Drew Rendall

What do animal signals mean

Animal communication studies often use analogies to human language and related constructs such as information encoding and transfer. This commonality is evident even when research goals are very different, for example when primate vocalizations are proposed to have word-like meaning, or sexually selected signals are proposed to convey information about a signaller’s underlying quality. We consider some of the ambiguities and limitations inherent in such informational approaches to animal communication as background to advocating alternatives. The alternatives eschew language-based metaphors and broader informational constructs and focus instead on concrete details of signal design as they reflect and interact with established sensory, physiological and psychological processes that support signalling and responding in listeners. The alternatives we advocate also explicitly acknowledge the different roles and often divergent interests of signallers and perceivers that can yield fundamental asymmetries in signalling interactions, and they therefore shift the focus of interpretations of animal communication from informing others to influencing others.

Pitch (F0) and formant profiles of human vowels and vowel-like baboon grunts: The role of vocalizer body size and voice-acoustic allometry

Key voice features--fundamental frequency (F0) and formant frequencies--can vary extensively between individuals. Much of the variation can be traced to differences in the size of the larynx and vocal-tract cavities, but whether these differences in turn simply reflect differences in speaker body size (i.e., neutral vocal allometry) remains unclear. Quantitative analyses were therefore undertaken to test the relationship between speaker body size and voice F0 and formant frequencies for human vowels. To test the taxonomic generality of the relationships, the same analyses were conducted on the vowel-like grunts of baboons, whose phylogenetic proximity to humans and similar vocal production biology and voice acoustic patterns recommend them for such comparative research. For adults of both species, males were larger than females and had lower mean voice F0 and formant frequencies. However, beyond this, F0 variation did not track body-size variation between the sexes in either species, nor within sexes in humans. In humans, formant variation correlated significantly with speaker height but only in males and not in females. Implications for general vocal allometry are discussed as are implications for speech origins theories, and challenges to them, related to laryngeal position and vocal tract length.

Factors Affecting Reproduction and Mortality Among Baboons in the Okavango Delta, Botswana

We present results of a 10-year study of free-ranging gray-footed chacma baboons (Papio ursinus griseipes) in the Okavango Delta of Botswana. The majority of deaths among adult females and juveniles were due to predation, while infants were more likely to die of infanticide. There were strong seasonal effects on birth and mortality, with the majority of conceptions occurring during the period of highest rainfall. Mortality due to predation and infanticide was highest during the 3-mo period when flooding was at its peak, when the group was more scattered and constrained to move along predictable routes. The reproductive parameters most likely to be associated with superior competitive ability—interbirth interval and infant growth rates—conferred a slight fitness advantage on high-ranking females. However, it was counterbalanced by the effects of infanticide and predation. Infanticide affected high- and low-ranking females more than middle-ranking females, while predation affected females of all ranks relatively equally. As a result, there were few rank-related differences in estimated female lifetime reproductive success.

Acoustic correlates of caller identity and affect intensity in the vowel-like grunt vocalizations of baboons

Comparative, production-based research on animal vocalizations can allow assessments of continuity in vocal communication processes across species, including humans, and may aid in the development of general frameworks relating specific constitutional attributes of callers to acoustic-structural details of their vocal output. Analyses were undertaken on vowel-like baboon grunts to examine variation attributable to caller identity and the intensity of the affective state underlying call production. Six hundred six grunts from eight adult females were analyzed. Grunts derived from 128 bouts of calling in two behavioral contexts: concerted group movements and social interactions involving mothers and their young infants. Each context was subdivided into a high- and low-arousal condition. Thirteen acoustic features variously predicted to reflect variation in either caller identity or arousal intensity were measured for each grunt bout, including tempo-, source- and filter-related features. Grunt bouts were highly individually distinctive, differing in a variety of acoustic dimensions but with some indication that filter-related features contributed disproportionately to individual distinctiveness. In contrast, variation according to arousal condition was associated primarily with tempo- and source-related features, many matching those identified as vehicles of affect expression in other nonhuman primate species and in human speech and other nonverbal vocal signals.

The role of vocal tract filtering in identity cueing in rhesus monkey (Macaca mulatta) vocalizations

The importance of individual identity and kinship has been demonstrated in the social behavior of many nonhuman primates, with some evidence suggesting that individually distinctive acoustic features are present in their vocalizations as well. In order to systematically test whether acoustic cues to identity are reliably present across the vocal repertoire of rhesus monkeys (Macaca mulatta), we examined coos, grunts, and noisy screams produced by adult females of two free-ranging groups. First, acoustic analyses were used to characterize spectral patterning, the fundamental frequency, and temporal characteristics of these three distinct call types. Vocalizations were then classified by caller identity, based on discriminant function analyses. Results showed that coos (rich, harmonically structured sounds) were markedly more distinctive by caller than were either grunts or noisy screams, and that spectral-patterning measures related to vocal tract filtering effects were the most reliable markers of individual identity. Grunts (pulsed, noisy calls) were classified at lower, but above-chance rates and spectral patterning cues were again critical in this sorting. Noisy screams (continuous, broadband noise bursts that could include a high-frequency, periodic component) could not be reliably sorted by caller. Playback experiments conducted with the screams showed no response differences when listening animals heard vocalizations produced by kin or nonkin individuals. This result was strikingly different from the corresponding outcome of a previous test with coo calls, but consistent with the acoustic analysis. Implications of these findings for vocal production mechanisms in nonhuman primates and previous studies of rhesus monkey vocalizations are discussed.

Social brains, simple minds: does social complexity really require cognitive complexity?

The social brain hypothesis is a well-accepted and well-supported evolutionary theory of enlarged brain size in the non-human primates. Nevertheless, it tends to emphasize an anthropocentric view of social life and cognition. This often leads to confusion between ultimate and proximate mechanisms, and an over-reliance on a Cartesian, narratively structured view of the mind and social life, which in turn lead to views of social complexity that are congenial to our views of ourselves, rather than necessarily representative of primate social worlds. In this paper, we argue for greater attention to embodied and distributed theories of cognition, which get us away from current fixations on ‘theory of mind’ and other high-level anthropocentric constructions, and allow for the generation of testable hypotheses that combine neurobiology, psychology and behaviour in a mutually reinforcing manner.

An Affect-Conditioning Model of Nonhuman Primate Vocal Signaling

We outline a model of nonhuman primate vocal behavior, proposing that the function of calling is to influence the behavior of conspecific receivers and that a Pavlovian conditioning framework can account for important aspects of how such influence occurs. Callers are suggested to use vocalizations to elicit affective responses in others, thereby altering the behavior of these individuals. Responses can either be unconditioned, being produced directly by the signal itself, or conditioned, resulting from past interactions in which the sender both called and produced affective responses in the receiver through other means.

Evolution of human vocal production

Determining the substrates required for the evolution of human speech is difficult as most traits thought to give rise to human speech — the vocal production apparatus and the brain — do not fossilize. Nor do we have any ‘proto-human’ sound tracks to analyse. The fossil record is also of limited utility for identifying indicators of pre-historic linguistic abilities. Ultimately, we are left with only one reliable way of investigating the biological mechanisms underlying the evolution of speech: the comparative method.

Two organizing principles of vocal production: Implications for nonhuman and human primates.

Vocal communication in nonhuman primates receives considerable research attention, with many investigators arguing for similarities between this calling and speech in humans. Data from development and neural organization show a central role of affect in monkey and ape sounds, however, suggesting that their calls are homologous to spontaneous human emotional vocalizations while having little relation to spoken language. Based on this evidence, we propose two principles that can be useful in evaluating the many and disparate empirical findings that bear on the nature of vocal production in nonhuman and human primates. One principle distinguishes production‐first from reception‐first vocal development, referring to the markedly different role of auditory‐motor experience in each case. The second highlights a phenomenon dubbed dual neural pathways, specifically that when a species with an existing vocal system evolves a new functionally distinct vocalization capability, it occurs through emergence of a second parallel neural pathway rather than through expansion of the extant circuitry. With these principles as a backdrop, we review evidence of acoustic modification of calling associated with background noise, conditioning effects, audience composition, and vocal convergence and divergence in nonhuman primates. Although each kind of evidence has been interpreted to show flexible cognitively mediated control over vocal production, we suggest that most are more consistent with affectively grounded mechanisms. The lone exception is production of simple, novel sounds in great apes, which is argued to reveal at least some degree of volitional vocal control. If also present in early hominins, the cortically based circuitry surmised to be associated with these rudimentary capabilities likely also provided the substrate for later emergence of the neural pathway allowing volitional production in modern humans. Am. J. Primatol. 73:530–544, 2011.

Lifting the curtain on the Wizard of Oz: Biased voice-based impressions of speaker size.

The consistent, but often wrong, impressions people form of the size of unseen speakers are not random but rather point to a consistent misattribution bias, one that the advertising, broadcasting, and entertainment industries also routinely exploit. The authors report 3 experiments examining the perceptual basis of this bias. The results indicate that, under controlled experimental conditions, listeners can make relative size distinctions between male speakers using reliable cues carried in voice formant frequencies (resonant frequencies, or timbre) but that this ability can be perturbed by discordant voice fundamental frequency (F-sub-0, or pitch) differences between speakers. The authors introduce 3 accounts for the perceptual pull that voice F-sub-0 can exert on our routine (mis)attributions of speaker size and consider the role that voice F-sub-0 plays in additional voice-based attributions that may or may not be reliable but that have clear size connotations.