Group Size In Primates Research Articles

Human group size puzzle: why it is odd that we live in large societies.

Human groups tend to be much larger than those of non-human primates. This is a puzzle. When ecological factors do not limit primate group size, the problem of coordination creates an upper threshold even when cooperation is guaranteed. This paper offers a model of group coordination towards behavioural synchrony to spell out the mechanics of group size limits, and thus shows why it is odd that humans live in large societies. The findings suggest that many of our species' evolved social behaviours and culturally maintained social technologies emerged as solutions to this problem.

'Dunbar's number' deconstructed.

A widespread and popular belief posits that humans possess a cognitive capacity that is limited to keeping track of and maintaining stable relationships with approximately 150 people. This influential number, ‘Dunbar's number’, originates from an extrapolation of a regression line describing the relationship between relative neocortex size and group size in primates. Here, we test if there is statistical support for this idea. Our analyses on complementary datasets using different methods yield wildly different numbers. Bayesian and generalized least-squares phylogenetic methods generate approximations of average group sizes between 69–109 and 16–42, respectively. However, enormous 95% confidence intervals (4–520 and 2–336, respectively) imply that specifying any one number is futile. A cognitive limit on human group size cannot be derived in this manner.

The social organization of Homo ergaster: Inferences from anti-predator responses in extant primates

Patterns of primate socioecology have been used to suggest that the first truly savanna-dwelling hominin, Homo ergaster, lived in sizeable groups. Here, we revisit these estimates and infer additional features of the social organization of these early hominins based on anti-predator responses observed across the primate taxon. We first show that the effect of habitat on primate group size, composition, and sexual dimorphism is negligible after controlling for substrate use and phylogeny: terrestrial species live in larger groups with more and bigger males than arboreal taxa. We next hypothesize that groups can only survive in open habitats if males are able to engage in joint counter-attacks against the large carnivorans typical of such environments. To test this, we analyze reports on primate counter-attacks against known predators and find these are indeed disproportionately frequent in terrestrial taxa living in open habitats, sometimes even involving the use of tentative weapons. If we subsequently only examine the taxa that are particularly adept at this (chimpanzees and baboons), we find an effect of habitat type on group size: groups on the savanna are larger than those in the forest. We thus infer that H. ergaster lived in very large groups with many males that jointly defended the group against carnivorans, and argue that these counter-attacks will readily have turned into confrontational scavenging and cooperative hunting, allowing Homo to move into the niche of social carnivore. These two features (life in very large multi-male groups and a switch to persistent carnivory) shaped the evolution of our lineage to such an extent that the social organization of H. ergaster may already have contained many key elements characterizing modern day foragers: male bonding, incipient male–female friendships with food sharing, a tendency toward endogamy, and the presence of large communities that eventually turned into the ethno-linguistic units we can still recognize today.

Assessing sources of error in comparative analyses of primate behavior: Intraspecific variation in group size and the social brain hypothesis

Phylogenetic comparative methods have become standard for investigating evolutionary hypotheses, including in studies of human evolution. While these methods account for the non-independence of trait data due to phylogeny, they often fail to consider intraspecific variation, which may lead to biased or erroneous results. We assessed the degree to which intraspecific variation impacts the results of comparative analyses by investigating the “social brain” hypothesis, which has provided a framework for explaining complex cognition and large brains in humans. This hypothesis suggests that group life imposes a cognitive challenge, with species living in larger social groups having comparably larger neocortex ratios than those living in smaller groups. Primates, however, vary considerably in group size within species, a fact that has been ignored in previous analyses. When within-species variation in group size is high, the common practice of using a mean value to represent the species may be inappropriate. We conducted regression and resampling analyses to ascertain whether the relationship between neocortex ratio and group size across primate species persists after controlling for within-species variation in group size. We found that in a sample of 23 primates, 70% of the variation in group size was due to between-species variation. Controlling for within-species variation in group size did not affect the results of phylogenetic analyses, which continued to show a positive relationship between neocortex ratio and group size. Analyses restricted to non-monogamous primates revealed considerable intraspecific variation in group size, but the positive association between neocortex ratio and group size remained even after controlling for within-species variation in group size. Our findings suggest that the relationship between neocortex size and group size in primates is robust. In addition, our methods and associated computer code provide a way to assess and account for intraspecific variation in other comparative analyses of primate evolution.

High dietary diversity supports large group size in Indo-Chinese gray langurs in Wuliangshan, Yunnan, China.

Identifying ecological factors underlying primate group size has been a central theme in primate behavioral ecology. The ecological constraints model proposes that increased group size leads to enhanced within-group feeding competition, necessitating increased travel to encounter additional or more productive feeding sites. Over the course of three years, we studied the largest known group (>80 individuals) of Trachypithecus crepusculus (Indo-Chinese gray langur) for 1738 hours during 213 days (including 96 full day follows) in Wuliangshan, China. During this period, group size increased from 81 individuals to over 90 individuals. The group consumed plant parts from 27 ± 8 (range: 15-51) species per month, and a total 148 plant species during the study. Based on time spent feeding, the diet was similar (54.2% leaves and 32.1% fruit and seeds) to that reported for other colobines. Despite occupying a home range several times larger than other groups of Trachypithecus, we found no evidence of an increase in daily path length with increasing group size, and the group's time budget and daily path length remained relatively constant despite marked monthly changes in dietary pattern (e.g. time spent exploiting fruit and seeds vs. buds and young leaves, vs. mature leaves). These results are inconsistent with many of the predictions of the ecological constraints model. Rather, our data suggest that high dietary diversity and the ability to exploit difficult to digest foods enabled Indo-Chinese gray langurs to reduce opportunities for feeding competition associated with increased group size.

Data Quality and the Comparative Method: The Case of Primate Group Size

The comparative method is frequently employed to study primate behavior and evolution. The method is used to infer adaptations, and considerable improvements have been made with respect to its implementation. Despite these advances, scant attention has been given to the nature of the data that are used in comparative analyses. This creates a potential problem as data are often compiled from studies conducted by multiple researchers, whose methods may differ, resulting in variation in data quality. In this article, we investigate the quality of data employed in studies of primate group size. Several issues concerning data quality arise when assembling data on group size. For example, data quality may be compromised if group sizes are estimated from censuses, unhabituated groups, or groups with unrecognized individuals. To mitigate these and other data quality issues, we gathered data from the literature on 23 monkeys and apes using well-defined and biologically relevant criteria for inclusion. We compare our results with those of eight published compilations of group size. Most studies did not provide details regarding the criteria for including data. We found that our group size values were uncorrelated or weakly correlated with those from three other studies and differed in a consistent fashion from those of one other study. Because conclusions derived from comparative analyses are only as accurate as the data that they use, future studies should provide details regarding data collection to ensure their reliability.

Unlike fellows – a review of primate–non‐primate associations

Throughout many regions of the tropics, non-primate animals - mainly birds and mammals - have been observed to follow primate groups and to exploit dropped food and flushed prey. The anecdotal nature of most of the numerous reports on these primate-non-primate associations (PNPAs) may obscure the biological significance of such associations. We review the existing literature and test predictions concerning the influence of primate traits (body size, activity patterns, dietary strategies, habitat, group size) on the occurrence of PNPAs. Furthermore, we examine the influence of non-primates' dietary strategies on the occurrence of PNPAs, and the distribution of benefits and costs. We detected a strong signal in the geographic distribution of PNPAs, with a larger number of such associations in the Neotropics compared to Africa and Asia. Madagascar lacks PNPAs altogether. Primate body size, activity patterns, habitat and dietary strategies as well as non-primate dietary strategies affect the occurrence of PNPAs, while primate group size did not play a role. Benefits are asymmetrically distributed and mainly accrue to non-primates. They consist of foraging benefits through the consumption of dropped leaves and fruits and flushed prey, and anti-predation benefits through eavesdropping on primate alarm calls and vigilance. Where quantitative information is available, it has been shown that benefits for non-primates can be substantial. The majority of PNPAs can thus be categorized as cases of commensalism, while mutualism is very rare. Our review provides evidence that the ecological function of primates extends beyond their manifold interactions with plants, but may remain underestimated.

Fission–Fusion Behavior in Yunnan Snub-Nosed Monkeys (Rhinopithecus bieti) in Yunnan, China

Several species of mammals live in relatively large social groups that temporarily fission or subgroup in response to changes in food availability, predation risk, foraging strategies, and mating competition. Although the dynamics of subgrouping are not well understood, evidence of facultative fission–fusion behavior in species that generally exhibit a highly cohesive group structure may help to clarify the set of ecological and social factors that constrain or affect group size in primates. We here examined patterns of subgrouping in Yunnan snub-nosed monkeys (Rhinopithecus bieti) inhabiting the Baimaxueshan National Nature Reserve in Yunnan, China. Rhinopithecus bieti lives in a large multilevel society consisting of 6–41 one-male, multifemale units (OMUs) and one all-male unit (AMU). Over a 5-yr period from 2003 through 2008, we tracked a group of >450 Rhinopithecus bieti. We recorded the location in the group’s home range where fission and fusion events occurred, the size and number of subgroups, days spent in subgroups, and food availability, and monitored predator sightings. The results indicate that the focal group underwent12 group fission events, all of which occurred between mid-June and July, and all in the same area of the group’s range. During these fissioning events, the AMU also showed subgrouping behavior. In all cases, the presence of bamboo shoots, an important seasonal component of the snub-nosed monkey diet, appeared to trigger subgrouping behavior. The subgroups reunited in other parts of their range after a period of 2–11 d. We found no evidence that the presence of predators had a direct affect on subgrouping behavior in Rhinopithecus bieti, as proposed in previous studies.

Within‐group social bonds in white‐faced saki monkeys (Pithecia pithecia) display male–female pair preference

White-faced saki monkeys (Pithecia pithecia) lack most of the behavioral and physical traits typical of primate monogamy [Fuentes, 1999]. In order to determine if social bonds in this species reflect patterns displayed by pair-bonded groups or larger multimale-multifemale groups, we draw on 17 months of data collected on wild white-faced sakis at Brownsberg Nature Park, Suriname. We analyzed within-group social bonds for three habituated groups (one two-adult and two multiadult groups) by measuring grooming, proximity, and approach/leave patterns between adult and subadult group members. We found that both two-adult and multiadult groups showed significantly stronger social bonds between a single male-female dyad within each group (deemed "primary dyads"). In all three groups, primary dyads were composed of the oldest adult male and a breeding female. These pairs had significantly higher levels of grooming than other within-group dyads and were also in close proximity (<1 m) more often than nonprimary dyads. Grooming in primary dyads was nonreciprocal, and consistently biased toward female investment. Grooming patterns in nonprimary dyads varied, but were often more reciprocal. Grooming and proximity of the primary dyad also changed in relation to infant development. Our results suggest that while white-faced sakis do not show behavioral and physical traits typical of monogamy or pair-bonding, social bonds are strongest between a single male-female pair. Pitheciine social systems range from small group monogamy in Callicebus to large multimale-multifemale groups in Chiropotes and Cacajao. As the middle taxon in this platyrrhine radiation, behavioral strategies of white-faced sakis provide a model for how social bonds and affiliation could be influenced by and affect the evolution of larger group size in primates.

Two girls for every boy: The effects of group size and composition on the reproductive success of male and female white‐faced capuchins

Many factors have been hypothesized to affect the size and adult sex ratios of primate groups and these, in turn, have been argued to influence birth rates. Using park-wide census data collected on a population of capuchins over a 25-year period, we examined whether group size and adult sex ratio affect the per capita reproductive success of male and female white-faced capuchins (Cebus capucinus) in Santa Rosa National Park, Costa Rica. We found that the reproductive success of females (measured as the observed minus the expected ratio of immatures to adult females in the group) decreased with increasing group size, whereas that of males was independent of group size. The proportion of adult males residing in groups had significant, yet contrasting effects on males and females. Male reproductive success was negatively associated with the proportion of males residing in groups whereas female reproductive success increased with the proportion of males. The latter finding supports the intersexual conflict hypothesis, which suggests that a conflict of interest occurs between males and females over adult sex ratios. The effects of group size and composition on the reproductive success of capuchins, a male-dispersed omnivorous species, are similar to those reported for howlers, a bisexually-dispersed folivorous species. One common factor between these taxa is that groups with low ratios of males to females are at greater risk of takeovers and resultant infanticide. Our results suggest that regardless of dietary preference and dispersal pattern, the threat of infanticide can constrain primate group size and composition.

Network cohesion, group size and neocortex size in female-bonded Old World primates

Most primates are intensely social and spend a large amount of time servicing social relationships. In this study, we use social network analysis to examine the relationship between primate group size, total brain size, neocortex ratio and several social network metrics concerned with network cohesion. Using female grooming networks from a number of Old World monkey species, we found that neocortex size was a better predictor of network characteristics than endocranial volumes. We further found that when we controlled for group size, neocortex ratio was negatively correlated with network density, connectivity, relative clan size and proportional clan membership, while there was no effect of neocortex ratio on change in connectivity following the removal of the most central female in the network. Thus, in species with larger neocortex ratios, females generally live in more fragmented networks, belong to smaller grooming clans and are members of relatively fewer clans despite living in a closely bonded group. However, even though groups are more fragmented to begin with among species with larger neocortices, the removal of the most central individual does cause groups to fall apart, suggesting that social complexity may ultimately involve the management of highly fragmented social groups while at the same time maintaining overall social cohesion. These results emphasize a need for more detailed brain data on a wider sample of primate species.

PL.02.01 Has serotonin a role in the development of the social brain?

Phylogenetic comparative methods have become standard for investigating evolutionary hypotheses, including in studies of human evolution. While these methods account for the non-independence of trait data due to phylogeny, they often fail to consider intraspecific variation, which may lead to biased or erroneous results. We assessed the degree to which intraspecific variation impacts the results of comparative analyses by investigating the “social brain” hypothesis, which has provided a framework for explaining complex cognition and large brains in humans. This hypothesis suggests that group life imposes a cognitive challenge, with species living in larger social groups having comparably larger neocortex ratios than those living in smaller groups. Primates, however, vary considerably in group size within species, a fact that has been ignored in previous analyses. When within-species variation in group size is high, the common practice of using a mean value to represent the species may be inappropriate. We conducted regression and resampling analyses to ascertain whether the relationship between neocortex ratio and group size across primate species persists after controlling for within-species variation in group size. We found that in a sample of 23 primates, 70% of the variation in group size was due to between-species variation. Controlling for within-species variation in group size did not affect the results of phylogenetic analyses, which continued to show a positive relationship between neocortex ratio and group size. Analyses restricted to non-monogamous primates revealed considerable intraspecific variation in group size, but the positive association between neocortex ratio and group size remained even after controlling for within-species variation in group size. Our findings suggest that the relationship between neocortex size and group size in primates is robust. In addition, our methods and associated computer code provide a way to assess and account for intraspecific variation in other comparative analyses of primate evolution.

Intersexual Conflict and Group Size in Alouatta palliata: A 23-year Evaluation

Models of optimal primate group size suggest that group formation and growth arise to benefit individual fitness, but that size is limited by costs. The ecological constraints hypothesis posits that group formation and growth is driven by protection from predation or the advantages of group foraging, while an upper limit on group size is constrained by travel costs and intragroup competition for food or other critical resources. Socioecological models also predict that individual reproductive success, hypothesized to decrease with increasing group size, also places an upper limit on the number of individuals in a group. Our analysis of 23 yr of group composition data on mantled howlers (Alouatta palliata) from a single Panamanian study site on Barro Colorado Island not only corroborates the socioecological model but also shows that female reproductive success increased, whereas that of males decreased, with the less female-biased sex ratios in larger groups. We suggest that the conflict of interest between the sexes over adult sex ratio, particularly the male proportion in a group, in combination with ecological factors, is an important determinant of group size and composition.

Primate Group Size and Abundance in the Caatinga Dry Forest, Northeastern Brazil

The Caatinga dry forest poses a series of ecological challenges for mammals in general and primates in particular. The erratic rainfall pattern impacts on plant diversity and phenological patterns; from year to year there is marked variability in fruit production and failure to fruit is common. The harshness apparently accounts for the impoverished mammalian fauna. However, data on primate abundance, distribution, and possible environmental effects on primate density are lacking in this type of dry forest. I censused the primate community in 3 habitats of the Serra da Capivara National Park, Piaui, NE Brazil, over a total distance of 318 km. Overall, the abundance of primates in the Caatinga dry forest is very low as a consequence of low abundance of food resources both in space and time. Alouatta caraya (predominantly folivorous) occurs at extremely low density, and during the dry season are apparently confined to canyon areas, where trees retain their leaves. Callithrix jacchus has morphological feeding specializations for gum-eating, and gum is an important resource during food bottleneck periods. Nonetheless, Callithrix jacchus occurs at comparatively low densities. Group sizes for howlers and marmosets in the Caatinga are significantly smaller than in other forest types. Contrarily, Cebus apella libidinosus had an average group size within the range reported for Amazonian and Atlantic forests. Researchers consider the generalized diet of capuchins as the explanation for their similar abundance in different habitats, indicating relative independence from ecological constraints. However, I suggest that capuchin foraging style and cognitive abilities are important factors accounting for their unreduced group size and density even under extreme conditions.

Primate group size and interpreting socioecological models: Do folivores really play by different rules?

Because primates display such remarkable diversity, they are an ideal taxon within which to examine the evolutionary significance of group living and the ecological factors responsible for variation in social organization. However, as with any social vertebrate, the ecological determinants of primate social variability are not easily identified. Interspecific variation in group size and social organization results from the compromises required to accommodate the associative and dissociative forces of many factors, including predation,1–3 conspecific harassment and infanticide,4–6 foraging competition1, 7 and cooperation,8 dominance interactions,9 reproductive strategies, and socialization.10–12 Causative explanations have emerged primarily through the construction of theoretical models that organize the observed variation in primate social organization and group size relative to measurable ecological variation.

Feeding Competition and Group Size in Alouatta pigra

Researchers consider group size in primates to be determined by complex relationships among numerous ecological forces. Antipredator benefits and better resource defense are the primary pressures for large groups. Conversely, intragroup limited food availability, can result in greater intragroup feeding competition and individual energy expenditure in larger groups, creating energetic advantages for individuals in small groups and placing an upper limit group size. However, the extent to which food availability constrains group size remains unclear for many species, including black howlers (Alouatta pigra), which ubiquitously live in small social groups (≤10 individuals). We studied the relationship between group size and 2 key indices of feeding competition—day journey length and activity budgets—in 3 groups of wild Alouatta pigra at a hurricane-damaged site in Belize, Central America. We controlled for differences in food availability between home ranges (food tree density) and compared both indicators of feeding competition directly with temporal variation in food availability for each group. Our results show no consistent association between resource availability, group size, and either index of competition, indicating that feeding competition does not limit group size at the site—i.e., that larger groups can form without increased costs of feeding competition. The results support the search for other explanations, possibly social ones, for small group size in the primates, and we conclude with suggestions and evidence for such alternative explanations.

A unimodal curve for primate group size along a gradient of population density

AbstractWhen plotted along a gradient of population density, the mean group size in populations of several primate species has a unimodal distribution, i.e., mean group size is greater at intermediate population densities than at higher or lower population densities. In this study I present a mathematical model to clarify the cause of this relationship. Population density is assumed to affect group size by enhancing between‐ or within‐group competition and by changing the number of neighboring groups around each group. The mean group size is predicted to decline as population density increases above a critical value, owing to the increasing number of neighboring groups.

Neocortex Size Predicts Group Size in Carnivores and Some Insectivores

AbstractNeocortex size has been shown to correlate with group size in primates. Data for carnivores and insectivores are used to test the generality of this relationship. The data suggest that carnivores lie on the same grade as the primates, but that insectivores lie on a separate grade to the left of these two orders. Among the insectivores, there appears to be a distinction between the ‘advanced’ genera (which show a relationship between group size and neocortex size) and the ‘basal’ genera (which do not).

Brain-Behavior Relations in Primates and Cetaceans: Implications for the Ubiquity of Factors Leading to the Evolution of Complex Intelligence

AbstractThis paper describes the results of a project aimed at addressing questions about the evolution of intelligence by utilizing comparisons of brain-behavior relations in cetaceans (dolphins and whales) and primates. Three specific questions were asked. First, have any cetaceans reached a hominid level of encephalization? Second, are cetacean-primate comparisons of brain organization useful for grappling with the differences that are bound to exist between any extraterrestrial organism and ourselves? Third, are there similarities in brain-behavior relations in cetaceans and primates that suggest behavioral evolution is shaped by general factors? Brain and body weights for cetacean and primate species were collected and compared with each other and with data on brain organization and social behavior. The results revealed that a hominid level of encephalization is not unique in mammalian history. Furthermore, cetacean-primate comparisons can be useful in understanding the different ways in which information processing systems can be organized. Finally, a comparison of the relation between encephalization and social group size in primates and cetaceans preliminarily suggests that similar mental capacities can be achieved through different independent evolutionary paths and that there may be factors common to behavioral evolution in all intelligent organisms.

Neocortical size and language

AbstractIn my target article, I argued (1) that the relationship between neocortical size and group size in primates implies that there is a cognitive limit on the size of human groups, and (2) that time constraints forced the evolution of language as a more efficient means of bonding the large groups that humans evolved. The doubts about these claims raised by these additional commentaries largely reflect misinterpretation of my original claims.