Stephen C. Pratt

Capturing qualitative features of animal behavior requires sampling occurrences of behavior over time. A continuously sampled dataset can capture many qualitative features of animal behavior; however, it can be very time‐consuming and sometimes infeasible. Instantaneous sampling can reduce the amount of labor required but will miss the fine structure present in continuously recorded datasets. We therefore explored a synthesis of these techniques which we call piecewise‐continuous sampling, where there are multiple continuous samples during randomly dispersed time intervals. To test the efficacy of this technique, we collected a continuous behavioral dataset of harvester‐ant workers and then randomly sampled from this dataset using continuous sampling, instantaneous sampling, and piecewise‐continuous sampling, each with approximately the same total amount of observation time. We then measured errors associated with measuring various attributes of animal behavior for each type of sampling technique. The different sampling techniques had dissimilar strengths and weaknesses. Using a multi‐objective optimization technique (desirability functions), we show that piecewise‐continuous sampling can be used to explore the gradient between continuous sampling and instantaneous sampling to find the strategy which simultaneously minimizes effort and error.

The collective transport of massive food items by ant teams is a striking example of biological cooperation, but it remains unclear how these decentralized teams coordinate to overcome the various challenges associated with transport. Previous research has focused on transport across horizontal surfaces and very shallow inclines, disregarding the complexity of natural foraging environments. In the ant Oecophylla smaragdina, prey are routinely carried up tree trunks to arboreal nests. Using this species, we induced collective transport over a variety of angled surfaces with varying prey masses to investigate how ants respond to inclines. We found that weight and incline pose qualitatively different challenges during transport. Prey were carried over vertical and inclined surfaces faster than across horizontal surfaces, even though inclines were associated with longer routes and a higher probability of dropping the load. This additional speed was associated with more transporters being allocated to loads on steeper inclines and not with the persistence of individual ants. Ant teams also regulated a stable prey delivery rate (rate of return per transporter) across all treatments. Our proposed constrained optimization model suggests a possible explanation for these results; theoretically, prey intake rate at the colony level is maximized when the allocation of transporters yields a similar prey delivery rate across loads.

Ants have evolved diverse recruitment methods to guide colony members to valuable resources, such as food or nest sites. One of these methods, tandem running, consists of an informed leader directly guiding a naive follower every step of the way from nest to resource. Although this behavior appears superficially similar in the different ant taxa in which it has independently evolved, this similarity could conceal underlying functional and mechanistic differences. Here we present a combined network and information-theoretic analysis, which reveals fundamental differences in the tandem recruitment between two distantly related ant genera, Temnothorax and Diacamma. Temnothorax uses tandem running to recruit additional recruiters, whereas Diacamma uses it principally to move the passive majority of the colony, a task that Temnothorax accomplishes with a different behavior, social carrying. Accordingly, the structure of the tandem run recruitment networks of Diacamma was different from those of Temnothorax , with Diacamma networks more closely resembling the social carrying networks of Temnothorax . Furthermore, an information-theoretic analysis of the spatial trajectories of leaders and followers revealed that Diacamma tandem runs lack bidirectional information flow, the signature of route-learning in Temnothorax tandem runs. These results suggest that Diacamma uses tandem runs not to share information, but to transport nestmates. By quantifying the cryptic diversity of communication behavior, this study increases the resolution of our understanding of animal societies.

The collective transport of massive food items by ant teams is a striking example of biological cooperation, but it remains unclear how these decentralized teams coordinate to overcome the various challenges associated with transport. Previous research has focused on transport across horizontal surfaces and very shallow inclines, disregarding the complexity of natural foraging environments. In the ant Oecophylla smaragdina , prey are routinely carried up tree trunks to arboreal nests. Using this species, we induce collective transport over a variety of angled surfaces with varying prey masses to investigate how ants respond to inclines. We found that weight and incline pose qualitatively different challenges during transport. Prey were carried over vertical inclines faster than across horizontal surfaces even though inclines were associated with longer routes and a higher probability of dropping the load. This additional speed is associated with more transporters being allocated to vertical loads and not from the persistence of individual ants. Ant teams also regulated a stable “prey delivery rate” (rate of return per transporter) across all treatments. Our proposed constrained optimization model explains these results: prey intake rate at the colony level is maximized when the allocation of transporters yields a similar prey delivery rate across loads.

Group-living animals that rely on stable foraging or migratory routes can develop behavioural traditions to pass route information down to inexperienced individuals. Striking a balance between exploitation of social information and exploration for better alternatives is essential to prevent the spread of maladaptive traditions. We investigated this balance during cumulative route development in the homing pigeonColumba livia. We quantified information transfer within pairs of birds in a transmission-chain experiment and determined how birds with different levels of experience contributed to the exploration–exploitation trade-off. Newly introduced naïve individuals were initially more likely to initiate exploration than experienced birds, but the pair soon settled into a pattern of alternating leadership with both birds contributing equally. Experimental pairs showed an oscillating pattern of exploration over generations that might facilitate the discovery of more efficient routes. Our results introduce a new perspective on the roles of leadership and information pooling in the context of collective learning.

In collective animal motion, coordination is often achieved by feedback between leaders and followers. For stable coordination, a leader's signals and a follower's responses are hypothesized to be attuned to each other. However, their roles are difficult to disentangle in species with highly coordinated movements, hiding potential diversity of behavioural mechanisms for collective behaviour. Here, we show that twoCoptotermestermite species achieve a similar level of coordination via distinct sets of complementary leader–follower interactions. Even thoughC. gestroifemales produce less pheromone thanC. formosanus, tandem runs of both species were stable. Heterospecific pairs withC. gestroimales were also stable, but not those withC. formosanusmales. We attributed this to the males' adaptation to the conspecific females;C. gestroimales have a unique capacity to follow females with small amounts of pheromone, whileC. formosanusmales rejectC. gestroifemales as unsuitable but are competitive over females with large amounts of pheromone. An information-theoretic analysis supported this conclusion by detecting information flow from female to male only in stable tandems. Our study highlights cryptic interspecific variation in movement coordination, a source of novelty for the evolution of social interactions.

Group-living animals that rely on stable foraging or migratory routes can develop behavioural traditions to pass route information down to inexperienced individuals. Striking a balance between exploitation of social information and exploration for better alternatives is essential to prevent the spread of maladaptive traditions. We investigated this balance during cumulative route development in the homing pigeon Columba livia . We quantified information transfer within pairs of birds in a transmission-chain experiment and determined how birds with different levels of experience contributed to the exploration– exploitation trade-off. Newly introduced naïve individuals were initially more likely to initiate exploration than experienced birds, but the pair soon settled into a pattern of alternating leadership with both birds contributing equally. Experimental pairs showed an oscillating pattern of exploration over generations that might facilitate the discovery of more efficient routes. Our results introduce a new perspective on the roles of leadership and information pooling in the context of collective learning.

Leadership of animal group movements depends on social feedback, hence leader’s signals and follower’s responses should be attuned to each other. However, leader and follower roles are difficult to disentangle in species with high levels of coordination. To overcome this challenge, we investigated a simple case of movement coordination: termite pairs in which a female leads a male as they search for a nest site. To tease apart leader and follower roles, we created conspecific and heterospecific pairs of Coptotermes gestroi and C. formosanus , which share a pairing pheromone so that males follow females of either species. Conspecific pairs were stable for both species, even though C. gestroi females produce less pheromone than C. formosanus . Heterospecific pairs with C. gestroi males were also stable, but not those with C. formosanus males. We attributed this difference to the C. gestroi male’s unique capacity to follow females that release small amounts of pheromone; C. formosanus males cannot follow or reject C. gestroi females as unsuitable. This conclusion was supported by an information-theoretic analysis that detected information flow from C. formosanus females to C. gestroi males as in conspecific pairs, but not from C. gestroi females to C. formosanus males. Despite their following ability, C. gestroi males lost to C. formosanus males in competitions to follow C. formosanus females. Thus, partner selection has shaped the species-specific association of mating pairs. Our results demonstrate that a similar level of coordination can emerge from distinct sets of complementary sender-receiver interactions.