Fission-fusion Societies Research Articles

Chimpanzees employ context-specific behavioral strategies within fission-fusion societies.

Fission-fusion social systems allow individuals to make flexible choices about where, with whom, and in what contexts to spend their time in response to competing social and ecological pressures. The ability for fission-fusion societies to support individual behavioral strategies that vary across contexts has been suggested, but the potential function of such context-specific social choices remains largely understudied. We adopted the concept of social niche construction to explore possible differences in social complexity at the individual and group level across feeding contexts. Specifically, we examined patterns of co-attendance across two common ecological contexts in wild Central African chimpanzees (Pan troglodytes troglodytes) in the Goualougo Triangle, Republic of Congo. From data compiled over 6years, we used multidimensional social network analysis to study the patterns of co-attendance generated from 436 group scans at Ficus and 4527 visits to termite mounds. These two contexts were chosen, because they are both fixed spatial features across the landscape that serve as well-defined points to compare association patterns. We identified context-specific social niche construction in a fission-fusion chimpanzee society that produce different patterns of relationships and social complexity that are consistent in their expression over many years, and offer functional benefits. While enhancing our understanding of chimpanzee behavioral strategies, culture, and conservation, our investigation also indicates that the social niche construction framework aids in elucidating the evolutionary advantages of fission-fusion sociality by accounting for intra- and interindividual variability, cognition, and choice in newfound ways.

Sex-specific social aging in wild African lions

There is a growing interest in social behavior change with age,1,2,3,4,5 and the impacts of sociality on longevity,6,7,8 but current knowledge is broadly limited to primates, societies structured by dominance hierarchies, or single-sex studies. It is less clear how social aging patterns emerge in carnivores. The African lion (Panthera leo), a species that lives in egalitarian fission-fusion societies, presents an exceptional opportunity to examine social aging. Across felids, lions are unique in their dependence on conspecifics for many essential processes,9,10,11 and there is vast knowledge of lion behavioral ecology,10,11,12,13,14 including documented reproductive senescence in both sexes.14,15 Applying spatial-social network analyses across 30 years of data on the wild Serengeti lion population, we show that sex strongly modulates patterns of social aging and longevity. Group size increased with age for both sexes, but only males experienced significant changes in associate numbers (degree), specifically to females, which peaked in mid-life before declining. While aging females experienced declines in intra-sex connectivity (strength) and bond strength (mean strength), they peaked in both to males during mid-life. Male inter-sex strength also peaked in mid-life, while conversely their intra-sex strength and mean strength significantly dipped in mid-life. Although social associations were important for survival in both sexes, the investment diverged significantly: females' overall network connectivity was key for longevity, while the number of associates was important for males. These findings illustrate important potential effects of social aging in a wild carnivore and demonstrate how these diverge strongly between the sexes.

Effect of ecological and anthropogenic factors on grouping patterns in African lions across Kenya.

Social carnivores frequently live in fission-fusion societies, where individuals that share a common territory or home range may be found alone, in subgroups, or altogether. Absolute group size and subgroup size is expected to vary according to resource distribution, but for species that are susceptible to anthropogenic pressures, other factors may be important drivers. African lions (Panthera leo) are the only truly social felid and lion prides are characterized by fission-fusion dynamics with social groups frequently splitting and reforming, and subgroup membership can change continuously and frequently. The number of individuals in a group can be reflective of social, ecological, and anthropogenic conditions. This dynamic behavior makes understanding lion grouping patterns crucial for tailoring conservation measures. The evolution of group living in lions has been the topic of numerous studies, and we drew on these to formulate hypotheses relating to group size and subgroup size variation. Based on data collected from 199 lion groups across eight sites in Kenya, we found that group sizes were smaller when lions were closer to human settlements, suggesting that edge effects are impacting lions at a national scale. Smaller groups were also more likely when they were far from water, and were associated with very low and very high levels of non-tree vegetation. We found significant differences between the study sites, with the Maasai Mara having the largest groups (mean ± SD = 7.7 ± 4.7, range = 1-19), and Amboseli conservation area the smallest (4.3 ± 3.5, range = 1-14). While long-term studies within a single site are well suited to thoroughly differentiate between absolute group size and subgroup size, our study provides unique insight into the correlates of grouping patterns in a vulnerable species at a national scale.

Functional convergence of genomic and transcriptomic architecture underlies schooling behaviour in a live-bearing fish.

The organization and coordination of fish schools provide a valuable model to investigate the genetic architecture of affiliative behaviours and dissect the mechanisms underlying social behaviours and personalities. Here we used replicate guppy selection lines that vary in schooling propensity and combine quantitative genetics with genomic and transcriptomic analyses to investigate the genetic basis of sociability phenotypes. We show that consistent with findings in collective motion patterns, experimental evolution of schooling propensity increased the sociability of female, but not male, guppies when swimming with unfamiliar conspecifics. This finding highlights a relevant link between coordinated motion and sociability for species forming fission-fusion societies in which both group size and the type of social interactions are dynamic across space and time. We further show that alignment and attraction, the two major traits forming the sociability personality axis in this species, showed heritability estimates at the upper end of the range previously described for social behaviours, with important variation across sexes. The results from both Pool-seq and RNA-seq data indicated that genes involved in neuron migration and synaptic function were instrumental in the evolution of sociability, highlighting a crucial role of glutamatergic synaptic function and calcium-dependent signalling processes in the evolution of schooling.

Accelerometer-based predictions of behaviour elucidate factors affecting the daily activity patterns of spotted hyenas.

Animal activity patterns are highly variable and influenced by internal and external factors, including social processes. Quantifying activity patterns in natural settings can be challenging, as it is difficult to monitor animals over long time periods. Here, we developed and validated a machine-learning-based classifier to identify behavioural states from accelerometer data of wild spotted hyenas (Crocuta crocuta), social carnivores that live in large fission-fusion societies. By combining this classifier with continuous collar-based accelerometer data from five hyenas, we generated a complete record of activity patterns over more than one month. We used these continuous behavioural sequences to investigate how past activity, individual idiosyncrasies, and social synchronization influence hyena activity patterns. We found that hyenas exhibit characteristic crepuscular-nocturnal daily activity patterns. Time spent active was independent of activity level on previous days, suggesting that hyenas do not show activity compensation. We also found limited evidence for an effect of individual identity on activity, and showed that pairs of hyenas who synchronized their activity patterns must have spent more time together. This study sheds light on the patterns and drivers of activity in spotted hyena societies, and also provides a useful tool for quantifying behavioural sequences from accelerometer data.

Volume of foliage tents affects harem size in Cynopterus sphinx living in a fission-fusion society

Volume of foliage tents affects harem size in Cynopterus sphinx living in a fission-fusion society

Fission-fusion dynamics in the social networks of a North American pitviper.

Many animal species exist in fission-fusion societies, where the size and composition of conspecific groups change spatially and temporally. To help investigate such phenomena, social network analysis (SNA) has emerged as a powerful conceptual and analytical framework for assessing patterns of interconnectedness and quantifying group-level interactions. We leveraged behavioral observations via radiotelemetry and genotypic data from a long-term (>10 years) study on the pitviper Crotalus atrox (western diamondback rattlesnake) and used SNA to quantify the first robust demonstration of social network structures for any free-living snake. Group-level interactions among adults in this population resulted in structurally modular networks (i.e., distinct clusters of interacting individuals) for fidelis use of communal winter dens (denning network), mating behaviors (pairing network), and offspring production (parentage network). Although the structure of each network was similar, the size and composition of groups varied among them. Specifically, adults associated with moderately sized social groups at winter dens but often engaged in reproductive behaviors-both at and away from dens-with different and fewer partners. Additionally, modules formed by individuals in the pairing network were frequently different from those in the parentage network, likely due to multiple mating, long-term sperm storage by females, and resultant multiple paternity. Further evidence for fission-fusion dynamics exhibited by this population-interactions were rare when snakes were dispersing to and traversing their spring-summer home ranges (to which individuals show high fidelity), despite ample opportunities to associate with numerous conspecifics that had highly overlapping ranges. Taken together, we show that long-term datasets incorporating SNA with spatial and genetic information provide robust and unique insights to understanding the social structure of cryptic taxa that are understudied.

Factors affecting captive female giraffe stress response: Male presence, small enclosure, and low temperature.

To improve animal welfare based on suitable social housing conditions, it is important to understand the factors that trigger high-stress responses. Wild giraffes live in a fission-fusion society and males and females are rarely in the same herd for a long period. The captive condition of belonging to a herd with the same individuals for months or years is uncommon in nature. To understand the effect of male presence on female stress levels, fecal glucocorticoid metabolite (fGCM) levels and social interactions in two captive female giraffes were investigated. Additionally, the effect of enclosure size and temperature on fGCM level and social interactions were examined. The results showed no significant difference in the fGCM levels of females based on male presence. The frequency of agonistic behavior by the dominant female toward the subordinate female was significantly increased when a male was present. The subordinate female was significantly less likely to approach the dominant female and showed decreased affiliative and agonistic interactions toward the dominant female when a male was present. The frequencies of agonistic interactions between females were higher in the small enclosure regardless of male presence. Low temperature triggered higher fGCM levels and increased agonistic interaction in an aged female. The findings of this study suggest that these multiple factors should be considered individually to promote the welfare of captive giraffes.

Cooperation increases bottlenose dolphins' (Tursiops truncatus) social affiliation.

Dolphins live in a fission-fusion society, where strong social bonds and alliances can last for decades. However, the mechanism that allows dolphins to form such strong social bonds is still unclear. Here, we hypothesized the existence of a positive feedback mechanism in which social affiliation promotes dolphins' cooperation, which in turn promotes their social affiliation. To test it, we stimulated the cooperation of the 11 dolphins studied by providing a cooperative enrichment tool based on a rope-pulling task to access a resource. Then we measured the social affiliation [simple ratio index (SRI)] of each possible pair of dolphins and evaluated whether it increased after cooperation. We also evaluated whether, before cooperation, pairs that cooperated had a higher SRI than those that did not cooperate. Our findings showed that the 11 cooperating pairs had significantly stronger social affiliation before cooperation than the 15 non-cooperating pairs. Furthermore, cooperating pairs significantly increased their social affiliation after cooperation, while non-cooperating pairs did not. As a result, our findings provide support to our hypothesis, and suggest that the previous social affiliation between dolphins facilitates cooperation, which in turn promotes their social affiliation.

Bonobo optimizer algorithm for optimum design of truss structures with static constraints

In this study, a very recently developed intelligent algorithm called Bonobo Optimizer (BO) algorithm is implemented for the sizing optimization of the truss structures with discrete and continuous variables. The BO algorithm imitates the social behaviour and reproductive schemes of the bonobos. Like the several other primates, the bonobos also follow the fission–fusion group strategy, where they form several small parties of various sizes and move separately in their territory. The sizing optimization searches for the minimum weight of the truss structures subject to the multiple loading conditions under the required constraints on the member stresses and nodal displacements. Five well-known truss examples with fixed-geometry and up to 160 elements are tested to verify the reliability and the robustness of the BO algorithm. The results clearly indicate that the BO algorithm is a powerful search and optimization technique for the truss structures imposed by the static constraints for the discrete and continuous variables.

Long-Term Assessment of Captive Chimpanzees: Influence of Social Group Composition, Seasonality and Biographic Background.

Wild chimpanzees (Pan troglodytes) live in flexible fission-fusion societies with frequent changes in both group size and composition. These changes depend mostly on resource availability and individual social preferences yet in captivity are determined by housing organizations. During a period of seven years, we observed a group of sanctuary chimpanzees, focusing on how they adapted to changes in the group composition over time. Using linear mixed models (LMMs), factors such as group size, sex ratio, seasonality, and the individuals' sex and origin (wild caught vs. captive born) were considered in order to evaluate the impact on the chimpanzees' activity levels, the occurrence of undesired behaviors (abnormal and self-directed behaviors) and the social grooming networks. Our results indicate that the activity levels and the occurrence of undesired behaviors were impacted by changes in group composition and the individuals' biographic background. The colder season was marked by higher levels of activity and more time spent grooming. Moreover, it was the individuals' origin but not changes in group composition that affected social grooming, with wild-caught chimpanzees grooming far less frequently. Long-term observations are essential to evaluate, predict and detect potential benefits and/or issues of housing conditions while considering the social and physical environment.

Network analysis of a northern long-eared bat (Myotis septentrionalis) maternity colony in a suburban forest patch

Abstract Many bat species are highly social, forming groups of conspecifics, particularly during the maternity season. In temperate North America, these social groups are typically comprised of closely related individuals or individuals that share some common trait (i.e. reproductive state or shared hibernacula from the previous winter). In the summer, when bats use forests for day-roosts, these social groups often demonstrate nonrandom patterns of periodically associating in common roosts and disassociating using different roosts as a ‘fission–fusion society’. As cave hibernating bat species in North America continue to decline due to the impacts of White-nose Syndrome, opportunities to describe these dynamics are becoming rare. Unfortunately, these patterns often are still poorly documented, yet understanding these behaviors is critical for species-specific habitat conservation and management. In our study, we tracked female northern long-eared bats (Myotis septentrionalis) to their day-roosts in a small, suburban forest fragment in coastal New York, USA, in the summers of 2018 and 2019. We confirmed that the bats shared roost sites and, using network analyses, analyzed social dynamics and space use. In contrast to previous research on this imperiled species in large, unfragmented core forests, we found a more dense, connected roost network that concentrated around forest patch edges. Unusual for this species, primary roosts were anthropogenic structures. Our findings suggest that northern long-eared bats can utilize small forest patches and that incorporation of specific types of anthropogenic roosts might be an effective strategy for long-term conservation in more urbanized landscapes where forest management actions to enhance day-roosting conditions are impractical and the risk of stochastic loss of roosts is high.

A pachyderm perfume: odour encodes identity and group membership in African elephants

Group-living animals that live in complex social systems require effective modes of communication to maintain social cohesion, and several acoustic, olfactory and visual signaling systems have been described. Individuals need to discriminate between in- and out-group odour to both avoid inbreeding and to identify recipients for reciprocal behaviour. The presence of a unique group odour, identified in several social mammals, is a proposed mechanism whereby conspecifics can distinguish group from non-group members. African elephants (Loxodonta africana) live in stable, socially complex, multi-female, fission–fusion groups, characterized by female philopatry, male dispersal and linear dominance hierarchies. Elephant social behaviour suggests that individuals use odour to monitor the sex, reproductive status, location, health, identity and social status of conspecifics. To date, it is not clear what fixed or variable information is contained in African elephant secretions, and whether odour encodes kinship or group membership information. Here we use SPME GC–MS generated semiochemical profiles for temporal, buccal and genital secretions for 113 wild African elephants and test their relationship with measures of genetic relatedness. Our results reveal the existence of individual identity odour profiles in African elephants as well as a signature for age encoded in temporal gland and buccal secretions. Olfactory signatures for genetic relatedness were found in labial secretions of adult sisters. While group odour was not correlated with group genetic relatedness, our analysis identified “group membership” as a significant factor explaining chemical differences between social groups. Saturated and short-chain fatty acids (SCFAs), derived from key volatile compounds from bacterial fermentation, were identified in temporal, buccal and genital secretions suggesting that group odour in African elephants may be the result of bacterial elements of the gut microbiome. The frequent affiliative behavior of African elephants is posited as a likely mechanism for bacterial transmission. Our findings favour flexible group-specific bacterial odours, which have already been proposed for other social mammals and present a useful form of olfactory communication that promotes bond group cohesion among non-relatives in fission–fusion mammals.

Strategic intergroup alliances increase access to a contested resource in male bottlenose dolphins

Efforts to understand human social evolution rely largely on comparisons with nonhuman primates. However, a population of bottlenose dolphins in Shark Bay, Western Australia, combines a chimpanzee-like fission-fusion grouping pattern, mating system, and life history with the only nonhuman example of strategic multilevel male alliances. Unrelated male dolphins form three alliance levels, or "orders", in competition over females: both within-group alliances (i.e., first- and second-order) and between-group alliances (third-order), based on cooperation between two or more second-order alliances against other groups. Both sexes navigate an open society with a continuous mosaic of overlapping home ranges. Here, we use comprehensive association and consortship data to examine fine-scale alliance relationships among 121 adult males. This analysis reveals the largest nonhuman alliance network known, with highly differentiated relationships among individuals. Each male is connected, directly or indirectly, to every other male, including direct connections with adult males outside of their three-level alliance network. We further show that the duration with which males consort females is dependent upon being well connected with third-order allies, independently of the effect of their second-order alliance connections, i.e., alliances between groups increase access to a contested resource, thereby increasing reproductive success. Models of human social evolution traditionally link intergroup alliances to other divergent human traits, such as pair bonds, but our study reveals that intergroup male alliances can arise directly from a chimpanzee-like, promiscuous mating system without one-male units, pair bonds, or male parental care.

Sound Visualization Demonstrates Velopharyngeal Coupling and Complex Spectral Variability in Asian Elephants.

Simple SummaryWhether a vocalization is emitted though the mouth or nose impacts on its acoustic shape through specific vocal tract resonances. In human language, nasalized compared to oral vowels can change the meaning of words. African elephants mainly use low-frequency rumbles to communicate and can utter them either through the mouth or trunk. In this study, we used an acoustic camera to visualize the sound emission of rumbles in Asian elephants, which have been studied considerably less than African elephants. We recorded nine captive female Asian elephants and analyzed the acoustic structure of 203 calls. We found that most rumbles (64%) were emitted through the trunk, 21% through the mouth, and surprisingly, 13% simultaneously through the mouth and trunk. Some of the rumbles were combined with orally emitted roars. The nasal rumbles concentrated most spectral energy in lower frequencies, whereas the oral and mixed rumbles showed a broader spectral energy distribution and concentration in higher frequencies, and were louder. The roars were the loudest, broadest and highest in frequency. This study is the first to demonstrate coupled oral and nasal emission in a non–human animal, thereby setting an important framework to study the functions of this acoustic variability in elephant communication and the evolution of vocal flexibility across species.Sound production mechanisms set the parameter space available for transmitting biologically relevant information in vocal signals. Low–frequency rumbles play a crucial role in coordinating social interactions in elephants’ complex fission–fusion societies. By emitting rumbles through either the oral or the three-times longer nasal vocal tract, African elephants alter their spectral shape significantly. In this study, we used an acoustic camera to visualize the sound emission of rumbles in Asian elephants, which have received far less research attention than African elephants. We recorded nine adult captive females and analyzed the spectral parameters of 203 calls, including vocal tract resonances (formants). We found that the majority of rumbles (64%) were nasally emitted, 21% orally, and 13% simultaneously through the mouth and trunk, demonstrating velopharyngeal coupling. Some of the rumbles were combined with orally emitted roars. The nasal rumbles concentrated most spectral energy in lower frequencies exhibiting two formants, whereas the oral and mixed rumbles contained higher formants, higher spectral energy concentrations and were louder. The roars were the loudest, highest and broadest in frequency. This study is the first to demonstrate velopharyngeal coupling in a non-human animal. Our findings provide a foundation for future research into the adaptive functions of the elephant acoustic variability for information coding, localizability or sound transmission, as well as vocal flexibility across species.

The partial merger of two dolphin societies.

Interactions between mammalian social groups are generally antagonistic as individuals in groups cooperate to defend resources from non-members. Members of the family Delphinidae inhabit a three-dimensional habitat where resource defence is usually impractical. Here, we describe a long-term partial fusion of two communities of Atlantic spotted dolphins (Stenella frontalis). The northern community, studied for 30 years, immigrated 160 km to the range of the southern community, observed for 20 years. Both communities featured fission–fusion grouping patterns, strongest associations between adult males, and frequent affiliative contact between individuals. For the 5-year period following the immigration, we found members of all age classes and both sexes in mixed groups, but there was a strong bias toward finding immigrant males in mixed groups. Some association levels between males, and males and females, from different communities were as high as the highest within-community associations. Affiliative contacts indicate that these individuals were forming social relationships. The mixing of two separate social groups with new bond formation is rare in terrestrial mammal groups. Such mixing between spotted dolphin groups suggests that adaptations to respond aggressively to ‘outsiders’ are diminished in this species and possibly other ecologically similar dolphins.

Cascading indirect genetic effects in a clonal vertebrate.

Understanding how individual differences arise and how their effects propagate through groups are fundamental issues in biology. Individual differences can arise from indirect genetic effects (IGE): genetically based variation in the conspecifics with which an individual interacts. Using a clonal species, the Amazon molly (Poecilia formosa), we test the hypothesis that IGE can propagate to influence phenotypes of the individuals that do not experience them firsthand. We tested this by exposing genetically identical Amazon mollies to conspecific social partners of different clonal lineages, and then moving these focal individuals to new social groups in which they were the only member to have experienced the IGE. We found that genetically different social environments resulted in the focal animals experiencing different levels of aggression, and that these IGE carried over into new social groups to influence the behaviour of naive individuals. These data reveal that IGE can cascade beyond the individuals that experience them. Opportunity for cascading IGE is ubiquitous, especially in species with long-distance dispersal or fission-fusion group dynamics. Cascades could amplify (or mitigate) the effects of IGE on trait variation and on evolutionary trajectories. Expansion of the IGE framework to include cascading and other types of carry-over effects will therefore improve understanding of individual variation and social evolution and allow more accurate prediction of population response to changing environments.

Olfactory Inspection of Female Reproductive States in Chimpanzees

In mammalian species, olfactory cues are important for within and between species communication. These cues can be part of multimodal signals indicating, for example, female fertility potentially perceived by male conspecifics. However, a large gap exists in our understanding of multimodal signaling in non-human catarrhines. Chimpanzees (Pan troglodytes) live in fission-fusion societies, mate promiscuously and express high levels of male-male competition. Females show a striking visual signal, an exaggerated sexual swelling, known to be a proxy of ovulation, while the maximum swelling is not matching the exact time of ovulation. The question remains if males use additional olfactory cues when being able to approach females closely. This would allow males to pinpoint the exact timing of ovulation and could be one reason why high-ranking males sire offspring more successfully than other males. Here, we present the first systematic test of such multimodal signaling by investigating male sniffing behavior directed to females in relation to their fertility in a group of 13 captive chimpanzees. Our results show that male sniffing behavior significantly increased with female swelling size, with female age as well as when fewer male competitors were present. Hence, odors might be part of a multimodal fertility cue, supporting the idea that males monitor both visual and olfactory cues to gain comprehensive information on female fertility.

Demographic characteristics shape patterns of dawn swarming during roost switching in tree-dwelling Daubenton’s bat

Frequent roost switching in fission–fusion societies of tree-dwelling bats is closely associated with swarming behaviour entailing ritualised night-time displays around the roost tree and/or at the roost entrance to signal its actual location, particularly immediately prior to sunrise. However, effects of demographic characteristics of individuals in this social behaviour remain unanswered. Using passive integrated transponders (PIT) and automatic readers, we recorded swarming activity of members of a Daubenton’s bat (Myotis daubentonii) maternity colony in the vicinity of their roosts. In total, 59,622 activity events of 281 PIT-tagged individuals were recorded on ten monitored roosts during three summer seasons. We found a gradual increase of swarming activity from midnight to sunrise in old adult females, whereas young females and juveniles primarily swarmed later at dawn. We attribute this difference to the learning status of younger bats, which are not yet able to perform a defined pattern of swarming activity, whereas older bats likely take a more active role in signalling the position of the roost. Old males exhibited the least swarming activity at maternity roosts, which mostly occurred between crepuscular periods, presumably due to their solitary lives. A negative correlation between genetic distance and swarming activity suggests an important role of kinship in the formation of the maternity colony as well as group cohesion during roost switching.

Runaway Social Selection in Human Evolution

Darwin posited that social competition among conspecifics could be a powerful selective pressure. Alexander proposed a model of human evolution involving a runaway process of social competition based on Darwin’s insight. Here we briefly review Alexander’s logic, and then expand upon his model by elucidating six core arenas of social selection that involve runaway, positive-feedback processes, and that were likely involved in the evolution of the remarkable combination of adaptations in humans. We discuss how these ideas fit with the hypothesis that a key life history innovation that opened the door to runaway social selection, and cumulative culture, during hominin evolution was increased cooperation among individuals in small fission-fusion groups.