harassment should not occur in unisexuals. While P. formosa are at a disadvantage compared to P. mexicana females due to male mate choice (leading to sperm limitation), this could be traded-off by suffering less from sexual harassment. In our experiment, we found males to direct significantly more pre-copulatory mating behaviour towards conspecific females, whereas actual mating attempts did not differ between species. Contrary to our prediction, both types of females started feeding later and spent less time feeding in the presence of a male partner compared to the time spent feeding with another female, suggesting that females of both species suffer from male harassment. The focal females' feeding time declined with increasing body size of the female competitor, and the same pattern was found when a male was present. We discuss that—besides sexual harassment—other factors such as food competition and female mate choice may affect female feeding efficiency.     

Effects of feeding experience in the copepod Eucalanus pileatus: a cinematographic study

High-speed microcinematography was used to examine the effects of prior experience with particular cell types on the feeding efficiency of a calanoid copepod. Female Eucalanus pileatus were fed monocultures of either the 5-m diatom Thalassiosira pseudonana or the 11-m diatom T. weissflogii during a 2-to 3-d preconditioning period. The smaller diatoms are accumulated passively by the second maxillae while the larger diatoms are detected and actively captured as individual cells. Four females from each preconditioning culture were transferred to a monoculture of the large cells and their behavior filmed at five intervals over a 24-h period to determine whether a loss of efficiency occurs when the copepods must shift capture modes. Ingestion rates for females experienced with the larger cells were approximately 2.5 times higher than those of inexperienced females. Six sequential behavioral steps in the feeding process could alter ingestion rates: (1) amount of time spent flapping the feeding appendages. (2) rate of flapping of the feeding appendages, (3) ability to detect individual cells, (4) success rate of capture attempts, (5) capture and handling time per cell and (6) rejection rate of captured cells. An increased ability to detect cells and a decreased rejection rate contributed significantly to the higher ingestion rate of experienced feeders, indicating that copepods have the ability to learn during the feeding process. Grazing rates may be seriously underestimated in experiments which do not include a preconditioning period, especially those which calculate ingestion over short time intervals. Such effects may also influence the feeding of copepods in the field when encountering changes in particle spectra through vertical migration or horizontal displacement.     

Vigilance in wild Thomas's langurs (Presbytis thomasi ): the importance of infanticide risk

This study examines vigilance as a behavioural indicator of the importance of infanticide risk by comparing the infanticide avoidance hypothesis with the predation avoidance and mate defence hypotheses for wild Thomas's langurs (Presbytis thomasi) in Sumatra. We found that all individuals were more vigilant in situations of high predation risk, i.e. lower in the trees and in the absence of neighbours. Females were also more vigilant on the periphery of the group. However, there were variations in vigilance levels that could not be accounted for by the predation avoidance hypothesis. Males without infants showed higher levels of vigilance in areas of home range overlap than in non-overlap areas during the early phase of their tenure, strongly suggesting mate defence. In these areas of home range overlap where Thomas's langur groups can interact, males may attack females and infants, and so the infanticide risk for males and females with infants is likely to be high in these areas. Only females with infants, but not males with infants or females without infants, showed higher vigilance levels in overlap areas than in non-overlap areas; in addition, in overlap areas, females with an infant were more vigilant than females without an infant, while this was not the case in non-overlap areas. Both females and males with infants were more vigilant high in the trees than at medium heights in overlap areas but not elsewhere. These findings can only be explained by the infanticide avoidance hypothesis. In contrast to predator attacks, infanticidal male attacks come from high in the canopy, and only occur in overlap areas. There was a significant sex difference in vigilance, but males were only more vigilant than females without an infant, and not more vigilant than females with an infant. We conclude that vigilance varied mainly in relation to the risk of predation and infanticide. Mate competition only played a role for males during the early phase of their tenure. Predation risk seems to offer the best explanation for vigilance for all individuals in the absence of infants. Both predation risk and infanticide risk played a role for females and males with infants. Received: 4 April 1998 / Accepted after revision: 6 September 1998     

Effects of male sexual harassment on female time budgets,feeding behavior,and metabolic rates in a tropical livebearing fish (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>)

Sexual harassment by males has the potential to affect almost any aspect of female behavior and life history. Using Atlantic mollies (Poecilia mexicana) as a model—a species in which males do not court but almost constantly try to forcefully mate with females—we asked whether and how male harassment influences (a) females’ time budgets and (b) feeding rates (e.g., through frequent flight from male approaches), and (c) whether metabolic rates are increased as a response to stress. Field observations in a natural P. mexicana population revealed that males (average feeding rate 15%) spent far less time feeding than females (60%), and clearly traded off frequent pursuit of females with foraging. Most importantly, females’ feeding times were dramatically reduced when being pursued by a harassing male. Also in standardized lab experiments, females spent significantly less time feeding when accompanied by a male as compared to being in the presence of another female. This effect was also observed when partner fish (male or female) were presented only visually, but could not interact physically with the focal female. It seems, therefore, that females increase vigilance when a harassing male is around, which keeps them from feeding even before males actually approach them. Based on the latter result, we asked whether a stress-induced increase in metabolic rates would be discernible. We measured oxygen consumption and gill ventilation frequencies (opercular rates) of females in different social contexts (alone, with another female, or a male). The predicted, strong body mass dependency of both physiological parameters was uncovered, but no evidence for an effect of social context was detected. We argue that male harassment represents such a constant (but non-lethal) stressor for poeciliid females that their metabolic stress responses have adapted to this through habituation.     

Current indirect fitness benefits associated with philopatry in juvenile prairie voles

Summary Prairie vole (Microtus ochrogaster) family groups were examined to determine possible indirect fitness benefits from the presence of juvenile helpers at the natal nest. The reproductive performance of family groups retaining two juveniles was compared with that of families in which all juveniles were removed when the subsequent litter was born. Litter sizes at birth of the second litter and pup survival rates were the same in the two treatments. Offspring quality was affected by the presence of juveniles however. Pups reared with juveniles weighed 13% more at weaning than pups reared without juveniles. Pups also opened their eyes sooner when juveniles were present. Differences in growth and development may have been affected by the amount of time pups were alone in the nest since pups in families with juveniles were left alone less frequently than were pups without juveniles. Maternal behavior patterns were not affected by the presence of juveniles. In contrast, fathers in families with juveniles spent more time in non-parental behaviors such as feeding, drinking, and foraging. In families with large litters, mothers delivered a subsequent litter sooner if juveniles were present. Subsequent litter sizes were the same in both treatments. Overall, both infants and parents benefited from the presence of juveniles, suggesting that helping may enhance the helper's indirect fitness in multiple ways.     

Spatial and temporal patterns in the feeding behavior of a fiddler crab

Fiddler crabs consume the surficial microphytobenthos around their burrows during low tide. We studied the spatial and temporal feeding patterns in the species Uca uruguayensis by using sequences of digital pictures of feeding pellets accumulation. Data from 61 crabs, feeding without the interference of neighbors, were fitted to different models using Maximum Likelihood and Bayesian approaches. Initial feeding location was independent from the emergence location, and then, crabs continued holding a main feeding direction (clockwise or counterclockwise), suggesting a systematic mechanism that may avoid feeding over already processed sediment. Crabs used at least half of their potential feeding area, but these areas were heterogeneous. Both sexes developed similar feeding areas; however, females were faster and needed less time to feed than males, suggesting that males are time restricted. Our work also highlights the importance of incorporating other underlying mechanisms of the behavior of species into the study of feeding strategies.     

Seasonal variation in feeding behavior, competition and female social relationships in a forest dwelling guenon, the blue monkey (Cercopithecus mitis stuhlmanni), in the Kakamega Forest, Kenya

Socioecological models relate differences in feeding strategies to variation in the nature of female social relationships. Among the African forest guenons, females consume large quantities of fruit and other plant reproductive parts, resources which are thought to promote contest competition, yet these monkeys have been characterized as having agonistically undifferentiated relationships in which rank, if discernible at all, does not correlate with fitness benefits. To determine whether female relationships become more hierarchical under relevant ecological conditions, we monitored the adult females of two blue monkey groups (Cercopithecus mitis stuhlmanni) over a complete annual cycle in the Kakamega Forest, Kenya. Females competed aggressively for plant reproductive parts more often than any other resource type, and in both groups we detected linear dominance hierarchies. Nonetheless, agonism rates remained low throughout our study, and did not vary with changes in ecological conditions. Rather, when plant reproductive parts were scarce, subordinate females spent more time feeding and less time resting in an apparent attempt to compensate for a reduced efficiency of food intake. The effects of rank and food abundance were not reflected, however, in the distribution of grooming. The use of alternative feeding strategies appeared to blunt competition – females of all ranks were unlikely to be near others while feeding and spent more time consuming alternative resources when plant reproductive parts were scarce. The diverse diet of this species may allow females to avoid conflict so that dominance has only subtle effects that are difficult to detect. While socioecological models often simplify the connection between resources and female interactions, our results emphasize that the behavior of animals consuming particular resources, and not the resources themselves, are critical predictors of social patterns.     

Communal suckling in the cavy Galea musteloides

We quantified the extent of communal suckling in the cavy Galea musteloides. Six groups of animals were held in large indoor enclosures and suckling behavior was recorded over 113 h of observation. The groups contained 2–6 lactating females and 3–14 sucking pups. Due to the relative synchronization of births, 73% of the pups present in each group during lactation were non-offspring. Each of the 22 lactating females in the six groups suckled non-offspring in addition to her own offspring. On average, females suckled 86% of non-offspring present in their groups. Thus, 98% of all pups (n = 47) received milk from non-mothers. Although suckling frequencies were significantly higher for mothers with their own individual offspring than with non-offspring individuals, females invested more total time suckling all non-offspring than did suckling just their own; this was possible because for each mother many more non-offspring than offspring pups were present during lactation. Suckling bouts were significantly longer for mothers with their own individual offspring than with non-offspring individuals. The proportion of non-offspring suckling of mothers correlated negatively with the proportion of own young among the pups of a group. Non-offspring suckling did not affect future reproduction of females. Our observations demonstrate extensive practice of communal suckling in G. musteloides under laboratory conditions. Probably because all mothers of a group participated more or less equally in communal suckling behavior, the obvious cost of giving energetically expensive milk to non-offspring did not result in reduced (future) reproductive success. Potential benefits directly involved with communal suckling are unclear. More indirectly, communal suckling as well as birth synchrony might contribute to the formation of advantageous multi-litter kindergardens.     

Development of food preferences: social learning by Belding's ground squirrels Spermophilus beldingi

Summary The mechanisms by which food selection behavior develops may constrain the evolution of optimal foraging, yet these mechanisms have received relatively little attention in recent optimal foraging studies. We used cafeteria-style feeding trials to examine the role of social learning in the development of food preferences by a generalist mammalian herbivore, Spermophilus beldingi. Naive young spent relatively little time feeding and showed no preferences among five plant species offered in their feeding trials. This random pattern persisted for one set of young during two subsequent trials in the absence of their mother. A second group of young squirrels was tested initially alone, once with their mothers, and then once more alone. These squirrels fed more than those in the control group during their final trials, and showed significant preferences among plant species. These paralleled the preferences of their mothers. These results suggest that social learning may be important in the development of feeding behavior in ground squirrels, and provide a possible mechanism for cultural differences in food preferences among populations.     

Maternal investment and sex ratio adjustment in American bison calves

Summary Cow-calf behavior was observed in American bison (Bison bison) to determine if mothers invested differentially in sons and daughters. Cows nursed sons significantly longer than they did daughters in their first three months of life. The increased nursing time for sons was not compensated for by increased grazing time by their mothers. Grazing and activity patterns did not differ significantly between sons and daughters. Cows that had sons bred later in the breeding season than nulliparous cows, barren cows, or cows with daughters. Nine yearling sons compared to only two yearling daughters continued to suckle from their mothers for up to 15 months of age. Cows that had sons the previous year were more apt to be barren in the current year than cows that had daughters in the previous year.     

Changes in the activity budgets of yellow baboons (Papio cynocephalus) during sexual consortships

Summary The energetic cost of guarding estrous females has been assumed to be a potentially major factor in limiting the duration of male consortships in a number of polygamous species. This issue has been examined in the present study by assessing changes in the usual activity budgets of 13 male and 20 female yellow baboons which occurred during their consortships.The percent of time adult males spent feeding was strongly related to their agonistic rank: Higher-ranking males fed proportionately more than lower-ranking males while not in consort, and they showed a relatively greater decrease in feeding time during consortships.The proportion of time females spent feeding was related to their agonistic rank and (to a lesser degree) to their parity. Higher-ranking females fed more than lower-ranking females outside of consortships, and showed a relatively greater decrease in time spent feeding while in consort. Nulliparous females tended to feed more than parous females while not in consort; during consortships, the feeding time of nulliparous females decreased, while that of parous females increased slightly.The activity of male and female consort partners was most coordinated during the fertile cycle stage of the female partner, and least coordinated once the female had passed the fertile stage.Although reductions in feeding time during consortships were generally much more pronounced in males than in females, neither males nor females appeared to attain the undisturbed optimal feeding level for their sex while in consort.     

Infant care,infanticide, and female reproductive strategies in polygynous groups of common marmosets (Callithrix jacchus)

Reproduction in marmoset and tamarin groups is typically restricted to a single dominant female, but it is unclear why subordinate females tolerate delayed reproduction. The presence of two breeding females in free-ranging groups of common marmosets (Callithrix jacchus) presented a unique opportunity to examine differences in the reproductive strategies of dominant and subordinate females. Three groups were monitored for 12–18 months at a forest reserve in northeastern Brazil. Data on infant care were collected during two consecutive all-day follows every 10 days until the infants were 2 months old. Carrying patterns for infants born to dominant females were similar to those observed in groups containing a single breeding female. All group members over 5 months of age participated in infant care, and dominant females allowed some group members to carry their infants from the 1 st day of life. In contrast, subordinate females were protective of their offspring and were their sole caretakers for at least a week following birth. One infant born to a subordinate was killed in an attack involving the dominant female. Overall, dominant females gave birth to more infants and had higher infant survival than did subordinate breeding females. subordinate females were successful in rearing young only when the timing of births was such that they did not overlap with the dependency period of infants born to the dominant female. These patterns suggest that subordinate females may face resource competition, especially over access to helpers. In times of severe competition, subordinate females that delay reproduction may be avoiding a wasted reproductive effort.     

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.     

“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.     

The development of sex differences in ring-tailed lemur feeding ecology

Sex differences in feeding ecology may develop in response to fluctuations in physiological costs to females over their reproductive cycles, or to sexual size dimorphism, or function to minimize feeding competition within a group via resource partitioning. For most mammal species, it is unknown how these factors contribute to sex differences in feeding, or how the development of males and females reflects these intraspecific feeding differences. We show changes in dietary composition, diversity, overlap, and foraging behavior throughout development in ring-tailed lemurs (Lemur catta) and test how the development of sex differences in feeding is related to female costs of reproduction and year-round resource partitioning. Sex differences in dietary composition were only present when females were lactating, but sex differences in other aspects of feeding, including dietary diversity, and relative time spent feeding and foraging, developed at or near the time of weaning. Sex difference in juveniles and subadults, when present, were similar to the differences found in adults. The low year-round dietary overlap and early differences in dietary diversity indicate that some resource partitioning may begin with young individuals and fluctuate throughout development. The major differences between males and females in dietary composition suggest that these larger changes in diet are closely tied to female reproductive state when females must shift their diet to meet energetic and nutritional requirements.