Inferential Communication: Bridging the Gap Between Intentional and Ostensive Communication in Non-human Primates

Communication, when defined as an act intended to affect the psychological state of another individual, demands the use of inference. Either the signaler, the recipient, or both must make leaps of understanding which surpass the semantic information available and draw from pragmatic clues to fully imbue and interpret meaning. While research into human communication and the evolution of language has long been comfortable with mentalistic interpretations of communicative exchanges, including rich attributions of mental state, research into animal communication has balked at theoretical models which describe mentalized cognitive mechanisms. We submit a new theoretical perspective on animal communication: the model of inferential communication. For use when existing proximate models of animal communication are not sufficient to fully explain the complex, flexible, and intentional communication documented in certain species, specifically non-human primates, we present our model as a bridge between shallower, less cognitive descriptions of communicative behavior and the perhaps otherwise inaccessible mentalistic interpretations of communication found in theoretical considerations of human language. Inferential communication is a framework that builds on existing evidence of referentiality, intentionality, and social inference in primates. It allows that they might be capable of applying social inferences to a communicative setting, which could explain some of the cognitive processes that enable the complexity and flexibility of primate communication systems. While historical models of animal communication focus on the means-ends process of behavior and apparent cognitive outcomes, inferential communication invites consideration of the mentalistic processes that must underlie those outcomes. We propose a mentalized approach to questions, investigations, and interpretations of non-human primate communication. We include an overview of both ultimate and proximate models of animal communication, which contextualize the role and utility of our inferential communication model, and provide a detailed breakdown of the possible levels of cognitive complexity which could be investigated using this framework. Finally, we present some possible applications of inferential communication in the field of non-human primate communication and highlight the role it could play in advancing progress toward an increasingly precise understanding of the cognitive capabilities of our closest living relatives.

Concerted variation in melanogenesis genes underlies emergent patterning of plumage in capuchino seedeaters

Coloration traits are central to animal communication; they often govern mate choice, promote reproductive isolation and catalyse speciation. Specific genetic changes can cause variation in coloration, yet far less is known about how overall coloration patterns—which involve combinations of multiple colour patches across the body—can arise and are genomically controlled. We performed genome-wide association analyses to link genomic changes to variation in melanin (eumelanin and pheomelanin) concentration in feathers from different body parts in the capuchino seedeaters, an avian radiation with diverse colour patterns despite remarkably low genetic differentiation across species. Cross-species colour variation in each plumage patch is associated with unique combinations of variants at a few genomic regions, which include mostly non-coding (presumably regulatory) areas close to known pigmentation genes. Genotype–phenotype associations can vary depending on patch colour and are stronger for eumelanin pigmentation, suggesting eumelanin production is tightly regulated. Although some genes are involved in colour variation in multiple patches, in some cases, the SNPs associated with colour changes in different patches segregate spatially. These results suggest that coloration patterning in capuchinos is generated by the modular combination of variants that regulate multiple melanogenesis genes, a mechanism that may have promoted this rapid radiation.

Human aggression and music evolution: a model

Together with language, music is perhaps our most distinctive behavioral trait. Following the lead of paleolinguistic research, different hypotheses have been proposed to explain why only humans perform music and how this ability might have evolved in the species. In this paper, we advance a new model of music evolution that builds on the theory of self-domestication, according to which the human phenotype is, at least in part, the outcome of a process similar to mammal domestication, triggered by a progressive reduction in reactive aggression levels in response to environmental changes. In the paper, we specifically argue that changes in aggression management through the course of human cultural evolution can account for the behaviors conducive to the emergence and evolution of music. We hypothesize 4 stages in the evolutionary development of music under the influence of environmental changes and evolution of social organization: starting from musilanguage, proto-music gave rise to personal and private forms of timbre-oriented music, then to small-group ensembles of pitch-oriented music, at first of indefinite and then definite pitch, and finally to collective (tonal) music. These stages parallel what has been hypothesized for languages and encompass the diversity of music types and genres described worldwide. Overall, music complexity emerges in a gradual fashion under the effects of enhanced abilities for cultural niche construction, resulting from the stable trend of reduction in reactive aggression towards the end of the Pleistocene, leading to the rise of hospitality codes, and succeeded by increase in proactive aggression from the beginning of the Holocene onward. This paper addresses numerous controversies in the literature on the evolution of music by providing a clear structural definition of music, identifying its structural features that distinguish it from oral language, and summarizing the typology of operational functions of music and formats of its transmission. The proposed framework of structural approach to music arms a researcher with means to identify and comparatively analyze different schemes of tonal organization of music, placing them in the context of human social and cultural evolution. Especially valuable contribution to the understanding of transition from animal communication to human music and language is the theory of so-called “personal song”, described and analyzed here from ethological, social, cultural, cognitive, and musicological perspectives. The emergence of personal song and its development into a social institution are interlinked with the evolution of kinship and placed into the timeline of cultural evolution, based on totality of ethnographic, archaeological, anthropological, genetic, and paleoclimatic data.

Sexual and individual signatures are encoded in the temporal rate of Cape gannet display calls

Vocalisations play a vital role in animal communication, as they are involved in many biological functions. Seabirds often breed in large and dense colonies, making successful recognition between mates or between parents-and offspring crucial for reproductive success. Most seabird species, including Cape gannets (Morus capensis), are monomorphic and likely rely on acoustic signals for mate selection and mate recognition. This study aimed to better understand the use of vocalisations for sex and individual recognition in Cape gannets by describing the acoustic structure of their display calls at the nest. Vocalisations of nesting Cape gannets were recorded and acoustic measurements were extracted in both temporal and frequency domains. Values of the fundamental frequency and the average of Inter-Onset-Interval appeared to be the most important acoustic variables for sex determination. Both temporal and frequency parameters showed a potential for individual identity coding, with the average units Inter-Onset-Interval being the most important variable for individual identification for both sexes. This study provides the first evidence of sex-specific and individual vocal signatures in adult breeding Cape gannets. From an applied perspective, identified sex specific differences could potentially be used as a non-invasive method for field-based sex-determination in research and monitoring projects on Cape gannets.

Overlooked evidence for semantic compositionality and signal reduction in wild chimpanzees (Pan troglodytes)

Recent discoveries of semantic compositionality in Japanese tits have enlivened the discussions on the presence of this phenomenon in wild animal communication. Data on semantic compositionality in wild apes are lacking, even though language experiments with captive apes have demonstrated they are capable of semantic compositionality. In this paper, I revisit the study by Boesch (Hum. Evol. 6:81–89, 1991) who investigated drumming sequences by an alpha male in a chimpanzee (Pan troglodytes) community in the Taï National Park, Côte d’Ivoire. A reanalysis of the data reveals that the alpha male produced semantically compositional combined messages of travel direction change and resting period initiation. Unlike the Japanese tits, the elements of the compositional expression were not simply juxtaposed but displayed structural reduction, while one of the two elements in the expression coded the meanings of both elements. These processes show relative resemblance to blending and fusion in human languages. Also unlike the tits, the elements of the compositional expression did not have a fixed order, although there was a fixed distribution of drumming events across the trees used for drumming. Because the elements of the expression appear to carry verb-like meanings, the compositional expression also resembles simple verb-verb constructions and short paratactic combinations of two clauses found across languages. In conclusion, the reanalysis suggests that semantic compositionality and phenomena resembling paratactic combinations of two clauses might have been present in the communication of the last common ancestor of chimpanzees and humans, not necessarily in the vocal modality.

Chimpanzee vowel-like sounds and voice quality suggest formant space expansion through the hominoid lineage

The origins of human speech are obscure; it is still unclear what aspects are unique to our species or shared with our evolutionary cousins, in part due to a lack of a common framework for comparison. We asked what chimpanzee and human vocal production acoustics have in common. We examined visible supra-laryngeal articulators of four major chimpanzee vocalizations (hoos, grunts, barks, screams) and their associated acoustic structures, using techniques from human phonetic and animal communication analysis. Data were collected from wild adult chimpanzees, Taï National Park, Ivory Coast. Both discriminant and principal component classification procedures revealed classification of call types. Discriminating acoustic features include voice quality and formant structure, mirroring phonetic features in human speech. Chimpanzee lip and jaw articulation variables also offered similar discrimination of call types. Formant maps distinguished call types with different vowel-like sounds. Comparing our results with published primate data, humans show less F1–F2 correlation and further expansion of the vowel space, particularly for [i] sounds. Unlike recent studies suggesting monkeys achieve human vowel space, we conclude from our results that supra-laryngeal articulatory capacities show moderate evolutionary change, with vowel space expansion continuing through hominoid evolution. Studies on more primate species will be required to substantiate this. This article is part of the theme issue ‘Voice modulation: from origin and mechanism to social impact (Part II)’.

Recursion: What is it, who has it, and how did it evolve?

Recursion is a topic of considerable controversy in linguistics, which stems from its varying definitions and its key features, such as its universality, uniqueness to human language, and evolution. Currently, there appear to be at least two common senses of recursion: (1) embeddedness of phrases within other phrases, which entails keeping track of long-distance dependencies among phrases and (2) the specification of the computed output string itself, including meta-recursion, where recursion is both the recipe for an utterance and the overarching process that creates and executes the recipes. There are also at least two evolutionary scenarios for the adaptive value of recursion in human language. The gradualist position posits precursors, such as animal communication and protolanguages, and holds that the selective purpose of recursion was for communication. The saltationist position assumes no gradual development of recursion and posits that it evolved for reasons other than communication. In the latter view, some heritable event associated with a cognitive prerequisite of language, such as Theory of Mind or working memory capacity, allowed recursive utterances. Evolutionary adaptive reasons for recursive thoughts were also proffered, including diplomatic speech, perlocutionary acts, and prospective cognitions.