Tandem running by foraging Pachycondyla striata workers in field conditions vary in response to food type, food distance, and environmental conditions

Ants show collective and individual behavioural flexibility in their response to immediate context, choosing for example between different foraging strategies. In Pachycondyla striata, workers can forage solitarily or recruit and guide nestmates to larger food sources through tandem running. Although considered more ancestral and less efficient than pheromone trail-laying, this strategy is common especially in species with small colony size. What is not known is how the decision to recruit or follow varies according to the immediate context. That is, how fine adjustments in information transfer affect immediate foraging decisions at the colony level. Here, we studied individually marked workers and evaluated their foraging decisions when food items varied in nature (protein vs carbohydrate), size, and distance from the nest at different temperatures and humidity levels. Our results show that tandem run leaders and potential followers adjust their behaviour according to a combination of external factors. While 84.2% of trips were solitary, most ants (81%) performed at least one tandem run. However, tandem runs were more frequent for nearby resources and at higher relative humidity. Interestingly, when food items were located far away, tandem runs were more successful when heading to protein sources (75%) compared to carbohydrate sources (42%). Our results suggest that the social information transfer between leaders and followers conveys more information than previously thought, and also relies on their experience and motivation.

Large body size variation is associated with low communication success in tandem running ants

Diversity in animal groups is often assumed to increase group performance. In insect colonies, genetic, behavioural and morphological variation among workers can improve colony functioning and resilience. However, it has been hypothesized that during communication processes, differences between workers, e.g. in body size, could also have negative effects. Tandem running is a common recruitment strategy in ants and allows a leader to guide a nestmate follower to resources. A substantial proportion of tandem runs fail because leader and follower lose contact. Using the ant Temnothorax nylanderi as a model system, we tested the hypothesis that tandem running success is impaired if leader and follower differ in size. Indeed, we found that the success rate of tandem pairs drops considerably as size variation increases: tandem runs were unsuccessful when the leader–follower size difference exceeded 10%, whereas ~ 80% of tandem runs were successful when ants differed less than 5% in body length. Possible explanations are that size differences are linked to differences in walking speed or sensory perception. Ants did not choose partners of similar size, but extranidal workers were larger than intranidal workers, which could reduce recruitment mistakes because it reduced the chance that very large and very small ants perform tandem runs together. Our results suggest that phenotypic differences between interacting workers can have negative effects on the efficiency of communication processes. Whether phenotypic variation has positive or negative effects is likely to depend on the task and the phenotypic trait that shows variation. Significance statement Diversity is often assumed to increase colony performance in social insects. However, phenotypic differences among workers could also have negative effects, e.g. during communication. Tandem running is a common recruitment strategy in ants, but tandem runs often fail when ants lose contact. We used the ant Temnothorax nylanderi to test the hypothesis that body size differences between tandem leader and follower impair tandem communication. We show that the success rate of tandem pairs drops considerably as size variation increases, possibly because ants of varying size also differ in walking speed. Our study supports the hypothesis that phenotypic variation among workers might not always be beneficial and can negatively impact the efficiency of communication processes.

The adaptive value of tandem communication in ants: insights from an agent-based model

Social animals often share information about the location of resources, such as a food source or a new nest-site. One well-studied communication strategy in ants is tandem running, whereby a leader guides a recruit to a resource. Tandem running is considered an example of animal teaching because a leader adjusts her behaviour and invests time to help another ant to learn the location of a resource more efficiently. Tandem running also has costs, such as waiting inside the nest for a leader and a reduced walking speed. Whether and when these costs outweigh the benefits of tandem running is not well understood. We developed an agent-based simulation model to investigate the conditions that favour communication by tandem running during foraging. We predicted that the spatio-temporal distribution of food sources, colony size and the ratio of scouts and recruits affect colony foraging success. Our results suggest that communication is favoured when food sources are hard to find, of variable quality and long lasting. These results mirror the findings of simulations of honeybee communication. Scouts locate food sources faster than tandem followers in some environments, suggesting that tandem running may fulfil the criteria of teaching only in some situations. Furthermore, tandem running was only beneficial above a critical colony size threshold. Taken together, our model suggests that there is a considerable parameter range that favours colonies that do not use communication, which could explain why many social insects with small colony sizes forage solitarily.

Revealing the structure of information flows discriminates similar animal social behaviors

Behavioral correlations stretching over time are an essential but often neglected aspect of interactions among animals. These correlations pose a challenge to current behavioral-analysis methods that lack effective means to analyze complex series of interactions. Here we show that non-invasive information-theoretic tools can be used to reveal communication protocols that guide complex social interactions by measuring simultaneous flows of different types of information between subjects. We demonstrate this approach by showing that the tandem-running behavior of the ant Temnothorax rugatulus and that of the termites Coptotermes formosanus and Reticulitermes speratus are governed by different communication protocols. Our discovery reconciles the diverse ultimate causes of tandem running across these two taxa with their apparently similar signaling mechanisms. We show that bidirectional flow of information is present only in ants and is consistent with the use of acknowledgement signals to regulate the flow of directional information.

Large body size variation is linked to low communication success in tandem running ants

Diversity in animal groups is often assumed to increase group performance. In insect colonies, genetic, behavioral and morphological variation among workers can improve colony functioning and resilience. However, it has been hypothesized that during communication processes, differences between workers, e.g. in body size, could also have negative effects. Tandem running is a common recruitment strategy in ants and allows a leader to guide a nestmate follower to resources. A substantial proportion of tandem runs fail because leader and follower loose contact. Using the ant Temnothorax nylanderi as a model system, we tested the hypothesis that tandem running success is impaired if leader and follower differ in size. Indeed, we found that the success rate of tandem pairs drops considerably as size variation increases: only ~7% of tandem runs were successful when the leader-follower size difference exceeded 10%, whereas 80% of tandem runs were successful when ants differed less than 5% in body length. One possible explanation is that ant size is linked to the preferred walking speed. Ants did not choose partners of similar size, but extranidal workers were larger than intranidal workers, which could reduce recruitment mistakes because it reduced the chance that very large and very small ants perform tandem runs together. Our results suggest that phenotypic differences between interacting workers can have negative effects on the efficiency of communication processes. Whether phenotypic variation has positive or negative effects is likely to depend on the task and the phenotypic trait that shows variation.