The relative importance of social information use for population abundance in group-living and non-grouping prey.

Predator-prey relationships are fundamental components of ecosystem functioning, within which the spatial consequences of prey social organization can alter predation rates. Group-living (GL) species are known to exploit inadvertent social information (ISI) that facilitates population persistence under predation risk. Still, the extent to which non-grouping (NG) prey can benefit from similar processes is unknown. Here we built an individual-based model to explore and compare the population-level consequences of ISI use in GL and NG prey. We differentiated between GL and NG prey only by the presence or absence of social attraction toward conspecifics that drives individual movement patterns. We found that the extent of the benefits of socially acquired predator information in NG highly depends on the prey's ability to detect nearby predators, prey density and the occurrence of false alarms. Conversely, even moderate probabilities of ISI use and predator detection can lead to maximal population-level benefits in GL prey. This theoretical work provides additional insights into the conditions under which ISI use can facilitate population persistence irrespective of prey social organisation.

Alarm communication networks as a driver of community structure in African savannah herbivores.

Social information networks have the potential to shape the spatial structure of ecological communities by promoting the formation of mixed-species groups. However, what actually drives social affinity between species in the wild will depend on the characteristics of the species available to group. Here we first present an agent-based model that predicts trait-related survival benefits from mixed-species group formation in a multi-species community and we then test the model predictions in a community-wide field study of African savannah herbivores using multi-layered network analysis. We reveal benefits from information transfer about predators as a key determinant of mixed-species group formation, and that dilution benefits alone are not enough to explain patterns in interspecific sociality. The findings highlight the limitations of classical ecological approaches focusing only on direct trophic interactions when analysing community structure and suggest that declines in species occupying central social network positions, such as key informants, can have significant repercussions throughout communities.

Strategies for integrating disparate social information.

Social information use is widespread in the animal kingdom, helping individuals rapidly acquire useful knowledge and adjust to novel circumstances. In humans, the highly interconnected world provides ample opportunities to benefit from social information but also requires navigating complex social environments with people holding disparate or conflicting views. It is, however, still largely unclear how people integrate information from multiple social sources that (dis)agree with them, and among each other. We address this issue in three steps. First, we present a judgement task in which participants could adjust their judgements after observing the judgements of three peers. We experimentally varied the distribution of this social information, systematically manipulating its variance (extent of agreement among peers) and its skewness (peer judgements clustering either near or far from the participant's judgement). As expected, higher variance among peers reduced their impact on behaviour. Importantly, observing a single peer confirming a participant's own judgement markedly decreased the influence of other-more distant-peers. Second, we develop a framework for modelling the cognitive processes underlying the integration of disparate social information, combining Bayesian updating with simple heuristics. Our model accurately accounts for observed adjustment strategies and reveals that people particularly heed social information that confirms personal judgements. Moreover, the model exposes strong inter-individual differences in strategy use. Third, using simulations, we explore the possible implications of the observed strategies for belief updating. These simulations show how confirmation-based weighting can hamper the influence of disparate social information, exacerbate filter bubble effects and deepen group polarization. Overall, our results clarify what aspects of the social environment are, and are not, conducive to changing people's minds.

Diurnal variation in the production of vocal information about food supports a model of social adjustment in wild songbirds.

Wintering songbirds have been widely shown to make economic foraging decisions to manage the changing balance of risks from predation and starvation over the course of the day. In this study, we ask whether the communication and use of information about food availability differ throughout the day. First, we assessed temporal variation in food-related vocal information produced in foraging flocks of tits ( Paridae) using audio recordings at radio-frequency identification-equipped feeding stations. Vocal activity was highest in the morning and decreased into the afternoon. This pattern was not explained by there being fewer birds present, as we found that group sizes increased over the course of the day. Next, we experimentally tested the underlying causes for this diurnal calling pattern. We set up bird feeders with or without playback of calls from tits, either in the morning or in the afternoon, and compared latency to feeder discovery, accumulation of flock members, and total number of birds visiting the feeder. Irrespective of time of day, playbacks had a strong effect on all three response measures when compared to silent control trials, demonstrating that tits will readily use vocal information to improve food detection throughout the day. Thus, the diurnal pattern of foraging behaviour did not appear to affect use and production of food-related vocalizations. Instead, we suggest that, as the day progresses and foraging group sizes increase, the costs of producing calls at the food source (e.g. competition and attraction of predators) outweigh the benefits of recruiting group members (i.e. adding individuals to large groups only marginally increases safety in numbers), causing the observed decrease in vocal activity into the afternoon. Our findings imply that individuals make economic social adjustments based on conditions of their social environment when deciding to vocally recruit group members.

Evidence of sociality in the timing and location of foraging in a colonial seabird.

Social foraging behaviours, which range from cooperative hunting to local enhancement, can result in increased prey capture and access to information, which may significantly reduce time and energy costs of acquiring prey. In colonial species, it has been proposed that the colony itself may act as a site of social information transfer and group formation. However, conclusive evidence from empirical studies is lacking. In particular, most studies in colonial species have generally focussed on behaviours either at the colony or at foraging sites in isolation, and have failed to directly connect social associations at the colony to social foraging. In this study, we simultaneously tracked 85% of a population of Australasian gannets (Morus serrator) over multiple foraging trips, to study social associations at the colony and test whether these associations influence the location of foraging sites. We found that gannets positively associate with conspecifics while departing from the colony and that co-departing gannets have more similar initial foraging patches than individuals that did not associate at the colony. These results provide strong evidence for the theory that the colony may provide a source of information that influences foraging location.

A frontopolar-temporal circuit determines the impact of social information in macaque decision making.

When choosing, primates are guided not only by personal experience of objects but also by social information such as others' attitudes toward the objects. Crucially, both sources of information-personal and socially derived-vary in reliability. To choose optimally, one must sometimes override choice guidance by personal experience and follow social cues instead, and sometimes one must do the opposite. The dorsomedial frontopolar cortex (dmFPC) tracks reliability of social information and determines whether it will be attended to guide behavior. To do this, dmFPC activity enters specific patterns of interaction with a region in the mid-superior temporal sulcus (mSTS). Reversible disruption of dmFPC activity with transcranial ultrasound stimulation (TUS) led macaques to fail to be guided by social information when it was reliable but to be more likely to use it when it was unreliable. By contrast, mSTS disruption uniformly downregulated the impact of social information on behavior.

Brain structure correlates of social information use: an exploratory machine learning approach.

Introduction: Individual differences in social learning impact many important decisions, from voting behavior to polarization. Prior research has found that there are consistent and stable individual differences in social information use. However, the underlying mechanisms of these individual differences are still poorly understood. Methods: We used two complementary exploratory machine learning approaches to identify brain volumes related to individual differences in social information use. Results and discussion: Using lasso regression and random forest regression we were able to capture linear and non-linear brain-behavior relationships. Consistent with previous studies, our results suggest there is a robust positive relationship between the volume of the left pars triangularis and social information use. Moreover, our results largely overlap with common social brain network regions, such as the medial prefrontal cortex, superior temporal sulcus, temporal parietal junction, and anterior cingulate cortex. Besides, our analyses also revealed several novel regions related to individual differences in social information use, such as the postcentral gyrus, the left caudal middle frontal gyrus, the left pallidum, and the entorhinal cortex. Together, these results provide novel insights into the neural mechanisms that underly individual differences in social learning and provide important new leads for future research.

The evolution of eavesdropping on heterospecific alarm calls: Relevance, reliability, and personal information.

Interceptive eavesdropping on the alarm calls of heterospecifics provides crucial information about predators. Previous research suggests predator discrimination, call relevance, reliability, and reception explain when eavesdropping will evolve. However, there has been no quantitative analysis to scrutinize these principles, or how they interact. We develop a mathematical framework that formalizes the study of the key principles thought to select for eavesdropping. Interceptive eavesdropping appears to be greatly affected by the threat faced by caller and eavesdropper, as well as presence of informational noise affecting the detection of calls and predators. Accordingly, our model uses signal detection theory to examine when selection will favor alarm calling by a sender species and fleeing by an eavesdropping receiver species. We find eavesdropping is most strongly selected when (1) the receiver faces substantial threats, (2) species are ecologically similar, (3) senders often correctly discriminate threats, (4) receivers often correctly perceive calls, and (5) the receiver's personal discrimination of threats is poor. Furthermore, we find (6) that very high predation levels can select against eavesdropping because prey cannot continuously flee and must conserve energy. Reliability of heterospecific calls for identifying threats is thought to be important in selecting for eavesdropping. Consequently, we formally define reliability, showing its connection to specificity and sensitivity, clarifying how these quantities can be measured. We find that high call relevance, due to similar vulnerability to predators between species, strongly favors eavesdropping. This is because senders trade-off false alarms and missed predator detections in a way that is also favorable for the eavesdropper, by producing less of the costlier error. Unexpectedly, highly relevant calls increase the total number of combined errors and so have lower reliability. Expectedly, when noise greatly affects personally gathered cues to threats, but not heterospecific calls or detection of predators, eavesdropping is favored.

From whom do animals learn? A meta-analysis on model-based social learning.

Social learning via the observation of or interaction with other individuals can allow animals to obtain information about the local environment. Once social information is obtained, animals may or may not act on and use this information. Animals may learn from others selectively based on particular characteristics (e.g., familiarity, age, dominance) of the information provider, which is thought to maximize the benefits of social learning. Biases to copy certain individuals over others plays an important role in how information is transmitted and used among individuals, and can influence the emergence of group-level behaviors (i.e., traditions). Two underlying factors can affect from whom animals learn: the population social dynamics - with whom you associate (e.g., familiar), and status of the demonstrator (e.g., dominant). We systematically surveyed the literature and conducted a meta-analysis to test whether demonstrator characteristics consistently influence social learning, and if social dynamics strategies differ from status strategies in their influence on social learning. We extracted effect sizes from papers that used an observer-demonstrator paradigm to test if the characteristics of the individual providing social information (i.e., the demonstrator) influence social information use by observers. We obtained 139 effect sizes on 33 species from 54 experiments. First, we found an effect of experimental design on the influence of demonstrator characteristics on social learning: between-subject designs had stronger effects compared to within-subject designs. Second, we found that demonstrator characteristics do indeed influence social learning. Characteristics based on social dynamics and characteristics based on status had a significant effect on social learning, especially when copying familiar and kin demonstrators. These results highlight the role that demonstrator characteristics play on social learning, which can have implications for the formation and establishment of behavioural traditions in animals.

Conceptual preferences can be transmitted via selective social information use between competing wild bird species.

Concept learning is considered a high-level adaptive ability. Thus far, it has been studied in laboratory via asocial trial and error learning. Yet, social information use is common among animals but it remains unknown whether concept learning by observing others occurs. We tested whether pied flycatchers (Ficedula hypoleuca) form conceptual relationships from the apparent choices of nest-site characteristics (geometric symbol attached to the nest-box) of great tits (Parus major). Each wild flycatcher female (n = 124) observed one tit pair that exhibited an apparent preference for either a large or a small symbol and was then allowed to choose between two nest-boxes with a large and a small symbol, but the symbol shape was different to that on the tit nest. Older flycatcher females were more likely to copy the symbol size preference of tits than yearling flycatcher females when there was a high number of visible eggs or a few partially visible eggs in the tit nest. However, this depended on the phenotype, copying switched to rejection as a function of increasing body size. Possibly the quality of and overlap in resource use with the tits affected flycatchers' decisions. Hence, our results suggest that conceptual preferences can be horizontally transmitted across coexisting animals, which may increase the performance of individuals that use concept learning abilities in their decision-making.

Viewing animal migration through a social lens.

Evidence of social learning is growing across the animal kingdom. Researchers have long hypothesized that social interactions play a key role in many animal migrations, but strong empirical support is scarce except in a few unique systems and species. In this review, we aim to catalyze advances in the study of social migrations by synthesizing research across disciplines and providing a framework for understanding when, how, and why social influences shape the decisions animals make during migration. Integrating research across the fields of social learning and migration ecology will advance our understanding of the complex behavioral phenomena of animal migration and help to inform conservation of animal migrations in a changing world.

Stress signalling in acellular slime moulds and its detection by conspecifics.

Unicellular organisms live in unpredictable environments. Therefore, they need to continuously assess environmental conditions and respond appropriately to survive and thrive. When subjected to rapid changes in their environment or to cellular damages, unicellular organisms such as bacteria exhibit strong physiological reactions called stress responses that can be sensed by conspecifics. The ability to detect and use stress-related cues released by conspecifics to acquire information about the environment constitutes an adaptive survival response by prompting the organism to avoid potential dangers. Here, we investigate stress signalling and its detection by conspecifics in a unicellular organism, Physarum polycephalum. Slime moulds were subjected to either biotic (i.e. nutritional) or abiotic (i.e. chemical and light) stressors or left undisturbed while they were exploring a homogeneous environment. Then, we observed the responses of slime moulds facing a choice between cues released by stressed clone mates and cues released by undisturbed ones. We found that slime moulds actively avoided environments previously explored by stressed clone mates. These results suggest that slime moulds, like bacteria or social amoeba, exhibit physiological responses to biotic and abiotic stresses that can be sensed by conspecifics. Our results establish slime moulds as a promising new model to investigate the use of social information in unicellular organisms. This article is part of the theme issue 'Signal detection theory in recognition systems: from evolving models to experimental tests'.

The signal detection problem of aposematic prey revisited: integrating prior social and personal experience.

Ever since Alfred R. Wallace suggested brightly coloured, toxic insects warn predators about their unprofitability, evolutionary biologists have searched for an explanation of how these aposematic prey evolve and are maintained in natural populations. Understanding how predators learn about this widespread prey defence is fundamental to addressing the problem, yet individuals differ in their foraging decisions and the predominant application of associative learning theory largely ignores predators' foraging context. Here we revisit the suggestion made 15 years ago that signal detection theory provides a useful framework to model predator learning by emphasizing the integration of prior information into predation decisions. Using multiple experiments where we modified the availability of social information using video playback, we show that personal information (sampling aposematic prey) improves how predators (great tits, Parus major) discriminate between novel aposematic and cryptic prey. However, this relationship was not linear and beyond a certain point personal encounters with aposematic prey were no longer informative for prey discrimination. Social information about prey unpalatability reduced attacks on aposematic prey across learning trials, but it did not influence the relationship between personal sampling and discrimination. Our results suggest therefore that acquiring social information does not influence the value of personal information, but more experiments are needed to manipulate pay-offs and disentangle whether information sources affect response thresholds or change discrimination. This article is part of the theme issue 'Signal detection theory in recognition systems: from evolving models to experimental tests'.

Egg-Cracking Vibration as a Cue for Stink Bug Siblings to Synchronize Hatching.

Egg clutches of many animals hatch synchronously due to parental control [1, 2] or environmental stimulation [3, 4]. In contrast, in some animals, embryos actively synchronize their hatching timing with their siblings to facilitate adaptive behavior in sibling groups, such as mass migration [5, 6]. These embryos require synchronization cues that are detectable from eggs and indicative of when the siblings hatch, such as pre-hatching vocalizations in birds and crocodiles [7, 8]. Previous studies, using methods including artificial presentation of non-specific mechanical stimuli, demonstrated that vibrations or other mechanical forces caused by sibling movements are cues used by some turtles and insects [9-13]. However, there is no evidence about which movements of tiny embryos or hatchlings, among multiple possibilities, can generate mechanical cues actually detectable through eggs. Here, we show that embryos of the brown marmorated stink bug, Halyomorpha halys, synchronize hatching by responding to single pulsed vibrations generated when siblings crack open their eggshells. An egg-cracking vibration seems to be transmitted to distant eggs within a clutch while still maintaining its function as a cue, thus leading to the highly synchronized hatching pattern previously reported [14]. In this species, it is possible that embryos attempt to hatch with short lags after earlier-hatched siblings to avoid egg cannibalism by them [14]. The present study illustrates the diversity of social-information use by animal embryos for success in the sibling group.

Sex differences in the use of social information emerge under conditions of risk.

Social learning provides an effective route to gaining up-to-date information, particularly when information is costly to obtain asocially. Theoretical work predicts that the willingness to switch between using asocial and social sources of information will vary between individuals according to their risk tolerance. We tested the prediction that, where there are sex differences in risk tolerance, altering the variance of the payoffs of using asocial and social information differentially influences the probability of social information use by sex. In a computer-based task that involved building a virtual spaceship, men and women (N = 88) were given the option of using either asocial or social sources of information to improve their performance. When the asocial option was risky (i.e., the participant's score could markedly increase or decrease) and the social option was safe (i.e., their score could slightly increase or remain the same), women, but not men, were more likely to use the social option than the asocial option. In all other conditions, both women and men preferentially used the asocial option to a similar degree. We therefore found both a sex difference in risk aversion and a sex difference in the preference for social information when relying on asocial information was risky, consistent with the hypothesis that levels of risk-aversion influence the use of social information.

Can video playback provide social information for foraging blue tits?

Video playback is becoming a common method for manipulating social stimuli in experiments. Parid tits are one of the most commonly studied groups of wild birds. However, it is not yet clear if tits respond to video playback or how their behavioural responses should be measured. Behaviours may also differ depending on what they observe demonstrators encountering. Here we present blue tits (Cyanistes caeruleus) videos of demonstrators discovering palatable or aversive prey (injected with bitter-tasting Bitrex) from coloured feeding cups. First we quantify variation in demonstrators' responses to the prey items: aversive prey provoked high rates of beak wiping and head shaking. We then show that focal blue tits respond differently to the presence of a demonstrator on a video screen, depending on whether demonstrators discover palatable or aversive prey. Focal birds faced the video screen more during aversive prey presentations, and made more head turns. Regardless of prey type, focal birds also hopped more frequently during the presence of a demonstrator (compared to a control video of a different coloured feeding cup in an empty cage). Finally, we tested if demonstrators' behaviour affected focal birds' food preferences by giving individuals a choice to forage from the same cup as a demonstrator, or from the cup in the control video. We found that only half of the individuals made their choice in accordance to social information in the videos, i.e., their foraging choices were not different from random. Individuals that chose in accordance with a demonstrator, however, made their choice faster than individuals that chose an alternative cup. Together, our results suggest that video playback can provide social cues to blue tits, but individuals vary greatly in how they use this information in their foraging decisions.

The Evolution of Individual and Cultural Variation in Social Learning.

It is often assumed in experiments and models that social learning abilities - how often individuals copy others, plus who and how they copy - are species-typical. Yet there is accruing evidence for systematic individual variation in social learning within species. Here we review evidence for this individual variation, placing it within a continuum of increasing phenotypic plasticity, from genetically polymorphic personality traits, to developmental plasticity via cues such as maternal stress, to the individual learning of social learning, and finally the social learning of social learning. The latter, possibly restricted to humans, can generate stable between-group cultural variation in social learning. More research is needed to understand the extent, causes, and consequences of this individual and cultural variation.

Collective decision making and social interaction rules in mixed-species flocks of songbirds.

Associations in mixed-species foraging groups are common in animals, yet have rarely been explored in the context of collective behaviour. Despite many investigations into the social and ecological conditions under which individuals should form groups, we still know little about the specific behavioural rules that individuals adopt in these contexts, or whether these can be generalized to heterospecifics. Here, we studied collective behaviour in flocks in a community of five species of woodland passerine birds. We adopted an automated data collection protocol, involving visits by RFID-tagged birds to feeding stations equipped with antennae, over two winters, recording 91 576 feeding events by 1904 individuals. We demonstrated highly synchronized feeding behaviour within patches, with birds moving towards areas of the patch with the largest proportion of the flock. Using a model of collective decision making, we then explored the underlying decision rule birds may be using when foraging in mixed-species flocks. The model tested whether birds used a different decision rule for conspecifics and heterospecifics, and whether the rules used by individuals of different species varied. We found that species differed in their response to the distribution of conspecifics and heterospecifics across foraging patches. However, simulating decisions using the different rules, which reproduced our data well, suggested that the outcome of using different decision rules by each species resulted in qualitatively similar overall patterns of movement. It is possible that the decision rules each species uses may be adjusted to variation in mean species abundance in order for individuals to maintain the same overall flock-level response. This is likely to be important for maintaining coordinated behaviour across species, and to result in quick and adaptive flock responses to food resources that are patchily distributed in space and time.