In Chapter 1, I summarize relevant past work on food and nest recruitment. Then I describe T. rugatulus and its recruitment behavior, tandem running, and I explain its suitability for these questions. In Chapter 2, I investigate whether food and nest recruiters behave differently. I report two novel behaviors used by recruiters during their interaction with nestmates. Food recruiters perform these behaviors more often than nest recruiters, suggesting that they convey information about target type. In Chapter 3, I investigate whether colonies respond to a tradeoff between foraging and emigration by allocating their workforce adaptively. I describe how colonies responded when I posed a tradeoff by manipulating colony need for food and shelter and presenting both resources simultaneously. Recruitment and visitation to each target partially matched the predictions of the tradeoff hypothesis. In Chapter 4, I address the tuned error hypothesis, which states that the error rate in recruitment is adaptively tuned to the patch area of the target. Food tandem leaders lost followers at a higher rate than nest tandem leaders. This supports the tuned error hypothesis, because food targets generally have larger patch areas than nest targets with small entrances.
This work shows that animal groups face tradeoffs as individual animals do. It also suggests that colonies spatially allocate their workforce according to resource type. Investigating recruitment for multiple resource types gives a better understanding of exploitation of each resource type, how colonies make collective decisions under conflicting goals, as well as how colonies manage the exploitation of multiple types of resources differently. This has implications for managing the health of economically important social insects such as honeybees or invasive fire ants.
This work investigates mechanisms that can extend the spatial reach of fertility signaling and reproductive regulation in three polydomous ant species. In Novomessor cockerelli, the presence of larvae but not eggs is shown to inhibit worker reproduction. Then, in Camponotus floridanus, 3-methylheptacosane found on the queen cuticle and queen-laid eggs is verified as a releaser pheromone sufficient to disrupt normally occurring aggressive behavior toward foreign workers. Finally, the volatile and cuticular hydrocarbon pheromones present on the cuticle of Oecophylla smaragdina queens are shown to release strong attraction response by workers; when coupled with previous work, this result suggests that these chemicals may underly both the formation of a worker retinue around the queen as well as egg-located mechanisms of reproductive regulation in distant satellite nests. Whereas most previous studies have focused on the short-range role of hydrocarbons on the cuticle of the queen, these studies demonstrate that eusocial insects may employ longer range regulatory mechanisms. Both queen volatiles and distributed brood can extend the range of queen fertility signaling, and the use of larvae for fertility signaling suggest that feeding itself may be a non-chemical mechanism for reproductive regulation. Although trail laying in mass-recruiting ants is often used as an example of complex communication, reproductive regulation in ants may be a similarly complex example of insect communication, especially in the case of large, polydomous ant colonies.
Collective behaviors in social insect societies often emerge from simple local rules. However, little is known about how these behaviors are dynamically regulated in response to environmental changes. Here, we use a compartmental modeling approach to identify factors that allow harvester ant colonies to regulate collective foraging activity in response to their environment. We propose a set of differential equations describing the dynamics of: (1) available foragers inside the nest, (2) active foragers outside the nest, and (3) successful returning foragers, to understand how colony-specific parameters, such as baseline number of foragers, interactions among foragers, food discovery rates, successful forager return rates, and foraging duration might influence collective foraging dynamics, while maintaining functional robustness to perturbations. Our analysis indicates that the model can undergo a forward (transcritical) bifurcation or a backward bifurcation depending on colony-specific parameters. In the former case, foraging activity persists when the average number of recruits per successful returning forager is larger than one. In the latter case, the backward bifurcation creates a region of bistability in which the size and fate of foraging activity depends on the distribution of the foraging workforce among the model׳s compartments. We validate the model with experimental data from harvester ants (Pogonomyrmex barbatus) and perform sensitivity analysis. Our model provides insights on how simple, local interactions can achieve an emergent and robust regulatory system of collective foraging activity in ant colonies.