Exploring the role of vocalizations in regulating group dynamics.

Because of the diverging needs of individuals, group life can lead to disputes and competition, but it also has many advantages, such as reduced predation risk, information sharing and increased hunting success. Social animals have to maintain group cohesion and need to synchronize activities, such as foraging, resting, social interactions and movements, in order to thrive in groups. Acoustic signals are highly relevant for social dynamics, some because they are long-ranging and others because they are short-ranging, which may serve important within-group functions. However, although there has been an increase in studies concentrating on acoustic communication within groups in the past decade, many aspects of how vocalizations relate to group dynamics are still poorly understood. The aim of this review is to present an overview of our current knowledge on the role of vocalizations in regulating social group dynamics, identify knowledge gaps and recommend potential future research directions. We review the role that vocalizations play in (i) collective movement, (ii) separation risk and cohesion maintenance, (iii) fission-fusion dynamics, and (iv) social networks. We recommend that future studies aim to increase the diversity of studied species and strengthen the integration of state-of-the-art tools to study social dynamics and acoustic signals. This article is part of the theme issue 'The power of sound: unravelling how acoustic communication shapes group dynamics'.

Consistent Individual Differences Drive Collective Movements in a Tibetan Macaque Group.

Collective movement has emerged as a key area of interest in animal behavior. While individual differences are often viewed as a potential threat to group cohesion, growing evidence suggests that these differences can actually influence an animal's behavior as an initiator or follower during collective movements, thereby driving the group's movement and decision-making processes. To resolve the divergence, we asked how personality can affect the dynamics of collective movements in one group of free-ranging Tibetan macaques (Macaca thibetana) in Huangshan, China. We assessed individual personality using principal component analysis and applied the generalized linear mixed model and linear mixed model to examine the influence of personality on decision making during collective movements. Our findings reveled three distinct personality types among Tibetan macaques: sociability, boldness, and anxiousness. Individuals with higher sociability scores and rank, or those with lower anxiousness scores, were more likely to initiate successful collective movements. Older individuals were less successful in initiating movements compared to young adults. Leaders with lower anxiousness scores or higher rank attracted more followers, with females attracting larger groups than males. As for followers, individuals with higher rank tended to join the collective movement earlier. Additionally, individuals with higher sociability or boldness scores had shorter joining latency in collective movement. Finally, there was a longer joining latency for middle-aged adults compared to young adults. These results suggest that individual differences are a potential driver of collective movements. We provide some insights into the relationships between personality and decision making in Tibetan macaques.

Leadership and the finder's advantage in mantled howler monkeys.

Leading collective movements and arriving first at feeding sites may improve food acquisition. Specifically, the first individual to discover and exploit a feeding site may gain a feeding advantage known as the "finder's advantage." The aim of this research was to verify if the probability of leading group movements to feeding sites in mantled howler monkeys (Alouatta palliata) varied by sex and reproductive status, and whether finders had higher foraging success than followers. We studied 18 adult individuals from two groups in La Flor de Catemaco over a year (978 h), and sampled group movements (n = 211) and foraging behavior (n = 215 feeding episodes). Gestating females were leaders and finders of group movements to feeding sites more often than expected but were also replaced in the leading position more frequently than individuals of other sex/reproductive states. Feeding behavior was not influenced by the order of arrival at feeding sites per se, but gestating females had higher food intake rate, bite rate, and feeding time when arriving earlier (i.e., occupying front group positions) than later. Therefore, leadership and the finder's advantage occur in this species and are probably employed by gestating females to maintain their energetic condition.

Rushing for "burned" food: Why and how does a group of patas monkeys (Erythrocebus patas) reach freshly burned areas?

Recently, considerable attention has been paid to animal adaptations to anthropogenic environments, such as foraging in burned areas where plants are promoted to regenerate by anthropogenic burning. However, among primates, reports on the utilization of resources that are available immediately after burning have been limited to a few primate species. In this study, we investigated and compared the activity budgets and food categories of a group of patas monkeys (Erythrocebus patas) in freshly burned areas by comparing them with those in previously burned areas and unburned areas. We also assessed the proportion of time spent in the freshly burned area before and after the fire: GPS collars were fitted to five of the six adults in the group, and their patterns when they traveled toward freshly burned and unburned feeding areas were compared. Patas monkeys spent more time in freshly burned areas after the fire, and they visited such areas mostly for feeding, particularly on roasted seeds of Cissus populnea. Furthermore, patas monkeys traveled faster and in a more synchronized way toward freshly burned areas. This "apparent goal-directed" travel began at least 1 h before arriving. Results indicate that the group recognized freshly burned areas as valuable, and the monkeys were able to travel in a goal-directed manner to them despite their variable locations. We suggest that smoke from freshly burned areas provides a visual cue with which to orient to the burned areas. Our results also support the notion that some primates are flexible enough to adapt to and benefit from anthropogenic environmental changes.

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.

Nearest Neighbour Node Deployment Algorithm for Mobile Sensor Networks.

Many animal aggregations display remarkable collective coordinated movements on a large scale, which emerge as a result of distributed local decision-making by individuals. The recent advances in modelling the collective motion of animals through the utilisation of Nearest Neighbour rules, without the need for centralised coordination, resulted in the development of self-deployment algorithms in Mobile Sensor Networks (MSNs) to achieve various types of coverage essential for different applications. However, the energy consumption associated with sensor movement to achieve the desired coverage remains a significant concern for the majority of algorithms reported in the literature. In this paper, the Nearest Neighbour Node Deployment (NNND) algorithm is proposed to efficiently provide blanket coverage across a given area while minimising energy consumption and enhancing fault tolerance. In contrast to other algorithms that sequentially move sensors, NNND leverages the power of parallelism by employing multiple streams of sensor motions, each directed towards a distinct section of the area. The cohesion of each stream is maintained by adaptively choosing a leader for each stream while collision avoidance is also ensured. These properties contribute to minimising the travel distance within each stream, resulting in decreased energy consumption. Additionally, the utilisation of multiple leaders in NNND eliminates the presence of a single point of failure, hence enhancing the fault tolerance of the area coverage. The results of our extensive simulation study demonstrate that NNND not only achieves lower energy consumption but also a higher percentage of k-coverage.

Social situations differ in their contribution to population-level social structure in griffon vultures.

Social relationships among animals emerge from interactions in multiple ecological and social situations. However, we seldom ask how each situation contributes to the global structure of a population, and whether different situations contribute different information about social relationships and the position of individuals within the social fabric. Griffon vultures (Gyps fulvus) interact socially in multiple situations, including communal roosting, joint flights, and co-feeding. These social interactions can influence population-level outcomes, such as disease transmission and information sharing that determine survival and response to changes. We examined the unique contribution of each social and ecological situation to the social structure of the population and individuals' positions within the overall social network using high-resolution GPS tracking. We found that the number of individuals each vulture interacted with (degree) was best predicted by diurnal interactions-both during flights and on the ground (such as when feeding). However, the strength of social bonds, that is, the number of interactions an individual had (strength), was best predicted by interactions on the ground-both during the day (e.g., while feeding) and at night (e.g., while roosting) but not by interactions while flying. Thus, social situations differ in their impact on the relationships that individuals form. By incorporating the ecological situations in which social interactions occur we gain a more complete view of how social relationships are formed and which situations are important for different types of interactions.

Inferring the interaction rules of complex systems with graph neural networks and approximate Bayesian computation.

Inferring the underlying processes that drive collective behaviour in biological and social systems is a significant statistical and computational challenge. While simulation models have been successful in qualitatively capturing many of the phenomena observed in these systems in a variety of domains, formally fitting these models to data remains intractable. Recently, approximate Bayesian computation (ABC) has been shown to be an effective approach to inference if the likelihood function for a model is unavailable. However, a key difficulty in successfully implementing ABC lies with the design, selection and weighting of appropriate summary statistics, a challenge that is especially acute when modelling high dimensional complex systems. In this work, we combine a Gaussian process accelerated ABC method with the automatic learning of summary statistics via graph neural networks. Our approach bypasses the need to design a model-specific set of summary statistics for inference. Instead, we encode relational inductive biases into a neural network using a graph embedding and then extract summary statistics automatically from simulation data. To evaluate our framework, we use a model of collective animal movement as a test bed and compare our method to a standard summary statistics approach and a linear regression-based algorithm.

Behavioural ecology at the spatial-social interface.

Spatial and social behaviour are fundamental aspects of an animal's biology, and their social and spatial environments are indelibly linked through mutual causes and shared consequences. We define the 'spatial-social interface' as intersection of social and spatial aspects of individuals' phenotypes and environments. Behavioural variation at the spatial-social interface has implications for ecological and evolutionary processes including pathogen transmission, population dynamics, and the evolution of social systems. We link spatial and social processes through a foundation of shared theory, vocabulary, and methods. We provide examples and future directions for the integration of spatial and social behaviour and environments. We introduce key concepts and approaches that either implicitly or explicitly integrate social and spatial processes, for example, graph theory, density-dependent habitat selection, and niche specialization. Finally, we discuss how movement ecology helps link the spatial-social interface. Our review integrates social and spatial behavioural ecology and identifies testable hypotheses at the spatial-social interface.

Field bands of marching locust juveniles show carbohydrate, not protein, limitation.

Locusts are grasshoppers that migrate en masse and devastate food security, yet little is known about the nutritional needs of marching bands in nature. While it has been hypothesized that protein limitation promotes locust marching behavior, migration is fueled by dietary carbohydrates. We studied South American Locust (Schistocerca cancellata) bands at eight sites across Argentina, Bolivia, and Paraguay. Bands ate most frequently from dishes containing carbohydrate artificial diets and minimally from balanced, protein, or control (vitamins and salts) dishes-indicating carbohydrate hunger. This hunger for carbohydrates is likely explained by the observation that local vegetation was generally protein-biased relative to locusts' preferred protein to carbohydrate ratio. This study highlights the importance of studying the nutritional ecology of animals in their environment and suggests that carbohydrate limitation may be a common pattern for migrating insect herbivores.

CoFee-L: A Model of Animal Displacement in Large Groups Combining Cohesion Maintenance, Feeding Area Search and Transient Leadership.

In the tropics, the conservation of tree species is closely linked to that of animals, as a large proportion of trees are zoochoric and therefore rely on the movement of animals to disperse their seeds in order to increase the chances of the survival of progeny and to allow migration in the face of climate change. Research into the prediction of animal movements is therefore critical but has so far focused only on particular features of collective behavior. In contrast, we included the concepts of cohesion maintenance, feeding area search and transient leadership in a single model, CoFee-L, and tested it to simulate the movement of a wild-ranging troop of primates (Macaca leonina). We analyzed and compared observations and simulations with a statistical physics tool (mean squared displacement) and with histograms and χ2 (for the step length and turning angle distributions). CoFee-L allowed us to simulate the physical properties of the troop's center of mass trajectory as well as the step length and angle distributions of the field data. The parametrization of CoFee-L was rather straightforward, as it was sufficient to fix a set of parameters easily observable in the field and then to adjust the values of four parameters that have biological meaning.

Alignment with neighbours enables escape from dead ends in flocking models.

Coordinated movement in animal groups (flocks, schools, herds, etc.) is a classic and well-studied form of collective behaviour. Most theoretical studies consider agents in unobstructed spaces; however, many animals move in often complicated environments and must navigate around and through obstacles. Here we consider simulated agents behaving according to typical flocking rules, with the addition of repulsion from obstacles, and study their collective behaviour in environments with concave obstacles (dead ends). We find that groups of such agents heading for a goal can spontaneously escape dead ends without wall-following or other specialized behaviours, in what we term 'flocking escapes'. The mechanism arises when agents align with one another while heading away from the goal, forming a self-stable cluster that persists long enough to exit the obstacle and avoids becoming trapped again when turning back towards the goal. Solitary agents under the same conditions are never observed to escape. We show that alignment with neighbours reduces the effective turning speed of the group while letting individuals maintain high manoeuvrability when needed. The relative robustness of flocking escapes in our studies suggests that this emergent behaviour may be relevant for a variety of animal species.

Does a high social status confer greater levels of trust from groupmates? An experimental study of leadership in domestic horses.

In collective movements, specific individuals may emerge as leaders. In this study on the domestic horse (Equus ferus caballus), we conducted experiments to establish if an individual is successfully followed due to its social status (including hierarchical rank and centrality). We first informed one horse about a hidden food location and recorded by how many it was followed when going back to this location. In this context, all horses lead their groupmates successfully. In a second step, we tested whether group members would trust some leaders more than others by removing the food before the informed individual led the group back to the food location. In addition, two control initiators with intermediate social status for which the food was not removed were tested. The results, confirmed by simulations, demonstrated that the proportions of followers for the unreliable initiator with highest social status are greater than the ones of the unreliable initiator with lowest social status. Our results suggest an existing relationship between having a high social status and a leadership role. Indeed, the status of a leader sometimes prevail at the detriment of the accuracy of the information, because an elevated social status apparently confers a high level of trust.

Come with me: experimental evidence for intentional recruitment in Tonkean macaques.

Recruitment is a process by which animals can initiate collective movements: the action of an individual prompts conspecifics to follow. Although it has been hypothesized that animals may be able to intentionally recruit others, there is no experimental evidence of this to date. We tested this hypothesis in two pairs of Tonkean macaques in a situation requiring the subjects to find a food site in a 2800 m2 area, and approach the site together to release rewards. Each subject was informed of the location of either highly or little-valued rewards. We recorded attention-action sequences in which an individual checked that his partner was attending to him before moving, and also simple departures (i.e., not preceded by eye contact). Analyses showed that sequences were more often followed by recruitment and leading the partner to a baited site than simple departures were. Moreover, subjects used attention-action sequences more frequently when informed of the location of the highly valued reward. This may be explained by the fact that the more motivated they were by the expected rewards, the more likely they were to actively recruit their partner. No such effect was found when subjects performed simple departures. We conclude that Tonkean macaques are capable of intentional recruitment because the subjects voluntarily behaved with the goal of influencing their partner's movement: they checked that the partner was paying attention to them and prompted him to follow by moving. Such performances can be accounted for either by associative learning or by intentional communication.

Collective movement decision-making in primates in crop-raiding contexts.

Crop-raiding by wildlife species often involves collective group movement and animal decision-making in this context is an important area of investigation as the risks and rewards associated with crop-raiding are greater than those that are likely to occur in wild food foraging situations. Yet, the form of consensus decision-making involved in wildlife crop-raiding has not been evaluated. In the current study, we assessed the decision-making process exhibited by rhesus macaques, a generalist primate species, in the context of crop-raiding. We predicted that rhesus macaques would display unshared consensus decision-making during the higher risk-entailed crop-raiding collective movements. We followed two groups of rhesus macaques and collected data on their movement and foraging behavior during crop-raiding events. The results from our study showed that collective movements in the context of crop-raiding were largely led by single individuals. However, individuals of all age-sex categories successfully led crop-raiding collective movements. Although adult individuals joined adult-initiated raid movements most often, they also joined raids initiated by other age classes. Contrary to our predictions, in crop-raiding contexts, rhesus macaques demonstrated a tendency towards equally shared decision-making for group movements.

Using Live Imaging to Examine Early Cardiac Development in Zebrafish.

Visualizing dynamic cellular behaviors using live imaging is critical to the study of cell movement and to the study of cellular and embryonic polarity. Similarly, live imaging can be vital to elucidating the pathology of genetic disorders and diseases. Model systems such as zebrafish, whose in vivo development is accessible to both the microscope and genetic manipulation, are particularly well-suited to the use of live imaging. Here we describe an overall approach to conducting live-imaging experiments with a specific emphasis on investigating cell movements during the early stages of heart development in zebrafish.

Defending in 4-4-2 or 5-3-2 formation? small differences in footballers' collective tactical behaviours.

This study explored footballers' tactical behaviours, based on their position data, as an effect of two defending formations, 4-4-2 and 5-3-2, using an experimental approach. Sixty-nine youth footballers participated in this 11-versus-11 study, performing 72 trials of attack-versus-defence. Players' position data were tracked using a local positioning system, and processed to calculate measures of collective movement. This was supplemented by the analysis of passing networks. The results showed small differences between the two conditions. Compared to a 4-4-2 formation, defending in 5-3-2 reduced dispersion (-0.69 m,p=0.012), midfield-forward distance (-0.81 m, p=0.047), and defence-forward distance (-1.29 m, p=0.038); the consequent effects on attacking teams included reduced team widths (-1.78 m, p=0.034), reduced necessity for back-passes to the goalkeeper, and less connectivity in the passing network. The effects of the two defending formations seem to have the greatest impact on fullbacks of the attacking teams, since they were main contributors of the reduced team widths, received more passes, and had higher betweenness centrality in the right-back position during 5-3-2 defending. In summary, the present study potentially demonstrates how the underlying mechanisms in players' collective movements and passing behaviours show that the 5-3-2 is more conservatively defensive than the 4-4-2.

Movement, Encounter Rate, and Collective Behavior in Ant Colonies.

Spatial patterns of movement regulate many aspects of social insect behavior, because how workers move around, and how many are there, determines how often they meet and interact. Interactions are usually olfactory; for example, in ants, by means of antennal contact in which one worker assesses the cuticular hydrocarbons of another. Encounter rates may be a simple outcome of local density: a worker experiences more encounters, the more other workers there are around it. This means that encounter rate can be used as a cue for overall density even though no individual can assess global density. Encounter rate as a cue for local density regulates many aspects of social insect behavior, including collective search, task allocation, nest choice, and traffic flow. As colonies grow older and larger, encounter rates change, which leads to changes in task allocation. Nest size affects local density and movement patterns, which influences encounter rate, so that nest size and connectivity influence colony behavior. However, encounter rate is not a simple function of local density when individuals change their movement in response to encounters, thus influencing further encounter rates. Natural selection on the regulation of collective behavior can draw on variation within and among colonies in the relation of movement patterns, encounter rate, and response to encounters.

Seasonality impacts collective movements in a wild group-living bird.

BACKGROUND: A challenge faced by animals living in groups with stable long-term membership is to effectively coordinate their actions and maintain cohesion. However, as seasonal conditions alter the distribution of resources across a landscape, they can change the priority of group members and require groups to adapt and respond collectively across changing contexts. Little is known about how stable group-living animals collectively modify their movement behaviour in response to environment changes, such as those induced by seasonality. Further, it remains unclear how environment-induced changes in group-level movement behaviours might scale up to affect population-level properties, such as a population's footprint. METHODS: Here we studied the collective movement of each distinct social group in a population of vulturine guineafowl (Acryllium vulturinum), a largely terrestrial and non-territorial bird. We used high-resolution GPS tracking of group members over 22 months, combined with continuous time movement models, to capture how and where groups moved under varying conditions, driven by seasonality and drought. RESULTS: Groups used larger areas, travelled longer distances, and moved to new places more often during drier seasons, causing a three-fold increase in the area used at the population level when conditions turned to drought. By contrast, groups used smaller areas with more regular movements during wetter seasons. CONCLUSIONS: The consistent changes in collective outcomes we observed in response to different environments raise questions about the role of collective behaviour in facilitating, or impeding, the capacity for individuals to respond to novel environmental conditions. As droughts will be occurring more often under climate change, some group living animals may have to respond to them by expressing dramatic shifts in their regular movement patterns. These shifts can have consequences on their ranging behaviours that can scale up to alter the footprints of animal populations.

Spatio-temporal variation in the wintering associations of an alpine bird.

Many animals make behavioural changes to cope with winter conditions, being gregariousness a common strategy. Several factors have been invoked to explain why gregariousness may evolve during winter, with individuals coming together and separating as they trade off the different costs and benefits of living in groups. These trade-offs may, however, change over space and time as a response to varying environmental conditions. Despite its importance, little is known about the factors triggering gregarious behaviour during winter and its change in response to variation in weather conditions is poorly documented. Here, we aimed at quantifying large-scale patterns in wintering associations over 23 years of the white-winged snowfinch Montifringilla nivalis nivalis. We found that individuals gather in larger groups at sites with harsh wintering conditions. Individuals at colder sites reunite later and separate earlier in the season than at warmer sites. However, the magnitude and phenology of wintering associations are ruled by changes in weather conditions. When the temperature increased or the levels of precipitation decreased, group size substantially decreased, and individuals stayed united in groups for a shorter time. These results shed light on factors driving gregariousness and points to shifting winter climate as an important factor influencing this behaviour.