Function of Notification in Papio hamadryas

Notification is a form of ritualized greeting behavior performed by adult male hamadryas baboons (Papio hamadryas). Originally observed in the wild when harem leaders signaled the direction of travel to their followers on daily foraging marches, its true function is unknown. Notification involves the approach of an adult male baboon to another male, the presentation of his hindquarters, then retreat. Researchers have previously linked notification to aggression avoidance, appeasement, and the dominance-subordination relationship between males. We observed instances of notification over 6 mo among 8 adult males in a troop of 61 captive-bred hamadryas baboons at Paignton Zoo Environmental Park, Devon, UK. We compared observation sessions in which notification occurred with matched control periods. We classified different forms of notification according to the recipient’s response: presenting, mounting, touching of the buttocks or genitals, or ignoring. Results suggest that different forms of notification may have different functions, including submission, alliance, and peacekeeping but not general greeting, recruitment, or appeasement.     

Testicular Size,Mating System,and Maturation Schedules in Wild Anubis and Hamadryas Baboons

We report body mass and testicular size in 258 anubis (Papio anubis or P. hamadryas anubis) and 59 hamadryas (P. hamadryas or P. h. hamadryas) baboons, live-trapped in Ethiopia. As predicted by theories of sexual selection by sperm competition, among hamadryas baboons, which are monandrous, fully adult males have absolutely and relatively smaller testes than those of comparable males among anubis baboons, which are polyandrous. Male hamadryas are also ca. 10% smaller in bodily mass as adults. The intertaxonal difference in adults is due entirely to the fact that in male anubis baboons, testicular and bodily mass continue to grow up to full adulthood–the age at which most males emigrate from their natal troop and initiate a confrontational breeding strategy among unrelated animals. By contrast, male hamadryas baboons, which are usually philopatric, attain adult body mass and testicular size as subadults. In both species, juveniles experience rapid testicular growth peaking in rate at ca. 12kg body mass, but testicular descent and growth starts earlier in hamadryas than in anubis baboons. Juvenile hamadryas baboons have relatively larger testes than their anubis equivalents, perhaps because male philopatry allows the mating strategy of male hamadryas baboons to be initiated during juvenile life and therefore permits some sperm competition between juveniles and adults.     

Dominance relationships among male hamadryas baboons (Papio hamadryas)

The concept of dominance has been shown to be useful in describing and predicting social interactions in group-living animals. However, the dominance relationships among adult hamadryas males (Papio hamadryas) are poorly understood, and systematic data are missing from the literature. This study investigated dominance relationships among male hamadryas baboons by applying novel statistical techniques to systematic detailed data on agonistic interactions. We also analyzed the dominance relationship with male age and evaluated the association between dominance and access to mating partners (i.e. the number of adult females per one-male unit (OMU)), food resources (i.e. monopolization of feeding areas), and greeting interactions. The derived dominance indexes showed that, in general, leader males were dominant over followers, and that dominance ability did not correlate with male age. Individual dominance values were very close to each other, suggesting that dominance relationships among hamadryas males were not very rigid. In addition, dominance values were positively correlated with number of adult females per OMU but not with feeding priority. Finally, greeting interactions occurred more frequently between individuals with similar dominance values and with low levels of dominance decidedness. We suggest the need of further studies, especially in wild populations, to confirm our findings.     

Population genetic insights into the social organization of Guinea baboons (Papio papio): Evidence for female‐biased dispersal

Sex differences in philopatry and dispersal have important consequences on the genetic structure of populations, social groups, and social relationships within groups. Among mammals, male dispersal and female philopatry are most common and closely related taxa typically exhibit similar dispersal patterns. However, among four well‐studied species of baboons, only hamadryas baboons exhibit female dispersal, thus differing from their congenerics, which show female philopatry and close‐knit female social relationships. Until recently, knowledge of the Guinea baboon social system and dispersal pattern remained sparse. Previous observations suggested that the high degree of tolerance observed among male Guinea baboons could be due to kinship. This led us to hypothesize that this species exhibits male philopatry and female dispersal, conforming to the hamadryas pattern. We genotyped 165 individuals from five localities in the Niokolo‐Koba National Park, Senegal, at 14 autosomal microsatellite loci and sequenced a fragment of the mitochondrial hypervariable region I (HVRI) of 55 individuals. We found evidence for higher population structuring in males than in females, as expected if males are the more philopatric sex. A comparison of relatedness between male–male and female–female dyads within and among communities did not yield conclusive results. HVRI diversity within communities was high and did not differ between the sexes, also suggesting female gene flow. Our study is the first comprehensive analysis of the genetic population structure in Guinea baboons and provides evidence for female‐biased dispersal in this species. In conjunction with their multilevel social organization, this finding parallels the observations for human hunter‐gatherers and strengthens baboons as an intriguing model to elucidate the processes that shaped the highly cooperative societies of Homo. Am. J. Primatol. 77:878–889, 2015. © 2015 The Authors. American Journal of Primatology Published by Wiley Periodicals Inc.     

The Influence of Social Systems on Patterns of Mitochondrial DNA Variation in Baboons

Behavior is influenced by genes but can also shape the genetic structure of natural populations. Investigating this link is of great importance because behavioral processes can alter the genetic diversity on which selection acts. Gene flow is one of the main determinants of the genetic structure of a population and dispersal is the behavior that mediates gene flow. Baboons (genus Papio) are among the most intensely studied primate species and serve as a model system to investigate the evolution of social systems using a comparative approach. The general mammalian pattern of male dispersal and female philopatry has thus far been found in baboons, with the exception of hamadryas baboons (Papio hamadryas). As yet, the lack of data on Guinea baboons (Papio papio) creates a taxonomic gap in genus-wide comparative analyses. In our study we investigated the sex-biased dispersal pattern of Guinea baboons in comparison to hamadryas, olive, yellow, and chacma baboons using sequences of the maternally transmitted mitochondrial hypervariable region I. Analyzing whole-range georeferenced samples (N = 777), we found strong evidence for female-biased gene flow in Guinea baboons and confirmed this pattern for hamadryas baboons, as shown by a lack of genetic-geographic structuring. In addition, most genetic variation was found within and not among demes, in sharp contrast to the pattern observed in matrilocal primates including the other baboon taxa. Our results corroborate the notion that the Guinea baboons’ social system shares some important features with that of hamadryas baboons, suggesting similar evolutionary forces have acted to distinguish them from all other baboons.     

A Papionin Multilevel Society as a Model for Hominin Social Evolution

Multilevel societies are unique in their ability to facilitate the maintenance of strong and consistent social bonds among some individuals while allowing separation among others, which may be especially important when social and sexual bonds carry significant and reliable benefits to individuals within social groups. Here we examine the importance of social and sexual bonds in the multilevel society of hamadryas baboons (Papio hamadryas) and apply these principles to social evolution in Plio-Pleistocene hominins. The behavior, adaptations, and socioecology of baboons (Papio spp.) have long been recognized as providing an important comparative sample to elucidate the processes of human evolution, and the social system of hamadryas baboons in particular shares even more similarities with humans than that of other baboons. Here we draw parallels between processes during the evolution of hamadryas social organization and those characterizing late Pliocene or early Pleistocene hominins, most likely Homo erectus. The higher costs of reproduction faced by female Homo erectus, exacerbated by an increased reliance on difficult to acquire, nutrient-dense foods, are commonly thought to have been alleviated by a strengthening of male–female bonds (via male provisioning and the evolution of monogamy) or by the assistance of older, postreproductive females (via grandmothering). We suggest that both of these social arrangements could have been present in Plio-Pleistocene hominins if we assume the development of a multilevel society such as that in hamadryas baboons. The evolution of a multilevel society thus underlies the adaptive potential for the complexity that we see in modern human social organization.     

Group housing of hamadryas baboons: a new cage design based upon field studies of social organization

The hamadryas subspecies, which is utilized extensively for laboratory research, has a social structure unique among baboons. Field studies have shown that small one male units are the basic grouping in a complex multi-level social system. A housing design for a research colony of hamadryas baboons was conceived to reflect this unique social organization and related behavioral patterns. The outcome of combining such ethological considerations with practical dictates for housing large laboratory primates has been assessed in terms of reproductive performance, animal health and utilization for research. Various benefits of this approach to cage design for hamadryas baboons are identified in this report.     

Greeting,aggression, and coalitions between male baboons: Demographic correlates

Interactions of asymmetrical, symmetrical, and unreciprocated greeting, and episodes of aggression and coalitions observed between 18 males, members of a large multi-harem colony of baboons (Papio hamadryas, P. cynocephalus, and their hybrids) housed in the Madrid Zoo, were recorded throughout 11 time blocks between January 1983 and March 1985. The major question investigated was whether the demographic characteristics of the colony affected the frequency with which males used three behavioural strategies, namely, greeting, aggression, and coalitions. It was found that under demographic conditions eliciting more intense male-male competition, for example, when the ratio of oestrous cycles per adult male was lower, males exchanged frequent greetings, especially the symmetrical form (i.e. notifying) and solicited more coalitions, but did not show any significant change in rates of dyadic aggression. Male phenotype-related effects on rates of greeting and coalitions were found as well. Results are discussed in relation with the suggested role of greeting and of coalition behaviour as strategies used in the negotiation of changes in competitive social relationship (e.g. consort changeovers). These strategies are regarded as social tactics alternative to direct aggression.     

Early sexual maturity in male hamadryas baboons (Papio hamadryas hamadryas) and its reproductive implications

We present data on sexual maturity in young hamadryas baboon males (Papio hamadryas hamadryas) and its reproductive consequences in a large captive baboon colony. Hamadryas baboons live in a multilevel social system, with one-male units (OMUs) as the smallest social entity. Male leaders of OMUs are believed to monopolize matings within their OMUs; hence mating is believed to be polygynous and monandrous. In a captive colony of hamadryas baboons, we found evidence that young males less than 4 years old fathered at least 2.5% of 121 offspring born subsequent to vasectomy of all adult males, and males aged 4-5 years fathered at least 16.5% of the offspring. Additional evidence that these young males are able to sire offspring came from a morphological comparison of sperm from hamadryas males of different ages. The sperm of a 48-month-old hamadryas baboon were morphologically indistinguishable from viable sperm from adult males, whereas sperm from a 45-month-old male showed some aberrations. If successful copulations by adolescent males constitute a regular pattern even in free-ranging hamadryas baboons, a hamadryas male's chances to reproduce would not be limited to his role as an OMU leader as previously assumed, and a male's reproductive career would consist of two phases: the adolescent phase, and the OMU leader male phase.     

Affiliation Among Females in Wild Hamadryas Baboons (Papio hamadryas hamadryas)

Previous researchers of hamadryas baboons have described a star-shaped sociogram, whereby the strongest social bonds within hamadryas one-male units are between a leader male and his females and bonds among females are weak by comparison. This type of social organization is also known as cross-bonding to distinguish it from the female-bonding found in most papionin monkeys. Models of female primate socioecology suggest that hamadryas baboons lack female bonding due to their reliance on scarce, widely-dispersed food resources. Here, I report observational data from a wild population of hamadryas baboons in Ethiopia indicating that, while females varied widely in their frequency of social interaction with other females, most females spent about as much social time with other females as they did with the leader male and some females even crossed unit boundaries to interact with one another. The size of a unit was positively correlated with the tendency of its females to interact with other females and was negatively correlated with the tendency of its females to interact with the leader male. Females were equally likely to spend social time with other females whether or not the leader male was available for social interaction at the time. Overall, this study suggests that a star-shaped sociogram does not characterize all hamadryas baboons and that female hamadryas may be, to some extent, female-bonded as well as cross-bonded. The lack of more pronounced female bonding in hamadryas is probably due to the behavior of males rather than to ecological factors.     

Social relationships of mountain baboons: Leadership and affiliation in a non-female-bonded monkey

Instead of close and differentiated relationship among adult females, the accepted norm for savanna baboons, groups of Drakensberg mountain baboons (Papio ursinus) showed strong affiliation of females towards a single male. The same male was usually the decision-making animal in controlling group movements. Lactating or pregnant females focused their grooming on this “leader” male, producing a radially patterned sociogram, as in the desert baboon (P. hamadryas); the leader male supported young animals in the group against aggression and protected them against external threats. Unlike typical savanna baboons, these mountain baboons rarely displayed approach-retreat or triadic interactions, and entirely lacked coalitions among adult females. Both groups studied were reproductively one-male; male-female relationships in one were like those in a unit of hamadryas male at his peak, while the other group resembled the unit of an old hamadryas male, who still led the group, with a male follower starting to build up a new unit and already monoplizing mating. In their mountain environment, where the low population density suggests conditions as harsh for baboons as in deserts, adults in these groups kept unusually large distances apart during ranging; kin tended to range apart, and spacing of adults was greatest at the end of the dry, winter season. These facts support the hypothesis that sparse food is responsible for convergence with hamadryas social organization. It is suggested that all baboons, though matrilocal, are better categorized as “cross-sex-bonded” than “female bonded”.     

Dental eruption schedules of wild and captive baboons

Dental eruption schedules previously used to age wild baboons have in the past derived from studies of captive animals housed under standard conditions and fed standard laboratory diets. This paper reports for the first time eruption schedules derived from wild baboons, the yellow baboons (Papio hamadryas cynocephalus) of Mikumi National Park, Tanzania, and compares these schedules with those of other baboon subspecies inhabiting both similar and dissimilar environments. Eighteen males and twelve females from the Viramba groups, ranging in age from 21 to 103 months, were trapped, and dental impressions and notes were made of the state of eruption of each tooth. Eruption of all teeth were delayed at Mikumi relative to the baboon standards derived from the captive animals at the Southwest Foundation for Biomedical Research, San Antonio, Texas. Teeth of the canine-premolar 3 complex and third molars were most delayed, erupting up to a year and a half later than their counterparts from captive animals. Comparison with data on hamadryas baboons from Erer-Gota in Ethiopia revealed that both the hamadryas and yellow subspecies of baboons, with different genetic backgrounds and living under markedly different environmental conditions, followed the same schedule. This constancy of developmental schedules suggests that these Mikumi data may reasonably be used as standards for other wild baboon populations and that acceleration of dental maturation, as well us maturation of other somatic systems in captivity, is another manifestation of the short-term adaptive plasticity of the baboon species as a whole.     

Group Composition of Guinea Baboons (<Emphasis Type="Italic">Papio papio</Emphasis>) at a Water Place Suggests a Fluid Social Organization

Baboon social systems are among the most studied in primates. Solid knowledge of the hamadryas and savannah baboon systems has accumulated, leading to a dichotomic view of baboon social systems. Hamadryas baboons live in multilayered troops based on 1-male units whereas savannah baboons live in multimale multifemale groups based on a network of related females. Less attention has been paid to their West African congenerics, the Guinea baboons, Papio papio. To fill this gap, in 2007 we initiated a long-term study of a baboon troop ranging in the Niokolo Koba National Park in southeastern Senegal. Earlier studies suggested a tendency for a multilayered social system in Guinea baboons, similar to the hamadryas baboon organization. Therefore, as a first approach to analyzing variability in party size and composition, we observed members of the troop crossing an open area from a fixed point for 3 mo during the dry and wet seasons. We counted individuals and recorded changes in composition of both arriving and departing parties. Party size and composition were highly variable on both a daily and a seasonal basis; 45.9% of the arriving parties changed in composition while crossing the open area, either splitting into smaller parties or fusing into larger ones, suggesting a fluid organization. Our data support the existence of neither a hamadryas baboon-like multilayered social organization nor a stable medium-sized multimale multifemale group as in savannah baboons. In light of our data we may need to revise the dichotomic view of baboon social systems and include space for greater variability of their social systems.     

One-male harems and female social dynamics in Guinea baboons

Little is known about the mating system and social organization of Guinea baboons. This study investigated whether Guinea baboons have a harem-based mating system similar to that of hamadryas and gelada baboons and whether one-male mating units also correspond to social units. Ten adult females in a captive multi-male multi-female group of Guinea baboons were focally observed 2 h per week for 12 weeks, and all observed copulations within the group were recorded. Some males copulated with a single female while others had harems of 2-4 females. All females copulated with a single male except 1 female that switched harems early in the study. The focal females had higher rates of social interaction with their harem members, especially their harem male, than with individuals outside the harem. Females appeared to be subordinate to the harem male but little or no physical aggression or herding behavior from the male was observed. Variation in female social interactions within the harem was not accounted for by their sexual interactions with the male or their genetic relatedness with the females. Females, however, appeared to maintain social relationships with their female relatives in other harems. Taken together, the results of this study show that both mating and affiliative interactions in Guinea baboons are concentrated within one-male units and that the social dynamics within and between these units share some similarities as well as differences with those of hamadryas and gelada baboons.     

Blood groups of baboons. Population genetics of feral animals

Blood and saliva from 495 Ethiopan baboons were collected in the field and tested for their human-type A-B-O groups while 493 blood samples were tested for their simian-type blood factors A^p, B^p, C^p, G^p, N^p, ca and hu. Four series of feral animals were tested: 194 olive baboons, a troop of 82 and another of 90 hamadryas baboons, and a series of 129 baboons classified as olive/hamadryas hybrids. In addition, 126 baboons from other sources were tested for their human-type A-B-O groups and 131 for their simian-type blood groups. Human-type groups A, B and AB but not group O were found in combined series of 621 animals. Gene frequency analysis also indicated the absence of group O. Population analysis of the data obtained for the 493 Ethiopian baboons has shown that the simian-type blood groups A^p and B^p are independent of one another. In contrast B^p and G^p appear to be determined by corresponding allelic genes; if confirmed by population data on additional series of animals, this would define the first baboon blood group system found. There is a close association between the blood group specificities ca and hu, the exact nature of which still remains to be clarified. Blood group ca, originally believed to be species specific, is found to be polymorphic in both olive baboons and hamadryas as well as in the hybrids; hu, on the other hand, present in all hamadryas tested, is polymorphic only in olive baboons.     

Growth and hormone characteristics of pubertal development in the hamadryas baboon

Abstract: The semi-longitudinal collection of growth measurements in male and female hamadryas baboons has enabled documentation of the timing of puberty and the development of sexually dimorphic growth patterns in body weight, crown-rump length (CRL), limb lengths, and muscle mass. In addition, another sexually dimorphic characteristic appears to be the presence of a pubertal growth spurt in body weight, and possibly CRL, in male but not female baboons. Serum testosterone levels rose during male development; however, there was a progressive decrease in dehydroepiandrosterone sulfate levels indicating the absence of adrenarche. Insulin-like growth factor-I (IGF-I) and its major binding protein, IGFBP-3, both rose during pubertal development; however, a simultaneous rise in the IGF-I:IGFBP-3 molar ratio suggests other factors may enhance the bioactivity of IGF-I during puberty. A distinct rise in serum osteocalcin levels was also associated with puberty in male baboons. These growth and hormonal changes during puberty in the hamadryas baboon indicate that this species provides a close primate model for human puberty.     

Migration of male hamadryas baboons into anubis groups in the Awash National Park,Ethiopia

Among “savanna” baboons, males are the dispersing sex, and females are philopatric. Despite clear evidence for migration of adult males at Erer-Gota, Ethiopia (Abegglen, 1984), it is generally believed that a different pattern-dispersal only by female transfer-is found in hamadryas baboons,Papio hamadryas hamadryas (Pusey and Packer, 1987; Pusey, 1988; Stammbach, 1987). Since the late 1960's, there have been isolated observations of hamadryas males migrating into anubis groups in the Awash National Park, Ethiopia (Nagel, 1973; Kawai and Sugawara, 1976; Sugawara, 1982). Since 1983, we have observed 11 individually identified adult hamadryas immigrants in four anubis groups above the Awash Falls and have trapped and tagged 9 of them. One subadult male was also captured and marked. Repeated visits to the study site allow us to document long-term residence of these “cross-migrant” males in their host groups. The longest-resident male has been in the same group for 5 years or more; a conservative estimate of the average length of residence is 3 years. We estimate that 25 hamadryas males have moved into this ozne over the last 15 years. Although larger than the hamadryas males captured in 1973, all but one of our cross-migrants appear phenotypically hamadryas. By comparing, the ages of our cross-migrants with Abegglen's account of the typical hamadryas male life-history, we have found that the adult hamadryas males seem to immigrate at ages consistent with having left their     

Group Composition and Adult Sex-ratio of Hamadryas Baboons (Papio hamadryas hamadryas) in Central Eritrea

During a survey of the geographical distribution and abundance of hamadryas baboons (Papio hamadryas hamadryas) in central Eritrea, we collected detailed demographic data on six bands at four sites in different ecogeographical zones. The proportions of age-sex classes within the six bands differed only with respect to juveniles. The general social organization of the Eritrean hamadryas baboons is similar to that reported for Ethiopia and Saudi Arabia. Eritrean hamadryas baboons live in a nested fission-fusion system, with one-male units as the basic social entity. Although the baboons were not provisioned, as they are in many places in Saudi Arabia, and habitat quality in Eritrea is lower than that in Ethiopia, sex-ratios and group composition corresponded more to those found in the Saudi Arabian population. Sex-ratios within the study population, in bands and also in one-male units were significantly more female-biased than in Ethiopian ones, and one-male units tended to be larger. Data from Eritrea suggest that these differences are due to a combination of a heavily fluctuating rainfall pattern and differential maturation of the sexes.     

Mitochondrial phylogeny and systematics of baboons (Papio)

Baboons (Papio, s.s.) comprise a series of parapatric allotaxa (subspecies or closely related species) widely distributed in sub-Saharan Africa. Despite extensive studies of their ecology, morphology, and behavior, disagreement about their phylogenetic relationships continues, as expressed in the current coexistence of at least three major, competing taxonomic treatments. To help resolve this situation, we sequenced approximately 900 bases of mitochondrial DNA of 40 individuals from five of the widely recognized "major" allotaxa. Total sequence diversity (>5%) is high compared to most primate species. Major mitochondrial clades correspond to recognized allotaxa, with the important exception that haplotypes from yellow and olive baboons form a single, monophyletic clade within which the two allotaxa do not comprise mutually exclusive clusters. The major clades fall unambiguously into the pattern: (chacma (Guinea (hamadryas (yellow + olive)))). This phylogeny does not support taxonomies that oppose hamadryas to all other baboons ("desert" vs. "savanna"), but is compatible with the view that all definable allotaxa should be recognized as coordinates, either as "phylogenetic" species or "biological" subspecies. The close relationship and unsegregated distribution of haplotypes from Kenyan and Tanzanian yellow and olive baboons are unexplained, but may reflect introgression across the documented hybrid zone. The overall phylogeny, when combined with paleontological data, suggests a southern African origin for extant Papio baboons, with all extant lineages sharing a common mitochondrial ancestor at approximately 1.8 Ma.     

Life history of hamadryas baboons: Physical development,infant mortality,reproductive parameters and family relationships

Demographic and life history parameters were estimated for a band of free-ranging hamadryas baboons, observed for 5.5 years in Ethiopia. Age-related changes in body weight and dentition were found to be delayed relative to laboratory-reared baboons. On the average, females reached menarche at 4.3 years of age and had their first infant at the age of 6.1 years. The mean interbirth interval was 24 months if the infant survived this period. The survival of infants and juveniles was higher compared to Amboseli yellow baboons, but somewhat lower than in gelada baboons in the Simen Mountains. Males acquired their first juvenile or adult female at the age of 8.5 to 11 years. Male-female pair-bonds lasted several years in most cases. The Cone Rock baboons were organized in a four-level social structure. The troop could split into bands, bands were divided into clans, and clans into one-male units with bachelor followers. The exchange of individuals between social units predominantly occurred within the band. All males of known origin became adult members of their presumed natal clan. Most females transferred also within clans, and juvenile females tended to remain in their natal clan. Females lost by one male to several rivals tended to reassemble in the same new one-male units later on.