Multilevel Societies Research Articles

Environmental Movement Interventions in Tourism and Energy Development in the North Atlantic

This article compares environmental movement engagement in energy and tourism development in Norway and Iceland by bridging the social movement societies (SMSoc) and the players and arenas perspectives. Results are based on field observation and interviews, as well as web-based textual analysis and a preliminary online survey. Results show that Norway is an institutionalized and multi-level social movement society with a mix of professionalized and grassroots local, national, and international organization. Iceland, by contrast, is a national and episodic social movement society where movement players operate at a national scale and engage in project-specific collaboration or opposition in tourism or energy development arenas. This analysis demonstrates the value of bridging the SMSoc and players and arenas perspectives for international comparative social movements research.

Diet and Activity Budget in Colobus angolensis ruwenzorii at Nabugabo, Uganda: Are They Energy Maximizers?

Introduction: Colobine monkeys are specialized folivores that use foregut fermentation to digest leaves. The slow process of fermentation forces them to spend a lot of time resting and to minimize their energy expenditure to subsist on a lower-quality diet. Methods: We recorded the diet and activity budget of Colobus angolensis ruwenzorii, which form a three-tiered multi-level society, at Lake Nabugabo, Uganda, over 12 months using scan sampling on adults and subadults, to determine whether they utilize the energy minimization strategy typical of colobines. Results: We found that the annual diet was primarily comprised of high-quality food resources (young leaves 65% and fruit 31%), and fruits were the only plant part the monkeysselected when available. Both the fruits and young leaves of some species were preferred food items in some months, and mature leaf consumption correlated negatively with preferred food availability. Mature leaves appear to be a fallback food for this population but are rarely relied upon (3%). The C. a. ruwenzoriiat Nabugabo spent less time resting (40%) and more time moving (25%) than is typical for other species of black-and-white colobus. Discussion/Conclusion: The high-quality diet of this population appears to allow them to utilize an energy maximization strategy. Their reliance on food items that tend to be clumped in space and time likely explains the frequent fission-fusion behaviour that we observe between core units. Our findings demonstrate that the foraging strategies of colobines may be more flexible than was previously thought and illustrate how food availability and distribution can impact primate social organization.

Inter-group relationships influence territorial defence in mountain gorillas.

Many species show territoriality, in which territory owners have exclusive or priority use of a region. In humans, tolerance of others within our space also depends greatly on our social relationships with them. This has been hypothesized as one potential driver of the evolution of long-term, inter-group relationships, through enabling shared access of resources and easing disputes over space. However, extremely little is known about the importance of social relationships between neighbouring groups in non-humans for how space is used and shared. Using 16years of data on the simultaneous movement and interaction patterns of 17 mountain gorilla groups, we investigated how the occurrence of aggressive and affiliative behaviour during inter-group encounters was influenced by both their social and spatial context. We found evidence of territorial defence, with rates of aggression increasing towards the centre of home ranges. Groups which had previously split from each other showed higher levels of affiliation during encounters with each other and experienced lower levels of aggression when within the other's peripheral home range. However, encounters within core areas of the home range consistently elicited higher aggression, regardless of the groups' history. Our findings indicate that not only are the social relationships between groups retained after they split from one another but also that these relationships enable groups to access certain areas with a reduced risk of aggression. This suggests that reduced aggression when accessing areas within neighbours' home ranges may be an advantage for the maintenance of inter-group relationships and a potential driver in the evolution of long-term, post-dispersal relationships and complex multi-level societies.

Male social rank and food competition in a primate multi-level society.

Social animals often have dominance hierarchies, with high rank conferring preferential access to resources. In primates, competition among males is often assumed to occur predominantly over reproductive opportunities. However, competition for food may occur during food shortages, such as in temperate species during winter. Higher-ranked males may thus gain preferential access to high-profitability food, which would enable them to spend longer engaged in activities other than feeding. We performed a field experiment with a breeding band of golden snub-nosed monkeys, a species that lives in a multi-level society in high-altitude forests in central China. We provisioned monkey's high-profitability food during winter when natural foods are limited, and then recorded the times individual adult males spent engaged in different behaviors. Higher-ranking males spent less time feeding overall and fed on provisioned foods at a higher rate than lower-ranking males. Higher-ranking males therefore had more time to spend on alternative behaviors. We found no significant difference according to rank in times spent moving or resting. However, high-ranking males spend significantly longer on affiliative behaviors with other members of their social sub-units, especially grooming and being groomed, behaviors known to promote social cohesion in primates. We show that preferential access to high-profitability foods likely relaxes time-budget constraints to higher-ranking males. High-ranking males thus spend more time on non-feeding activities, especially grooming, which may enhance social cohesion within their social sub-unit. We discuss the potential direct and indirect benefits to high-ranking males associated with preferential access to high-value food during winter.

Philopatry at the frontier: A demographically driven scenario for the evolution of multilevel societies in baboons (Papio)

The baboons (Papio sp.) exhibit marked interspecies variation in social behavior. The thesis presented here argues, first, that male philopatry is a crucial factor, arguably the crucial factor, underlying the other distinctive features (one-male units, multilevel society) shared by hamadryas and Guinea baboons, but not other species of Papio. The second suggestion is that male philopatry as a population norm was not an adaptation to a particular habitat or set of ecological circumstances but evolved in the common ancestor of hamadryas and Guinea baboons as a response to natural selection in the demographic context peculiar to the frontier of a rapidly expanding population. Other derived features of social structure (male-male tolerance, some facultative female dispersal) subsequently evolved to accommodate male philopatry. The mitochondrial genetic population structure of extant baboons preserves a footprint of the initial expansion of ‘modern’ Papio. Immediately after the expansion, male-philopatric, multilevel populations with a general physical and behavioral resemblance to Guinea baboons occupied the whole northern hemisphere range of the genus. Behavioral and physical autapomorphies of hamadryas baboons evolved in a subpopulation of this ancestral northern base, in response to a less productive habitat of the Horn of Africa. Subsequently, ancestral olive baboons ‘reinvented’ male dispersal. They and yellow baboons, another male-dispersing species, then replaced most of the male-philopatric northern populations, by male-driven introgression and nuclear swamping.

The microbiome of captive hamadryas baboons

BackgroundThe hamadryas baboon (Papio hamadryas) is a highly social primate that lives in complex multilevel societies exhibiting a wide range of group behaviors akin to humans. In contrast to the widely studied human microbiome, there is a paucity of information on the host-associated microbiomes of nonhuman primates (NHPs). Here, our goal was to understand the microbial composition throughout different body sites of cohabiting baboons.ResultsWe analyzed 170 oral, oropharyngeal, cervical, uterine, vaginal, nasal and rectal samples from 16 hamadryas baboons via 16S rRNA gene sequencing. Additionally, raw Miseq sequencing data from 1041 comparable publicly available samples from the human oral cavity, gut and vagina were reanalyzed using the same pipeline. We compared the baboon and human microbiome of the oral cavity, gut and vagina, showing that the baboon microbiome is distinct from the human. Baboon cohabitants share similar microbial profiles in their cervix, uterus, vagina, and gut. The oral cavity, gut and vagina shared more bacterial amplicon sequence variants (ASVs) in group living baboons than in humans. The shared ASVs had significantly positive correlations between most body sites, suggesting a potential bacterial exchange throughout the body. No significant differences in gut microbiome composition were detected within the maternity line and between maternity lines, suggesting that the offspring gut microbiota is shaped primarily through bacterial exchange among cohabitants. Finally, Lactobacillus was not so predominant in baboon vagina as in the human vagina but was the most abundant genus in the baboon gut.ConclusionsThis study is the first to provide comprehensive analyses of the baboon microbiota across different body sites. We contrast this to human body sites and find substantially different microbiomes. This group of cohabitating baboons generally showed higher microbial diversity and remarkable similarities between body sites than were observed in humans. These data and findings from one group of baboons can form the basis of future microbiome studies in baboons and be used as a reference in research where the microbiome is expected to impact human modeling with baboons.

The vocal repertoire of an African colobine, Colobus angolensis ruwenzorii: a multi-level society compared to congeners in stable groups

Abstract Vocal behaviour offers a window into understanding the social life and evolution of animals. Colobine monkeys show great interspecific and interpopulation variation in their social organization and behaviour. Recent research has shown that Rwenzori Angolan colobus (Colobus angolensis ruwenzorii) differ substantially from other black-and-white colobus in forming a multi-level society. No previous research has been conducted on the communication of C. a. ruwenzorii, but the social complexity hypothesis for communication suggests that more complex societies should evolve more complex communication repertoires. Our objective was to catalogue the vocal repertoire of C. a. ruwenzorii at Nabugabo, Uganda, and to compare it with the data available on congeners regarding intergroup tolerance, vocal repertoire size, and acoustic and behavioural features of vocal communication. Vocalizations were subject to spectrographic and behavioural analysis, and a descriptive analysis of each vocalization type was made. The influence of a few environmental and social factors on calling rates was also examined. We describe five vocalizations (i.e., the snort, roar, squeak, scream and pok) and one non-vocal signal (i.e., the tongue click) in this subspecies and their contexts. Distinct alarm calls are made for dogs, and these are given more often near the edge of the forest where humans frequent. We did not find that C. a. ruwenzorii showed a greater vocal repertoire than C. guereza or C. polykomos, which do not live in multi-level societies. Further, preliminary data do not indicate greater calling rates in larger core units of C. a. ruwenzorii compared to smaller units. These findings support the view that these primates’ vocalizations tend to be relatively conserved despite large differences in social organization.

Spatiotemporal association patterns in a supergroup of Rwenzori black-and-white colobus (Colobus angolensis ruwenzorii) are consistent with a multilevel society.

Primates display broad diversity in their social organization. The social groups of a few primate species are organized in a multilevel fashion, with large groups composed of multiple, core one-male units (OMUs). A characteristic of multilevel societies is that the higher levels can include hundreds of individuals. The Rwenzori black-and-white colobus (Colobus angolensis ruwenzorii) in the montane forests of Rwanda form supergroups and have been suspected to exhibit multilevel social organization. Here we present the first data on the "anatomy" of a supergroup numbering 500+ individuals. We identified subgroups within the supergroup based on progression data, extracting the social network structure from the time-stamped spatiotemporal distribution of passing individuals identified to age-sex class, and selecting an optimal time window for each network using the two-step approach developed by Uddin, Choudhury, Farhad, and Rahman (2017). We detail the existence of core units-multi-male units (MMUs) with a mean of 1.7 adult males and 3.1 adult females, as well as OMUs, all-female units and bachelor units composed of adult and sub-adult males. More than two-thirds of units are MMUs. These grouping patterns conform to a multilevel society with predominantly multi-male core units, a social system that has recently also been described for a population of the same taxon in Uganda. Individual identification will be required to corroborate these interpretations.

Anogenital distance as a measure of male competitive ability in Rwenzori Angolan colobus.

Anogenital distance (AGD) is positively correlated to fetal androgen exposure and developmental masculinization in mammals. Independent of overall body size, AGD shows a strong positive correlation with male fertility and in rodents, AGD is a good indicator of male competitive ability and is associated with female choice. We hypothesized that AGD will also predict male competitive ability in non-human primates. To test this, we measured AGD noninvasively with a parallel laser in a wild population of Angolan colobus monkeys (Colobus angolensis ruwenzorii) in Uganda and correlated to it to their social structure. C. angolensis ruwenzorii form a multilevel society with both one-male/multifemale units (OMUs) and multimale/multifemale units (MMUs). We compared AGD in males from five OMUs and six MMUs and related it to their fecal androgen metabolite concentrations, dominance rank and body size, and to the number of females in their unit. Males in OMUs had greater access to females, so were predicted to have longer AGDs, but this was not found. AGD also did not correlate overall with mean fecal androgen metabolites in MMUs. However, AGD was correlated with dominance rank in MMUs, demonstrating that higher-ranking males in these multimale units had longer AGDs. Body size did not show the same relationship with dominance rank, suggesting that male rank was not just a reflection of absolute male size. Our findings indicate that AGD predicts male competitive ability in this species and that it may be a useful correlate throughout the non-human primates. These results also support the idea that prenatal androgen exposure increases the likelihood of the expression of behaviors that maintain high dominance rank.

Bachelor groups in primate multilevel society facilitate gene flow across fragmented habitats.

In the face of ongoing habitat fragmentation, many primate species have experienced reduced gene flow resulting in a reduction of genetic diversity, population bottlenecks, and inbreeding depression, including golden snub-nosed monkeys Rhinopithecus roxellana. Golden snub-nosed monkeys live in a multilevel society composed of several 1 male harem units that aggregate to form a cohesive breeding band, which is followed by one or more bachelor groups composed of juvenile, subadult, and adult male members. In this research, we examine the continuous landscape resistance surface, the genetic diversity and patterns of gene flow among 4 isolated breeding bands and 1 all-male band in the Qinling Mountains, China. Landscape surface modeling suggested that human activities and ecological factors severely limit the movement of individuals among breeding bands. Although these conditions are expected to result in reduced gene flow, reduced genetic diversity, and an increased opportunity for a genetic bottleneck, based on population genetic analyses of 13 microsatellite loci from 188 individuals inhabiting 4 isolated breeding bands and 1 all-male band, we found high levels of genetic diversity but low levels of genetic divergence, as well as high rates of gene flow between males residing in the all-male band and each of the 4 breeding bands. Our results indicate that the movement of bachelor males across the landscape, along with their association with several different breeding bands, appears to provide a mechanism for promoting gene flows and maintaining genetic diversity that may counteract the otherwise isolating effects of habitat fragmentation.

Coda repertoire and vocal clans of sperm whales in the western Atlantic Ocean

Sperm whales live in culture-based multilevel societies in which the fundamental social level is the nearly permanent social unit of females and immature individuals, and the largest level is the clan constituted by social units that share a common coda repertoire. This study describes the coda repertoire and vocal clans of sperm whales in Brazilian waters recorded between 2011 and 2016. Inter-click intervals of codas were used to quantify similarity between repertoires. The analysis showed two evident clans, in the north and the south, distinct in their repertoires. The clan of the north, termed the “5R” clan, presented a predominant production of codas containing five regularly spaced clicks, not found in the clan of the south, termed “D” clan, which repertoire was dominated by long codas with 10–13 clicks with descending inter-click intervals. Codas with five regularly spaced clicks with the same rhythm and tempo as those found in the “5R” clan of Brazil are also predominant in sperm whales recorded in the Island of Dominica, at a distance of approximately 2700 km from the north of Brazil, which corresponds to the extension of the areas of sympatric clans. The sharing of 5R type by the both distinct clans is likely the result of cultural transmission in which conformism through social learning homogenizes coda repertoire. The results of this study corroborated the hypothesis that social boundaries, here the clan level, are maintained by cultural identities.

Microhabitat Use in Angolan Colobus Monkeys (Colobus angolensis ruwenzorii) at Nabugabo, Uganda Demonstrates Intraspecific Variability

Large-scale anthropogenic changes to the environment are making it vital to investigate habitat use so that we can better understand species’ basic ecological needs in order to create successful conservation plans. Some species are very flexible in their habitat requirements. One such species may be Colobus angolensis ruwenzorii, monkeys that occupy montane and tropical lowland forest. In the mountains of southwestern Rwanda, this species spends a great deal of time on the ground and feeds mainly on mature leaves and lichens, but the habitat use of lower altitude populations is unknown. We examined the fine-scale habitat use of C. a. ruwenzorii at Nabugabo, Uganda (1136 m altitude), where they feed primarily on young leaves and fruits. We used data collected over 9 mo (94 days between July 2017 and April 2018) on 117–131 colobus in 12 core units in a multilevel society. We found that colobus individuals were rarely on the ground, and resting and social behavior occurred at higher heights and in larger trees than feeding and moving. Individual height in the canopy followed a daily pattern (higher at the start and end, with a decrease in the middle of the day) and colobus were at lower heights when group size was greater because of core unit fusion. Finally, we found that tree species were generally used relative to their availability in the forest and their value as food species. Given greater predation risk in smaller groups and lower in the canopy at this site, our results suggest that perceived risk has an important effect on microhabitat use in C. a. ruwenzorii at lower altitude. Our results demonstrate the ecological flexibility of this primate species, which bodes well for their continued conservation.

Leopard predation on gelada monkeys at Guassa, Ethiopia.

Predation is widely believed to exert strong selective pressure on primate behavior and ecology but is difficult to study and rarely observed. In this study, we describe seven encounters between lone wild leopards (Panthera pardus) and herds of geladas (Theropithecus gelada) over a 6-year period in an intact Afroalpine grassland ecosystem at the Guassa Community Conservation Area, Ethiopia. Three encounters consisted of attempted predation on geladas by leopards, one of which was successful. All three attacks occurred in low-visibility microhabitats (dominated by tussock graminoids, mima mounds, or tall shrubs) that provided leopards with hidden viewsheds from which to ambush geladas. An additional four encounters did not result in an attempted attack but still document the consistently fearful responses of geladas to leopards. In encounters with leopards, geladas typically gave alarm calls (n = 7 of 7 encounters), reduced interindividual distances (n = 5), and collectively fled towards or remained at their sleeping cliffs (n = 7), the only significant refugia in the open-country habitat at Guassa. Geladas did not engage in mobbing behavior towards leopards. Encounters with leopards tended to occur on days when gelada herd sizes were small, raising the possibility that leopards, as ambush hunters, might stalk geladas on days when fewer eyes and ears make them less likely to be detected. We compare the behavioral responses of geladas to leopards at Guassa with those previously reported at Arsi and the Simien Mountains and discuss how gelada vulnerability and responses to leopards compare with those of other primate species living in habitats containing more refugia. Lastly, we briefly consider how living in multilevel societies may represent an adaptive response by geladas and other open-country primates to predation pressure from leopards and other large carnivores.

Multilevel societies facilitate infanticide avoidance through increased extrapair matings

In mammals characterized by a mating system in which a single male monopolizes reproduction, infanticide is reported to occur following a male take-over, often resulting in females returning to oestrus more rapidly than if their infant has survived. However, over the course of a 17-year study of golden snub-nosed monkeys, Rhinopithecus roxellana, a polygynous colobine species, we found no behavioural or suspected evidence of infanticide. Based on social network and genetic analyses, we found that more than half of the infants were sired as a result of extrapair matings (EPMs). Female golden snub-nosed monkeys initiated EPMs with bachelor or neighbouring resident males. These were considered ‘sneaky’ copulations because they did not occur in the presence of the leader male of the female's one-male unit (OMU). We suggest that through a process of paternity confusion, females decreased the likelihood of infanticide after a male take-over. In contrast, in a group of the same species in which the number of bachelor males was artificially reduced, EPMs were infrequent and four cases of infanticide were observed, three of them following a male take-over. Golden snub-nosed monkeys reside in a large multilevel society that develops by the fusion of several independent OMUs to form a breeding band, which is characterized by increased social cohesion and intrasexual tolerance among leader males. Females play an active role in mate choice, and engage in a diverse set of mating tactics, including counterstrategies such as EPMs to avoid infanticide. These results offer new insights into the diversity of behavioural strategies facilitated by complex social structures in nonhuman primates, as well as other social mammals.

Beyond the group: how food, mates, and group size influence intergroup encounters in wild bonobos

AbstractIn social-living animals, interactions between groups are frequently agonistic, but they can also be tolerant and even cooperative. Intergroup tolerance and cooperation are regarded as a crucial step in the formation of highly structured multilevel societies. Behavioral ecological theory suggests that intergroup tolerance and cooperation can emerge either when the costs of hostility outweigh the benefits of exclusive resource access or when both groups gain fitness benefits through their interactions. However, the factors promoting intergroup tolerance are still unclear due to the paucity of data on intergroup interactions in tolerant species. Here, we examine how social and ecological factors affect the onset and termination of intercommunity encounters in two neighboring communities of wild bonobos, a species exhibiting flexible patterns of intergroup interactions, at Kokolopori Bonobo Reserve, Democratic Republic of the Congo. We recorded the timing and location of intercommunity encounters and measured fruit abundance and distribution, groups’ social characteristics, and space-use dynamics over a 19-month period. We found that intercommunity tolerance was facilitated by a decrease in feeding competition, with high fruit abundance increasing the likelihood of communities to encounter, and high clumpiness of fruit patches increasing the probability to terminate encounters likely due to increased contest. In addition, the possibility for extra-community mating, as well as the potential benefits of more efficient foraging in less familiar areas, reduced the probability that the communities terminated encounters. By investigating the factors involved in shaping relationships across groups, this study contributes to our understanding of how animal sociality can extend beyond the group level.

New approaches to modeling primate socioecology: Does small female group size BEGET loyal males?

Humans are unusual in having stable male-female breeding bonds within multi-level societies. Such societies are not found in other apes, but have evolved independently in other primates, including several African papionins: hamadryas and Guinea baboons and gelada monkeys. Stable breeding bonds have been proposed to evolve either (1) because males can monopolize females when food distribution forces females to forage in small groups or (2) because females exchange exclusive mating for male services, such as protection from infanticide. Comparative studies are needed to test these hypotheses. In the meantime, we used an agent-based computer model to test the plausibility of these hypotheses. We simulated primates living in social groups within a larger population using a model we call BEGET (Behavior, Ecology, Genetics, Evolution, and Tradeoffs), which employed decision vectors, experimental evolution, realistic trade-offs, and phenotypic plasticity. We employed experimental evolution to generate male genotypes that varied in their competitive ability and in their long-term mating strategy. “Rover” males searched for and mated with any sexually receptive females whereas “Loyalist” males formed stable associations with particular groups of females. Much like living primates, the virtual primates exhibited tradeoffs between contest and scramble competition for access to females: Loyalists evolved larger body size than Rovers. We tested the effect of female foraging group size and the presence of infanticide and infant protection on the relative success of these strategies. We found that Loyalists achieved greater reproductive success than Rovers only when females were in groups smaller than four. Both Rovers and Loyalists sometimes evolved infanticidal behavior, but the presence of infanticide benefited Rovers rather than Loyalists, suggesting that the evolution of stable breeding bonds depends on the spatial distribution of females, rather than the risk of infanticide.

A multi-level society comprised of one-male and multi-male core units in an African colobine (Colobus angolensis ruwenzorii).

Several mammalian species exhibit complex, nested social organizations, termed multi-level or modular societies. Multi-level societies comprise stable core units that fission and fuse with one another in a hierarchical manner, forming groups that vary in size over time. Among nonhuman primates, these social systems have been confirmed in several African papionin and Asian colobine species. We use data from August 2017 to July 2018 on individually-recognized Rwenzori Angolan colobus living near Lake Nabugabo, Uganda to document the first multi-level society in an African colobine. The study band comprised up to 135 individuals organized into 12 socially and spatially distinct core units that ranged in size from 4 to 23 individuals. These core units showed a strong affinity to one another, spending roughly 75% of their time together. Core units fissioned and fused non-randomly with one another throughout the day, leading to different combinations of core units being observed. Using association indices between core units, we employed hierarchical cluster analyses and permutation tests to show that some core units associated preferentially into clans. Thus, we confirm three tiers of social organization for Rwenzori Angolan colobus: core unit, clan, and band. The social organization of this subspecies is unlike any reported previously in a nonhuman primate, with about half the core units containing a single adult male and the others containing multiple reproductive adult males. The discovery of a unique primate multi-level society in a novel lineage could allow for a better understanding of the evolution of these complex social systems across the Animal Kingdom. Preliminary data show males transfer within the band and females transfer outside of the band, which is proposed for hominin multi-level societies. This subspecies could thus also provide insight into the selective pressures underlying multi-level societies in our own lineage.

Social evolution in Plio-Pleistocene hominins: Insights from hamadryas baboons and paleoecology

Reconstructions of hominin evolution have long benefited from comparisons with nonhuman primates, especially baboons and chimpanzees. The hamadryas baboon (Papio hamadryas) is arguably one of the best such models, as it exhibits both the male kin bonding and the cross-sex pair bonding thought to have been important in hominin evolution. Here we link processes of behavioral evolution in hamadryas baboons with those in a Plio-Pleistocene hominin, provisionally identified as Homo erectus (sensu lato) – a pivotal species in that its larger body and brain size and wider ranging patterns increased female costs of reproduction, increasing the importance of sociality. The combination of these higher costs of reproduction and shifts in diet and food acquisition have previously been argued to have been alleviated either via strengthening of male-female bonds (involving male provisioning and the evolution of monogamy) or via the assistance of older, post-reproductive females (leading to post-reproductive longevity in females, i.e., the grandmother hypothesis). We suggest that both arrangements could have been present in Plio-Pleistocene hominins if they lived in multilevel societies. Here we expand on our earlier scenario with two sets of recent data in support of it, (1) archaeological data from the 2 million year old Oldowan site of Kanjera South, Kenya and other sites that are suggestive of tool dependent foraging on nutrient dense resources (animal carcasses and plant underground storage organs), cooperation, and food sharing; and (2) a pattern of genetic variation in hamadryas baboons that suggests the operation of kin selection among both males and females at multiple levels of society. Taken together, these two sets of data strengthen our model and support the idea of a complex society linked by male-male, male-female, and female-female bonds at multiple levels of social organization in Plio-Pleistocene hominins.

How life in a tolerant society affects the usage of grunts: evidence from female and male Guinea baboons

Signals are used to regulate interactions between individuals. To disentangle how motivational disposition, the processing of social information, and the costs and benefits of putative outcomes of interactions affect signalling behaviour, we investigated the usage and function of grunts during approaches in wild Guinea baboons, Papio papio. Guinea baboons live in a tolerant multilevel society with female-biased dispersal, which allowed us to compare their grunt usage to that of other more despotic baboon species. We analysed approaches by female and male Guinea baboons living in the Niokolo Koba National Park, Senegal. When approaching baboons grunted, they were more likely to interact in an affiliative fashion and less likely to displace the partner. In females, the probability of grunting was higher when the relationship strength was low, but only when an infant was present. In males, relationship strength had no impact on the likelihood of grunting during approaches. Rank did not explain variation in grunt probability in females and could not be discerned in males, but males were also more likely to grunt when an infant was near a female partner. We suggest that grunt usage in baboons can be best conceived as a combination of a motivational and a strategic component. The motivational component expresses the increased disposition to interact in an affiliative fashion, while the strategic component refers to the modulation of grunt usage with regard to relationship quality and context. The motivational component appears to be shared between baboon species, while variation in despotism and social organization places different premiums on the benefits of signalling, resulting in variation in grunting patterns between species.

How life in a tolerant society affects the attention to social information in baboons

An animals' ability to classify relationships between conspecifics according to kinship, affiliation and dominance constitutes an important adaptation to life in social groups. Yet, variation in the degree of competition as well as in association patterns is assumed to put differential pressure on social cognition. Because little is known about social cognition in more tolerant societies, we investigated the social knowledge of Guinea baboons, Papio papio, ranging in the Niokolo Koba National Park, Senegal. Guinea baboons live in a fluid multilevel society with ‘units’, ‘parties’ and ‘gangs’. Units comprise a ‘primary’ male, a small number of females and young. Male–female associations may last between a few weeks to several years, and females freely transfer between males at all levels of the society. Male–male relationships are characterized by high spatial tolerance and low level of competition. We examined whether males tracked changes in male–female associations at the level of the unit. We played back grunt exchanges between a female and a primary male to 25 males and recorded male responses. The grunt sequences simulated an affiliative interaction between individuals from different units (potentially indicating a female transfer to a new male, hereafter ‘inconsistent’ condition) or from individuals of the same unit (‘consistent’ condition). Surprisingly, males looked longer when exposed to the consistent compared to the inconsistent condition. In the more competitive chacma baboons, Papio ursinus, in contrast, animals responded more strongly to experimentally simulated deviations from existing social patterns. Our study suggests that the value of social information indeed varies with the degree of competition of Guinea baboons: because of female choice, little is at stake for males when a female transfers to a new male. Furthermore, the high gregariousness of the animals may favour the classification of deviant interaction patterns, at least initially, as ‘social noise’.