Social Complexity Hypothesis Research Articles

Complexity in Chick‐a‐Dee Calls of Mountain Chickadees (Poecile gambeli): Call Variation Associated With Flock Size and Flight

ABSTRACTThe chick‐a‐dee call of chickadees, tits, and titmice is a vocal system used in a wide range of social contexts by both sexes throughout the year and is one of the more structurally complicated vocal systems outside of human language. Relatively little is known about the chick‐a‐dee calls of mountain chickadees, Poecile gambeli, however. This is an important species for increasing our comparative understanding of variation in chick‐a‐dee calls as they are one of the chickadee species with the largest naturally occurring flock sizes. Flock size relates to the social complexity of flocks, and the social complexity hypothesis for communication predicts that individuals in more complex social groups should communicate with greater complexity than individuals in simpler social groups. Correlational and experimental evidence in support of the hypothesis has been found in the calls of a wide range of species, including Carolina chickadees, P. carolinensis. Here, we provide the first description of the variation in note composition and note‐ordering rules in calls from mountain chickadee flocks in California and Colorado. California flocks were found to be significantly larger than Colorado flocks. Analysis of note‐type usage and transition probabilities between note types found that calls of California birds were more complex than calls of Colorado birds, supporting a key prediction of the social complexity hypothesis for communication. We also found relatively high rates of reversals of note‐ordering rules in mountain chickadee calls, which might help explain the complexity of the chick‐a‐dee calls of this species. Additionally, birds in flight produced calls with different note compositions when compared to perched birds. Generally, the note‐type ordering and transition probabilities of calls of mountain chickadees seem comparable to other better‐studied chickadee species, although their frequent note‐type order rule reversals suggest potential syntax‐like properties in this call system.

Social Brain Perspectives on the Social and Evolutionary Neuroscience of Human Language.

Human language and social cognition are two key disciplines that have traditionally been studied as separate domains. Nonetheless, an emerging view suggests an alternative perspective. Drawing on the theoretical underpinnings of the social brain hypothesis (thesis of the evolution of brain size and intelligence), the social complexity hypothesis (thesis of the evolution of communication), and empirical research from comparative animal behavior, human social behavior, language acquisition in children, social cognitive neuroscience, and the cognitive neuroscience of language, it is argued that social cognition and language are two significantly interconnected capacities of the human species. Here, evidence in support of this view reviews (1) recent developmental studies on language learning in infants and young children, pointing to the important crucial benefits associated with social stimulation for youngsters, including the quality and quantity of incoming linguistic information, dyadic infant/child-to-parent non-verbal and verbal interactions, and other important social cues integral for facilitating language learning and social bonding; (2) studies of the adult human brain, suggesting a high degree of specialization for sociolinguistic information processing, memory retrieval, and comprehension, suggesting that the function of these neural areas may connect social cognition with language and social bonding; (3) developmental deficits in language and social cognition, including autism spectrum disorder (ASD), illustrating a unique developmental profile, further linking language, social cognition, and social bonding; and (4) neural biomarkers that may help to identify early developmental disorders of language and social cognition. In effect, the social brain and social complexity hypotheses may jointly help to describe how neurotypical children and adults acquire language, why autistic children and adults exhibit simultaneous deficits in language and social cognition, and why nonhuman primates and other organisms with significant computational capacities cannot learn language. But perhaps most critically, the following article argues that this and related research will allow scientists to generate a holistic profile and deeper understanding of the healthy adult social brain while developing more innovative and effective diagnoses, prognoses, and treatments for maladies and deficits also associated with the social brain.

Vocal complexity in a socially complex corvid: gradation, diversity and lack of common call repertoire in male rooks.

Vocal communication is widespread in animals, with vocal repertoires of varying complexity. The social complexity hypothesis predicts that species may need high vocal complexity to deal with complex social organization (e.g. have a variety of different interindividual relations). We quantified the vocal complexity of two geographically distant captive colonies of rooks, a corvid species with complex social organization and cognitive performances, but understudied vocal abilities. We quantified the diversity and gradation of their repertoire, as well as the inter-individual similarity at the vocal unit level. We found that males produced call units with lower diversity and gradation than females, while song units did not differ between sexes. Surprisingly, while females produced highly similar call repertoires, even between colonies, each individual male produced almost completely different call repertoires from any other individual. These findings question the way male rooks communicate with their social partners. We suggest that each male may actively seek to remain vocally distinct, which could be an asset in their frequently changing social environment. We conclude that inter-individual similarity, an understudied aspect of vocal repertoires, should also be considered as a measure of vocal complexity.

Social uncertainty promotes signal complexity during approaches in wild chimpanzees (Pan troglodytes verus) and mangabeys (Cercocebus atys atys).

The social complexity hypothesis for the evolution of communication posits that complex social environments require greater communication complexity for individuals to effectively manage their relationships. We examined how different socially uncertain contexts, reflecting an increased level of social complexity, relate to variation in signalling within and between two species, which display varying levels of fission-fusion dynamics (sympatric-living chimpanzees and sooty mangabeys, Taï National Park, Ivory Coast). Combined signalling may improve message efficacy, notably when involving different perception channels, thus may increase in moments of high social uncertainty. We examined the probability of individuals to emit no signal, single or multisensory or combined (complex) signals, during social approaches which resulted in non-agonistic outcomes. In both species, individuals were more likely to use more combined and multisensory signals in post-conflict approaches with an opponent than in other contexts. The clearest impact of social uncertainty on signalling complexity was observed during chimpanzee fusions, where the likelihood of using complex signals tripled relative to other contexts. Overall, chimpanzees used more multisensory signals than mangabeys. Social uncertainty may shape detected species differences in variation in signalling complexity, thereby supporting the hypothesis that social complexity, particularly associated with high fission-fusion dynamics, promotes signalling complexity.

Higher social tolerance is associated with more complex facial behavior in macaques

The social complexity hypothesis for communicative complexity posits that animal societies with more complex social systems require more complex communication systems. We tested the social complexity hypothesis on three macaque species that vary in their degree of social tolerance and complexity. We coded facial behavior in >3000 social interactions across three social contexts (aggressive, submissive, affiliative) in 389 animals, using the Facial Action Coding System for macaques (MaqFACS). We quantified communicative complexity using three measures of uncertainty: entropy, specificity, and prediction error. We found that the relative entropy of facial behavior was higher for the more tolerant crested macaques as compared to the less tolerant Barbary and rhesus macaques across all social contexts, indicating that crested macaques more frequently use a higher diversity of facial behavior. The context specificity of facial behavior was higher in rhesus as compared to Barbary and crested macaques, demonstrating that Barbary and crested macaques used facial behavior more flexibly across different social contexts. Finally, a random forest classifier predicted social context from facial behavior with highest accuracy for rhesus and lowest for crested, indicating there is higher uncertainty and complexity in the facial behavior of crested macaques. Overall, our results support the social complexity hypothesis.

Higher social tolerance is associated with more complex facial behavior in macaques.

The social complexity hypothesis for communicative complexity posits that animal societies with more complex social systems require more complex communication systems. We tested the social complexity hypothesis on three macaque species that vary in their degree of social tolerance and complexity. We coded facial behavior in >3000 social interactions across three social contexts (aggressive, submissive, affiliative) in 389 animals, using the Facial Action Coding System for macaques (MaqFACS). We quantified communicative complexity using three measures of uncertainty: entropy, specificity, and prediction error. We found that the relative entropy of facial behavior was higher for the more tolerant crested macaques as compared to the less tolerant Barbary and rhesus macaques across all social contexts, indicating that crested macaques more frequently use a higher diversity of facial behavior. The context specificity of facial behavior was higher in rhesus as compared to Barbary and crested macaques, demonstrating that Barbary and crested macaques used facial behavior more flexibly across different social contexts. Finally, a random forest classifier predicted social context from facial behavior with highest accuracy for rhesus and lowest for crested, indicating there is higher uncertainty and complexity in the facial behavior of crested macaques. Overall, our results support the social complexity hypothesis.

The functions and evolution of graded complex calls in a treefrog

ABSTRACT Variation in signal complexity is common among different species. Understanding why some species have evolved extensive call complexity can offer insight into the evolution of complex signals. In this study, we investigated the functions of different call types in a treefrog using call network analysis, male playback experiments, and correlation analysis between call properties and body size. Our results show that the male treefrogs can produce three kinds of notes that can be combined to produce single-note call types as well as composite calls. We also identified an intermediate note type between the advertisement call and suppress call, indicating that male suppress calls may have evolved from advertisement calls. The call network revealed functional correlation and distinction between different calls. Additionally, our study revealed that the dominant frequencies of the advertisement calls and suppress calls can signal the size of calling males. Male frogs can combine different notes in different sequences to form graded complex calls, which may perform different communication functions. Our results indicate that different communication needs may drive the evolution of signal complexity, which provides new evidence to the social complexity hypothesis and deeper insights into the function and complexity of animal acoustic communication.

Slow development of vocal sequences through ontogeny in wild chimpanzees (Pan troglodytes verus).

The development of the unique, hierarchical, and endless combinatorial capacity in a human language requires neural maturation and learning through childhood. Compared with most non-human primates, where combinatorial capacity seems limited, chimpanzees present a complex vocal system comprising hundreds of vocal sequences. We investigated how such a complex vocal system develops and the processes involved. We recorded 10,929 vocal utterances of 98 wild chimpanzees aged 0-55 years, from Taï National Park, Ivory Coast. We developed customized Generalized non-Linear Models to estimate the ontogenetic trajectory of four structural components of vocal complexity: utterance length, diversity, probability of panting (requiring phonation across inhalation and exhalation), and probability of producing two adjacent panted units. We found chimpanzees need 10 years to reach adult levels of vocal complexity. In three variables, the steepest increase coincided with the age of first non-kin social interactions (2-5 years), and plateaued in sub-adults (8-10 years), as individuals integrate into adult social life. Producing two adjacent panted units may require more neuromuscular coordination of the articulators, as its emergence and steepest increase appear later in development. These results suggest prolonged maturational processes beyond those hitherto thought likely in species that do not learn their vocal repertoire. Our results suggest that multifaceted ontogenetic processes drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation. As humans live in a complex social world, empirical support for the "social complexity hypothesis" may have relevance for theories of language evolution. RESEARCH HIGHLIGHTS: Chimpanzees need around 10 years to develop the vocal structural complexity present in the adult repertoire, way beyond the age of emergence of every single vocal unit. Multifaceted ontogenetic processes may drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation. Non-linear increases in vocal complexity coincide with social developmental milestones. Vocal sequences requiring rapid articulatory change emerge later than other vocal sequences, suggesting neuro-muscular maturational processes continue through the juvenile years.

Flexible use of contact calls in a species with high fission-fusion dynamics.

The 'social complexity hypothesis' posits that complex social systems (which entail high uncertainty) require complex communicative systems (with high vocal flexibility). In species with fission-fusion dynamics, where the fluid composition of temporary subgroups increases the uncertainty with which group members must manage their social relationships, vocal communication must be particularly flexible. This study assessed whether contact call rates vary with caller and audience characteristics in free-living spider monkeys, as well as with fission and fusion events. Adult females and immature individuals called more when in small audience settings, while audience size did not influence adult males. Adults called more when in the presence of the opposite sex, whereas immatures vocalized more in subgroups composed only by females. Females also called more when with their mature sons. We found higher call rates in periods during which fission and fusion events took place than in periods with more stable compositions and when the composition after a fission or fusion event changed from one sex to two sexes. A flexible use of contact calls allows individuals to identify themselves when they join others, particularly if they are members of the opposite sex. This socio-spatial cohesion function reduces the uncertainty about subgroup composition. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

Body size and sexual selection shaped the evolution of parrot calls.

Morphology, habitat and various selective pressures (e.g. social and sexual selection) can influence the evolution of acoustic signals, but the relative importance of their effects is not well understood. The order Psittaciformes (parrots, sensu lato) is a large clade of very vocal and often gregarious species for which large-scale comparative studies of vocalizations are lacking. We measured acoustic traits (duration, sound frequency, frequency bandwidth and sound entropy) of the predominant call type for >200 parrot species to test: (1) for associations with body size; (2) the acoustic adaptation hypothesis (AAH) (predicting differences between forest and open-habitat species); (3) the social complexity hypothesis (predicting more complex calls in gregarious species) and (4) influences of sexual selection (predicting correlated evolution with colour ornamentation). Larger species had on average longer calls, lower sound frequency and wider frequency bandwidth. These associations with body size are all predicted by physical principles of sound production. We found no evidence for the acoustic adaptation and social complexity hypotheses, but perhaps social complexity is associated with vocal traits not studied here, such as call repertoire sizes. More sexually dichromatic species had on average simpler calls (shorter, with lower entropy and narrower frequency bandwidth) indicating an influence of sexual selection, namely an evolutionary negative correlation between colour ornamentation and elaborate acoustic signals, as predicted by the transference hypothesis. Our study is the first large-scale attempt at understanding acoustic diversity across the Psittaciformes, and indicates that body size and sexual selection influenced the evolution of species differences in vocal signals.

Does Social Complexity Drive Vocal Complexity? Insights from the Two African Elephant Species.

Simple SummaryThe social complexity hypothesis (SCH) for communication predicts that species with complex social systems exhibit complex communication systems. Testing the SHC in a broad range of species can contribute to a better understanding of human evolution because a co-evolutionary runaway process between social and vocal complexity may have shaped human language. Here we compare patterns of vocal complexity between the two species of African elephants: the savanna elephant exhibiting a complex social organization and the forest elephant exhibiting a simpler social organization. We review the existing literature and present novel insights into the vocal communication system of the elusive forest elephant, along with a first direct comparison with savanna elephants. Our findings suggest that the African elephants may contradict the SCH, as well as other factors potentially shaping patterns of vocal complexity across species. A better understanding of vocal complexity in the two species of African elephants will depend on continuing advancements in remote data collection technologies to overcome the challenges of observing forest elephants in their dense rainforest habitat, as well as the availability of comparable data quantifying both structural and contextual variability in the vocal production of both species of African elephants.The social complexity hypothesis (SCH) for communication states that the range and frequency of social interactions drive the evolution of complex communication systems. Surprisingly, few studies have empirically tested the SHC for vocal communication systems. Filling this gap is important because a co-evolutionary runaway process between social and vocal complexity may have shaped the most intricate communication system, human language. We here propose the African elephant Loxodonta spec. as an excellent study system to investigate the relationships between social and vocal complexity. We review how the distinct differences in social complexity between the two species of African elephants, the forest elephant L. cyclotis and the savanna elephant L. africana, relate to repertoire size and structure, as well as complex communication skills in the two species, such as call combination or intentional formant modulation including the trunk. Our findings suggest that Loxodonta may contradict the SCH, as well as other factors put forth to explain patterns of vocal complexity across species. We propose that life history traits, a factor that has gained little attention as a driver of vocal complexity, and the extensive parental care associated with a uniquely low and slow reproductive rate, may have led to the emergence of pronounced vocal complexity in the forest elephant despite their less complex social system compared to the savanna elephant. Conclusions must be drawn cautiously, however. A better understanding of vocal complexity in the genus Loxodonta will depend on continuing advancements in remote data collection technologies to overcome the challenges of observing forest elephants in their dense rainforest habitat, as well as the availability of directly comparable data and methods, quantifying both structural and contextual variability in the production of rumbles and other vocalizations in both species of African elephants.

A review of the social behaviour of the giraffeGiraffa camelopardalis: a misunderstood but socially complex species

AbstractUntil around 2000, giraffeGiraffa camelopardaliswere believed to have no social structure. Despite a resurgence of interest in giraffe behaviour in around 2010, most studies are of isolated populations, making it difficult to draw general conclusions. Although it is now well established that giraffe social organisation is non‐random, there is little consensus as to what influences preferred and avoided associations or the underpinning mechanisms.We test two hypotheses: first, giraffe have a complex cooperative social system, exhibited by 1) stable groups of females, 2) offspring that stay in their natal group for part or all of their lives, 3) support by non‐mothers in rearing young, and 4) non‐reproductive females in the group; and second, giraffe form matrilineal societies, evidenced by 1) male dispersal, 2) female philopatry, 3) assistance in raising or protecting offspring, and 4) individual benefits gained from social foraging.We reviewed 404 papers on giraffe behaviour and social organisation; captive studies were included where they supplemented information from free‐living populations.We show that giraffe exhibit many of the features typical of mammals with complex cooperative social systems and matrilineal societies. However, the social complexity hypothesis posits that such species also require complex communication systems to regulate interactions and relations among group members; giraffe communication systems are poorly understood.Quantifying the fitness and survival benefits of the giraffe’s social organisation is necessary to ensure its long‐term survival. Giraffe numbers have declined by 40% since 1985, they have been declared extinct in seven (possibly nine) countries and are listed as Vulnerable by the International Union for Conservation of Nature. We identify research areas that will advance our understanding of giraffe behaviour and conservation requirements.

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.

Tolerant and intolerant macaques show different levels of structural complexity in their vocal communication.

We tested the social complexity hypothesis which posits that animals living in complex social environments should use complex communication systems. We focused on two components of vocal complexity: diversity (number of categories of calls) and flexibility (degree of gradation between categories of calls). We compared the acoustic structure of vocal signals in groups of macaques belonging to four species with varying levels of uncertainty (i.e. complexity) in social tolerance (the higher the degree of tolerance, the higher the degree of uncertainty): two intolerant species, Japanese and rhesus macaques, and two tolerant species, Tonkean and crested macaques. We recorded the vocalizations emitted by adult females in affiliative, agonistic and neutral contexts. We analysed several acoustic variables: call duration, entropy, time and frequency energy quantiles. The results showed that tolerant macaques displayed higher levels of vocal diversity and flexibility than intolerant macaques in situations with a greater number of options and consequences, i.e. in agonistic and affiliative contexts. We found no significant differences between tolerant and intolerant macaques in the neutral context where individuals are not directly involved in social interaction. This shows that species experiencing more uncertain social interactions displayed greater vocal diversity and flexibility, which supports the social complexity hypothesis.

Facial expressions and pair bonds in hylobatids.

Facial expressions are an important component of primate communication that functions to transmit social information and modulate intentions and motivations. Chimpanzees and macaques, for example, produce a variety of facial expressions when communicating with conspecifics. Hylobatids also produce various facial expressions; however, the origin and function of these facial expressions are still largely unclear. It has been suggested that larger facial expression repertoires may have evolved in the context of social complexity, but this link has yet to be tested at a broader empirical basis. The social complexity hypothesis offers a possible explanation for the evolution of complex communicative signals such as facial expressions, because as the complexity of an individual's social environment increases so does the need for communicative signals. We used an intraspecies, pair-focused study design to test the link between facial expressions and sociality within hylobatids, specifically the strength of pair-bonds. The current study compared 206 hr of video and 103 hr of focal animal data for ten hylobatid pairs from three genera (Nomascus, Hoolock, and Hylobates) living at the Gibbon Conservation Center. Using video footage, we explored 5,969 facial expressions along three dimensions: repertoire use, repertoire breadth, and facial expression synchrony [FES]. We then used focal animal data to compare dimensions of facial expressiveness to pair bond strength and behavioral synchrony. Hylobatids in our study overlapped in only half of their facial expressions (50%) with the only other detailed, quantitative study of hylobatid facial expressions, while 27 facial expressions were uniquely observed in our study animals. Taken together, hylobatids have a large facial expression repertoire of at least 80 unique facial expressions. Contrary to our prediction, facial repertoire composition was not significantly correlated with pair bond strength, rates of territorial synchrony, or rates of behavioral synchrony. We found that FES was the strongest measure of hylobatid expressiveness and was significantly positively correlated with higher sociality index scores; however, FES showed no significant correlation with behavioral synchrony. No noticeable differences between pairs were found regarding rates of behavioral or territorial synchrony. Facial repertoire sizes and FES were not significantly correlated with rates of behavioral synchrony or territorial synchrony. Our study confirms an important role of facial expressions in maintaining pair bonds and coordinating activities in hylobatids. Data support the hypothesis that facial expressions and sociality have been linked in hylobatid and primate evolution. It is possible that larger facial repertoires may have contributed to strengthening pair bonds in primates, because richer facial repertoires provide more opportunities for FES which can effectively increase the "understanding" between partners through smoother coordination of interaction patterns. This study supports the social complexity hypothesis as the driving force for the evolution of complex communication signaling.

Sociality and communicative complexity: insights from the other insect societies.

Recognition and communication are essential processes when it comes to interaction of organisms with their biotic environment. As especially social interactions are coordinated by communication, it has been predicted that social evolution drives communicative complexity. However, studies comparing olfactory signals or receptor repertoires of solitary and eusocial insects found only mixed evidence for the social complexity hypothesis. We present some possible explanations and especially argue that our current knowledge of intermediate levels of sociality is insufficient to fully test the hypothesis, for which a more balanced comparative dataset would be required. We illustrate with chosen examples how complex communication within the other insect societies can be: Many messages are not unique to eusocial insects. Studying the other insect societies will provide us with a more detailed picture of the link between social and communicative complexity.

Vocal complexity and sociality in spotted paca (Cuniculus paca).

The evolution of sociality is related to many ecological factors that act on animals as selective forces, thus driving the formation of groups. Group size will depend on the payoffs of group living. The Social Complexity Hypothesis for Communication (SCHC) predicts that increases in group size will be related to increases in the complexity of the communication among individuals. This hypothesis, which was confirmed in some mammal societies, may be useful to trace sociality in the spotted paca (Cuniculus paca), a Neotropical caviomorph rodent reported as solitary. There are, however, sightings of groups in the wild, and farmers easily form groups of spotted paca in captivity. Thus, we aimed to describe the acoustic repertoire of captive spotted paca to test the SCHC and to obtain insights about the sociability of this species. Moreover, we aimed to verify the relationship between group size and acoustic repertoire size of caviomorph rodents, to better understand the evolution of sociality in this taxon. We predicted that spotted paca should display a complex acoustic repertoire, given their social behavior in captivity and group sightings in the wild. We also predicted that in caviomorph species the group size would increase with acoustic repertoire, supporting the SCHC. We performed a Linear Discriminant Analysis (LDA) based on acoustic parameters of the vocalizations recorded. In addition, we applied an independent contrasts approach to investigate sociality in spotted paca following the social complexity hypothesis, independent of phylogeny. Our analysis showed that the spotted paca’s acoustic repertoire contains seven vocal types and one mechanical signal. The broad acoustic repertoire of the spotted paca might have evolved given the species’ ability to live in groups. The relationship between group size and the size of the acoustic repertoires of caviomorph species was confirmed, providing additional support for the SCHC in yet another group of diverse mammals–caviomorph rodents.

Cooperative breeding influences the number and type of vocalizations in avian lineages.

Although communicative complexity is often predicted to correlate with social complexity in animal societies, few studies have employed large-scale comparative analyses to test whether socially complex species have more complex systems of communication. I tested this social complexity hypothesis in birds (Class: Aves) using the large amount of natural history information that describes both vocal repertoire and social system in these species. To do so, I marshalled data from primary and secondary records of avian vocal repertoires (n = 253), and for each of the species in the dataset I recorded the reported repertoire size and associated species information. Using phylogenetic comparative methods, I found that cooperative breeding was a strong and repeatable predictor of vocal repertoire size, while other social variables, e.g. group size and group stability, had little or no influence on repertoire size. Importantly, repertoire sizes expanded concurrently with the evolution of cooperative breeding, suggesting a direct link between these two traits. Cooperatively breeding species devoted significantly more of their repertoire to contact calls and alarm calls. Overall, these results therefore lend support to the hypothesis that social complexity via behavioural coordination leads to increases in vocal complexity.

Reciprocal greeting in chimpanzees ( Pan troglodytes ) at the Ngogo community

Abstract Human conversations are organized according to operative sequential features with different components of utterances being related to each other forming so-called adjacency pairings. An adjacency pair is a sequential structure produced by both speakers, consisting of two parts, with the second part being contingent upon and normatively obliged to the first part, such as greeting-greeting exchanges. The present paper investigates such ordered serial utterances in chimpanzee greeting interactions. Results show that reciprocal and structured communicative ‘discourse’ is relatable to the communicative modality that is chosen for greetings as well as to the social bonding between the greeting interactants. The findings of the present study are discussed in relation to the social complexity hypothesis and the current debate on the importance of the turn-taking system for language evolution.

A Dominant Social Comparison Heuristic Unites Alternative Mechanisms for the Evolution of Indirect Reciprocity.

Cooperation is a fundamental human trait but our understanding of how it functions remains incomplete. Indirect reciprocity is a particular case in point, where one-shot donations are made to unrelated beneficiaries without any guarantee of payback. Existing insights are largely from two independent perspectives: i) individual-level cognitive behaviour in decision making, and ii) identification of conditions that favour evolution of cooperation. We identify a fundamental connection between these two areas by examining social comparison as a means through which indirect reciprocity can evolve. Social comparison is well established as an inherent human disposition through which humans navigate the social world by self-referential evaluation of others. Donating to those that are at least as reputable as oneself emerges as a dominant heuristic, which represents aspirational homophily. This heuristic is found to be implicitly present in the current knowledge of conditions that favour indirect reciprocity. The effective social norms for updating reputation are also observed to support this heuristic. We hypothesise that the cognitive challenge associated with social comparison has contributed to cerebral expansion and the disproportionate human brain size, consistent with the social complexity hypothesis. The findings have relevance for the evolution of autonomous systems that are characterised by one-shot interactions.