Mapping vocal interactions in space and time differentiates signal broadcast versus signal exchange in meerkat groups.

Animal vocal communication research traditionally focuses on acoustic and contextual features of calls, yet substantial information is also contained in response selectivity and timing during vocalization events. By examining the spatiotemporal structure of vocal interactions, we can distinguish between 'broadcast' and 'exchange' signalling modes, with the former potentially serving to transmit signallers' general state and the latter reflecting more interactive signalling behaviour. Here, we tracked the movements and vocalizations of wild meerkat (Suricata suricatta) groups simultaneously using collars to explore this distinction. We found evidence that close calls (used for maintaining group cohesion) are given as signal exchanges. They are typically given in temporally structured call-response sequences and are also strongly affected by the social environment, with individuals calling more when they have more neighbours and juveniles responding more to adults than the reverse. In contrast, short note calls appear mainly in sequences produced by single individuals and show little dependence on social surroundings, suggesting a broadcast signalling mode. Despite these differences, both call categories show similar clustering in space and time at a group level. Our results highlight how the fine-scale structure of vocal interactions can give important insights into the usage and function of signals in social groups. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics.'

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.

Sensory collectives in natural systems.

Groups of animals inhabit vastly different sensory worlds, or umwelten, which shape fundamental aspects of their behaviour. Yet the sensory ecology of species is rarely incorporated into the emerging field of collective behaviour, which studies the movements, population-level behaviours, and emergent properties of animal groups. Here, we review the contributions of sensory ecology and collective behaviour to understanding how animals move and interact within the context of their social and physical environments. Our goal is to advance and bridge these two areas of inquiry and highlight the potential for their creative integration. To achieve this goal, we organise our review around the following themes: (1) identifying the promise of integrating collective behaviour and sensory ecology; (2) defining and exploring the concept of a 'sensory collective'; (3) considering the potential for sensory collectives to shape the evolution of sensory systems; (4) exploring examples from diverse taxa to illustrate neural circuits involved in sensing and collective behaviour; and (5) suggesting the need for creative conceptual and methodological advances to quantify 'sensescapes'. In the final section, (6) applications to biological conservation, we argue that these topics are timely, given the ongoing anthropogenic changes to sensory stimuli (e.g. via light, sound, and chemical pollution) which are anticipated to impact animal collectives and group-level behaviour and, in turn, ecosystem composition and function. Our synthesis seeks to provide a forward-looking perspective on how sensory ecologists and collective behaviourists can both learn from and inspire one another to advance our understanding of animal behaviour, ecology, adaptation, and evolution.

Correction to: 'Fission-fusion dynamics in sheep: the influence of resource distribution and temporal activity patterns' (2023) by Della Libera et al.

[This corrects the article DOI: 10.1098/10.1098/rsos.230402.][This corrects the article DOI: 10.1098/rspb.2021.1115.].

Humans strategically avoid connecting to others who agree and avert the emergence of network polarization in a coordination task.

Clusters of like-minded individuals can impede consensus in group decision-making. We implemented an online color coordination task to investigate whether control over communication links creates clusters impeding group consensus. In 244 6-member networks, individuals were incentivized to reach a consensus by agreeing on a color, but had conflicting incentives for which color to choose. We varied (1) if communication links were static, changed randomly over time, or were player-controlled; (2) whether links determined who was observed or addressed; and (3) whether a majority existed or equally many individuals preferred each color. We found that individuals preferentially selected links to previously unobserved and disagreeing others, avoiding links with agreeing others. This prevented cluster formation, sped up consensus formation rather than impeding it, and increased the probability that the group agreed on the majority incentive. Overall, participants with a consensus goal avoided clusters by applying strategies that resolved uncertainty about others.

Fission-fusion dynamics in sheep: the influence of resource distribution and temporal activity patterns.

Fission-fusion events, i.e. changes to the size and composition of animal social groups, are a mechanism to adjust the social environment in response to short-term changes in the cost-benefit ratio of group living. Furthermore, the time and location of fission-fusion events provide insight into the underlying drivers of these dynamics. Here, we describe a method for identifying group membership over time and for extracting fission-fusion events from animal tracking data. We applied this method to high-resolution GPS data of free-ranging sheep (Ovis aries). Group size was highest during times when sheep typically rest (midday and at night), and when anti-predator benefits of grouping are high while costs of competition are low. Consistent with this, fission and fusion frequencies were highest during early morning and late evening, suggesting that social restructuring occurs during periods of high activity. However, fission and fusion events were not more frequent near food patches and water resources when adjusted for overall space use. This suggests a limited role of resource competition. Our results elucidate the dynamics of grouping in response to social and ecological drivers, and we provide a tool for investigating these dynamics in other species.

Link updating strategies influence consensus decisions as a function of the direction of communication.

Consensus decision-making in social groups strongly depends on communication links that determine to whom individuals send, and from whom they receive, information. Here, we ask how consensus decisions are affected by strategic updating of links and how this effect varies with the direction of communication. We quantified the coevolution of link and opinion dynamics in a large population with binary opinions using mean-field numerical simulations of two voter-like models of opinion dynamics: an incoming model (IM) (where individuals choose who to receive opinions from) and an outgoing model (OM) (where individuals choose who to send opinions to). We show that individuals can bias group-level outcomes in their favour by breaking disagreeing links while receiving opinions (IM) and retaining disagreeing links while sending opinions (OM). Importantly, these biases can help the population avoid stalemates and achieve consensus. However, the role of disagreement avoidance is diluted in the presence of strong preferences-highly stubborn individuals can shape decisions to favour their preferences, giving rise to non-consensus outcomes. We conclude that collectively changing communication structures can bias consensus decisions, as a function of the strength of preferences and the direction of communication.

Directional speakers as a tool for animal vocal communication studies.

Audio playbacks are a common experimental tool in vocal communication research. However, low directionality of sound makes it hard to control the audience exposed to the stimuli. Parametric speakers offer a solution for transmitting directional audible signals by using ultrasonic carrier waves. The targeted transmission of vocal signals offers exciting opportunities for testing the diffusion of information in animal groups and mechanisms for resolving informational ambiguities. We have field tested the quality and directionality of a commercial parametric speaker, Soundlazer SL-01. Additionally, we assessed its usability for performing playback experiments by comparing behavioural responses of free-ranging meerkats (Suricata suricatta) with calls transmitted from conventional and parametric speakers. Our results show that the tested parametric speaker is highly directional. However, the acoustic structure of meerkat calls was strongly affected and low frequencies were not reliably reproduced by the parametric speaker. The playback trials elicited weakened behavioural responses probably due to the partial distortion of the signal but also indicating the potential importance of social facilitation for initiating mobbing events in meerkats. We conclude that parametric speakers can be useful tools for directed transmission of animals calls but after a careful assessment of signal fidelity.

The role of position in consensus dynamics of polarizable networks.

Communication constraints often complicate group decision-making. In this experiment, we investigate how the network position of opinionated group members determines both the speed and the outcome of group consensus in 7-member communication networks susceptible to polarization. To this end, we implemented an online version of a color coordination task within experimentally controlled communication networks. In 72 networks, one individual was incentivized to prefer one of two options. In 156 networks, two individuals were incentivized to prefer conflicting options. The network positions of incentivized individuals were varied. In networks with a single incentivized individual, network position played no significant role in either the speed or outcome of consensus decisions. For conflicts, the incentivized individual with more neighbors was more likely to sway the group to their preferred outcome. Furthermore, consensus emerged more slowly when the opponents had the same number of neighbors, but could not see each other's votes directly. These results suggest that the visibility of an opinion is key to wielding group influence, and that specific structures are sufficient to run communication networks into polarization, hindering a speedy consensus.

Disentangling influence over group speed and direction reveals multiple patterns of influence in moving meerkat groups.

Animals that travel together in groups must constantly come to consensus about both the direction and speed of movement, often simultaneously. Contributions to collective decisions may vary among group members, yet inferring who has influence over group decisions is challenging, largely due to the multifaceted nature of influence. Here we collected high-resolution GPS data from five habituated meerkat groups in their natural habitat during foraging and developed a method to quantify individual influence over both group direction and speed. We find that individual influence over direction and speed are correlated, but also exhibit substantial variation. Comparing patterns across social statuses reveals that dominant females have higher influence than other individuals over both group direction and speed. Individuals with high influence also tend to spend more time in the front of the group. We discuss our results in light of meerkat life-history and current literature on influence during group movement. Our method provides a general approach which can be applied to disentangle individual influence over group direction and speed in a wide range of species with cohesive movement, emphasizing the importance of integrating multiple lines of inquiry when inferring influence in moving animal groups.

Long-distance vocalizations of spotted hyenas contain individual, but not group, signatures.

In animal societies, identity signals are common, mediate interactions within groups, and allow individuals to discriminate group-mates from out-group competitors. However, individual recognition becomes increasingly challenging as group size increases and as signals must be transmitted over greater distances. Group vocal signatures may evolve when successful in-group/out-group distinctions are at the crux of fitness-relevant decisions, but group signatures alone are insufficient when differentiated within-group relationships are important for decision-making. Spotted hyenas are social carnivores that live in stable clans of less than 125 individuals composed of multiple unrelated matrilines. Clan members cooperate to defend resources and communal territories from neighbouring clans and other mega carnivores; this collective defence is mediated by long-range (up to 5 km range) recruitment vocalizations, called whoops. Here, we use machine learning to determine that spotted hyena whoops contain individual but not group signatures, and that fundamental frequency features which propagate well are critical for individual discrimination. For effective clan-level cooperation, hyenas face the cognitive challenge of remembering and recognizing individual voices at long range. We show that serial redundancy in whoop bouts increases individual classification accuracy and thus extended call bouts used by hyenas probably evolved to overcome the challenges of communicating individual identity at long distance.

A practical guide for generating unsupervised, spectrogram-based latent space representations of animal vocalizations.

BACKGROUND: The manual detection, analysis and classification of animal vocalizations in acoustic recordings is laborious and requires expert knowledge. Hence, there is a need for objective, generalizable methods that detect underlying patterns in these data, categorize sounds into distinct groups and quantify similarities between them. Among all computational methods that have been proposed to accomplish this, neighbourhood-based dimensionality reduction of spectrograms to produce a latent space representation of calls stands out for its conceptual simplicity and effectiveness. Goal of the study/what was done: Using a dataset of manually annotated meerkat Suricata suricatta vocalizations, we demonstrate how this method can be used to obtain meaningful latent space representations that reflect the established taxonomy of call types. We analyse strengths and weaknesses of the proposed approach, give recommendations for its usage and show application examples, such as the classification of ambiguous calls and the detection of mislabelled calls. What this means: All analyses are accompanied by example code to help researchers realize the potential of this method for the study of animal vocalizations.

Confirmation and variability of the Allee effect inDictyostelium discoideumcell populations, possible role of chemical signaling within cell clusters.

In studies of the unicellular eukaryoteDictyostelium discoideum, many have anecdotally observed that cell dilution below a certain 'threshold density' causes cells to undergo a period of slow growth (lag). However, little is documented about the slow growth phase and the reason for different growth dynamics below and above this threshold density. In this paper, we extend and correct our earlier work to report an extensive set of experiments, including the use of new cell counting technology, that set this slow-to-fast growth transition on a much firmer biological basis. We show that dilution below a certain density (around 104cells ml-1) causes cells to grow slower on average and exhibit a large degree of variability: sometimes a sample does not lag at all, while sometimes it takes many moderate density cell cycle times to recover back to fast growth. We perform conditioned media experiments to demonstrate that a chemical signal mediates this endogenous phenomenon. Finally, we argue that while simple models involving fluid transport of signal molecules or cluster-based signaling explain typical behavior, they do not capture the high degree of variability between samples but nevertheless favor an intra-cluster mechanism.

Vocal Turn-Taking in Meerkat Group Calling Sessions.

Coordination is a fundamental aspect of social living, underlying processes ranging from the maintenance of group cohesion to the avoidance of competition. Coordination can manifest as synchronization, where individuals perform the same action at the same time but can also take the form of anti-synchronization or turn-taking. Turn-taking has mainly been studied in the context of the development of language [1] due to the fact that it is a universal feature in all languages and has been found to appear early in infancy [2, 3]. Recently, turn-taking has received attention in animal communication research [4-7] as a potential foundation on which social communication was formed [1, 3]. In this study, we describe turn-taking in group-wide vocal interactions of meerkats (Suricata suricatta) during low-conflict sunning behavior, which is accompanied by the production of specific "sunning calls." We show that sunning-call production is socially stimulated and that at the group level, meerkats avoid overlap, thus fulfilling a key principle of turn-taking [8]. Through observational data and playback experiments, we show that these group-level patterns arise from two individual-level rules: call inhibition over short timescales, which prevents mutual interference, and call excitation over longer timescales, which stimulates further group calling. These simple rules suggest that hierarchy formation and turn allocation are not required for achieving group-wide coordination of communication. We also suggest that the potential bonding function of turn-taking shown in humans might have similar effects in animal interactions. VIDEO ABSTRACT.

Inferring influence and leadership in moving animal groups.

Collective decision-making is a daily occurrence in the lives of many group-living animals, and can have critical consequences for the fitness of individuals. Understanding how decisions are reached, including who has influence and the mechanisms by which information and preferences are integrated, has posed a fundamental challenge. Here, we provide a methodological framework for studying influence and leadership in groups. We propose that individuals have influence if their actions result in some behavioural change among their group-mates, and are leaders if they consistently influence others. We highlight three components of influence (influence instances, total influence and consistency of influence), which can be assessed at two levels (individual-to-individual and individual-to-group). We then review different methods, ranging from individual positioning within groups to information-theoretic approaches, by which influence has been operationally defined in empirical studies, as well as how such observations can be aggregated to give insight into the underlying decision-making process. We focus on the domain of collective movement, with a particular emphasis on methods that have recently been, or are being, developed to take advantage of simultaneous tracking data. We aim to provide a resource bringing together methodological tools currently available for studying leadership in moving animal groups, as well as to discuss the limitations of current methodologies and suggest productive avenues for future research.This article is part of the theme issue 'Collective movement ecology'.

Challenges and solutions for studying collective animal behaviour in the wild.

Mobile animal groups provide some of the most compelling examples of self-organization in the natural world. While field observations of songbird flocks wheeling in the sky or anchovy schools fleeing from predators have inspired considerable interest in the mechanics of collective motion, the challenge of simultaneously monitoring multiple animals in the field has historically limited our capacity to study collective behaviour of wild animal groups with precision. However, recent technological advancements now present exciting opportunities to overcome many of these limitations. Here we review existing methods used to collect data on the movements and interactions of multiple animals in a natural setting. We then survey emerging technologies that are poised to revolutionize the study of collective animal behaviour by extending the spatial and temporal scales of inquiry, increasing data volume and quality, and expediting the post-processing of raw data.This article is part of the theme issue 'Collective movement ecology'.

Habitat and social factors shape individual decisions and emergent group structure during baboon collective movement.

For group-living animals traveling through heterogeneous landscapes, collective movement can be influenced by both habitat structure and social interactions. Yet research in collective behavior has largely neglected habitat influences on movement. Here we integrate simultaneous, high-resolution, tracking of wild baboons within a troop with a 3-dimensional reconstruction of their habitat to identify key drivers of baboon movement. A previously unexplored social influence - baboons' preference for locations that other troop members have recently traversed - is the most important predictor of individual movement decisions. Habitat is shown to influence movement over multiple spatial scales, from long-range attraction and repulsion from the troop's sleeping site, to relatively local influences including road-following and a short-range avoidance of dense vegetation. Scaling to the collective level reveals a clear association between habitat features and the emergent structure of the group, highlighting the importance of habitat heterogeneity in shaping group coordination.

Both Nearest Neighbours and Long-term Affiliates Predict Individual Locations During Collective Movement in Wild Baboons.

In many animal societies, groups of individuals form stable social units that are shaped by well-delineated dominance hierarchies and a range of affiliative relationships. How do socially complex groups maintain cohesion and achieve collective movement? Using high-resolution GPS tracking of members of a wild baboon troop, we test whether collective movement in stable social groups is governed by interactions among local neighbours (commonly found in groups with largely anonymous memberships), social affiliates, and/or by individuals paying attention to global group structure. We construct candidate movement prediction models and evaluate their ability to predict the future trajectory of focal individuals. We find that baboon movements are best predicted by 4 to 6 neighbours. While these are generally individuals' nearest neighbours, we find that baboons have distinct preferences for particular neighbours, and that these social affiliates best predict individual location at longer time scales (>10 minutes). Our results support existing theoretical and empirical studies highlighting the importance of local rules in driving collective outcomes, such as collective departures, in primates. We extend previous studies by elucidating the rules that maintain cohesion in baboons 'on the move', as well as the different temporal scales of social interactions that are at play.

Estimating uncertainty and reliability of social network data using Bayesian inference.

Social network analysis provides a useful lens through which to view the structure of animal societies, and as a result its use is increasingly widespread. One challenge that many studies of animal social networks face is dealing with limited sample sizes, which introduces the potential for a high level of uncertainty in estimating the rates of association or interaction between individuals. We present a method based on Bayesian inference to incorporate uncertainty into network analyses. We test the reliability of this method at capturing both local and global properties of simulated networks, and compare it to a recently suggested method based on bootstrapping. Our results suggest that Bayesian inference can provide useful information about the underlying certainty in an observed network. When networks are well sampled, observed networks approach the real underlying social structure. However, when sampling is sparse, Bayesian inferred networks can provide realistic uncertainty estimates around edge weights. We also suggest a potential method for estimating the reliability of an observed network given the amount of sampling performed. This paper highlights how relatively simple procedures can be used to estimate uncertainty and reliability in studies using animal social network analysis.