Coupled information networks drive honeybee (Apis mellifera) collective foraging.

Collective behaviour by eusocial insect colonies is typically achieved through multiple communication networks that produce complex behaviour at the group level but often appear to provide redundant or even competing information. A classic example occurs in honeybee (Apis mellifera) colonies, where both the dance communication system and robust scent-based mechanisms contribute to the allocation of a colony's workforce by regulating the flow of experienced foragers among known food sources. Here we analysed social connectivity patterns during the reactivation of experienced foragers to familiar feeding sites to show that these social information pathways are not simply multiple means to achieve the same end but intersect to play complementary roles in guiding forager behaviour. Using artificial feeding stations, we mimicked a natural scenario in which two forager groups were simultaneously collecting from distinct patches containing different flowering species. We then observed the reactivation of these groups at their familiar feeding sites after interrupting their foraging. Social network analysis revealed that temporarily unemployed individuals interacted more often and for longer with foragers that advertised a familiar versus unfamiliar foraging site. Due to such resource-based assortative mixing, network-based diffusion analysis estimated that reactivation events primarily resulted from interactions among bees that had been trained to the same feeding station and less so from different-feeder interactions. Both scent- and dance-based interactions strongly contributed to reactivation decisions. However, each bout of dance-following had an especially strong effect on a follower's likelihood of reactivation, particularly when dances indicated locations familiar to followers. Our findings illustrate how honeybee foragers can alter their social connectivity in ways that are likely to enhance collective outcomes by enabling foragers to rapidly access up-to-date information about familiar foraging sites. In addition, our results highlight how reliance on multiple communication mechanisms enables social insect workers to utilise flexible information-use strategies that are robust to variation in the availability of social information.

Behavioural variation among workers promotes feed-forward loops in a simulated insect colony.

Coordinated responses in eusocial insect colonies arise from worker interaction networks that enable collective processing of ecologically relevant information. Previous studies have detected a structural motif in these networks known as the feed-forward loop, which functions to process information in other biological regulatory networks (e.g. transcriptional networks). However, the processes that generate feed-forward loops among workers and the consequences for information flow within the colony remain largely unexplored. We constructed an agent-based model to investigate how individual variation in activity and movement shaped the production of feed-forward loops in a simulated insect colony. We hypothesized that individual variation along these axes would generate feed-forward loops by driving variation in interaction frequency among workers. We found that among-individual variation in activity drove over-representation of feed-forward loops in the interaction networks by determining the directionality of interactions. However, despite previous work linking feed-forward loops with efficient information transfer, activity variation did not promote faster or more efficient information flow, thus providing no support for the hypothesis that feed-forward loops reflect selection for enhanced collective functioning. Conversely, individual variation in movement trajectory, despite playing no role in generating feed-forward loops, promoted fast and efficient information flow by linking together otherwise unconnected regions of the nest.

Detecting and quantifying social transmission using network-based diffusion analysis.

Although social learning capabilities are taxonomically widespread, demonstrating that freely interacting animals (whether wild or captive) rely on social learning has proved remarkably challenging. Network-based diffusion analysis (NBDA) offers a means for detecting social learning using observational data on freely interacting groups. Its core assumption is that if a target behaviour is socially transmitted, then its spread should follow the connections in a social network that reflects social learning opportunities. Here, we provide a comprehensive guide for using NBDA. We first introduce its underlying mathematical framework and present the types of questions that NBDA can address. We then guide researchers through the process of selecting an appropriate social network for their research question; determining which NBDA variant should be used; and incorporating other variables that may impact asocial and social learning. Finally, we discuss how to interpret an NBDA model's output and provide practical recommendations for model selection. Throughout, we highlight extensions to the basic NBDA framework, including incorporation of dynamic networks to capture changes in social relationships during a diffusion and using a multi-network NBDA to estimate information flow across multiple types of social relationship. Alongside this information, we provide worked examples and tutorials demonstrating how to perform analyses using the newly developed nbda package written in the R programming language.

Personality composition determines social learning pathways within shoaling fish.

In shaping how individuals explore their environment and interact with others, personality may mediate both individual and social learning. Yet increasing evidence indicates that personality expression is contingent on social context, suggesting that group personality composition may be key in determining how individuals learn about their environment. Here, we used recovery latency following simulated predator attacks to identify Trinidadian guppies (Poecilia reticulata) that acted in a consistently bold or shy manner. We then employed network-based diffusion analysis to track the spread of a novel foraging behaviour through groups containing different proportions of bold and shy fish. Informed associates promoted learning to a greater extent in bold individuals, but only within groups composed predominately of bold fish. As the proportion of shy fish within groups increased, bold individuals instead emerged as especially effective demonstrators that facilitated learning in others. Individuals were also more likely to learn overall within shy-dominated groups than in bold-dominated ones. We demonstrate that whether and how individuals learn is conditional on group personality composition, indicating that selection may favour traits enabling individuals to better match their behavioural phenotype to their social environment.

Network-based diffusion analysis reveals context-specific dominance of dance communication in foraging honeybees.

The honeybee (Apis mellifera) dance communication system is a marvel of collective behaviour, but the added value it brings to colony foraging efficiency is poorly understood. In temperate environments, preventing communication of foraging locations rarely decreases colony food intake, potentially because simultaneous transmission of olfactory information also plays a major role in foraging. Here, we employ social network analyses that quantify information flow across multiple temporally varying networks (each representing a different interaction type) to evaluate the relative contributions of dance communication and hive-based olfactory information transfer to honeybee recruitment events. We show that virtually all successful recruits to novel locations rely upon dance information rather than olfactory cues that could otherwise guide them to the same resource. Conversely, during reactivation to known sites, dances are relatively less important, as foragers are primarily guided by olfactory information. By disentangling the contributions of multiple information networks, the contexts in which dance communication truly matters amid a complex system full of redundancy can now be identified.

Honeybee Communication: There's More on the Dancefloor.

Honeybees transmit food-related information to nestmates via waggle dances and food-sharing. A new study reveals that learning about food scents can also be mediated by social contact alone, suggesting unexpected complexity in honeybee foraging networks.

Fear of predation shapes social network structure and the acquisition of foraging information in guppy shoals.

Spatio-temporal variation in predation risk is predicted to select for plastic anti-predator responses, which may in turn impact the fine-scale social structure of prey groups and processes mediated by that structure. To test these predictions, we manipulated the ambient predation risk experienced by Trinidadian guppy (Poecilia reticulata) groups before quantifying their social networks and recording individual latencies to approach and solve a novel foraging task. High-risk conditions drove the formation of social networks that were more strongly assorted by body size than those exposed to low ambient risk and promoted longer durations of contact between preferred partners. Additionally, high background predation risk reduced the probability individuals would approach and solve a novel foraging task. Network-based diffusion analysis revealed that while social transmission of the task solution from knowledgeable to naive individuals occurred at a higher rate within low-risk groups, individuals in high-risk groups were particularly likely to investigate the task while shoaling with preferred social partners. Taken together, our results suggest that the structure and functional importance of prey social networks may partly depend on local predation pressure. Furthermore, by influencing individuals' access to information, fear of predation may impact decision-making in a potentially wide array of behavioural contexts.

Environmental conditions associated with repetitive behavior in a group of African elephants.

Repetitive movement patterns are commonly observed in zoo elephants. The extent to which these behaviors constitute a welfare concern varies, as their expression ranges from stereotypies to potentially beneficial anticipatory behaviors. Nevertheless, their occurrence in zoo animals is often viewed negatively. To better identify conditions that prompt their performance, observations were conducted on six African elephants (Loxodonta africana) at the North Carolina Zoo. Individuals spent most of their time engaged in feeding, locomotion, resting, and repetitive behavior. Both generalized estimating equation and zero-inflated negative binomial models were used to identify factors associated with increased rates of repetitive behavior. Time of day in conjunction with location on- or off-exhibit best explained patterns of repetitive behavior. Repetitive behaviors occurred at a lower rate in the morning when on-exhibit, as compared to afternoons on-exhibit or at any time of day off-exhibit. Increased repetitive behavior rates observed on-exhibit in the afternoon prior to the evening transfer and feeding were possibly anticipatory responses towards those events. In contrast, consistently elevated frequencies of repetitive behavior off-exhibit at all times of day could be related to differences in exhibit complexity between off-exhibit and on-exhibit areas, as well as a lack of additional foraging opportunities. Our study contributes valuable information on captive elephant behavior and represents a good example of how behavioral research can be employed to improve management of zoo animals.