A statistical method for identifying different rules of interaction between individuals in moving animal groups.

The emergent patterns of collective motion are thought to arise from application of individual-level rules that govern how individuals adjust their velocity as a function of the relative position and behaviours of their neighbours. Empirical studies have sought to determine such rules of interaction applied by 'average' individuals by aggregating data from multiple individuals across multiple trajectory sets. In reality, some individuals within a group may interact differently from others, and such individual differences can have an effect on overall group movement. However, comparisons of rules of interaction used by individuals in different contexts have been largely qualitative. Here we introduce a set of randomization methods designed to determine statistical differences in the rules of interaction between individuals. We apply these methods to a case study of leaders and followers in pairs of freely exploring eastern mosquitofish (Gambusia holbrooki). We find that each of the randomization methods is reliable in terms of: repeatability of p-values, consistency in identification of significant differences and similarity between distributions of randomization-based test statistics. We observe convergence of the distributions of randomization-based test statistics across repeat calculations, and resolution of any ambiguities regarding significant differences as the number of randomization iterations increases.

Cohesion, order and information flow in the collective motion of mixed-species shoals.

Despite the frequency with which mixed-species groups are observed in nature, studies of collective behaviour typically focus on single-species groups. Here, we quantify and compare the patterns of interactions between three fish species, threespine sticklebacks (Gasterosteus aculeatus), ninespine sticklebacks (Pungitius pungitius) and roach (Rutilus rutilus) in both single- and mixed-species shoals in the laboratory. Pilot data confirmed that the three species form both single- and mixed-species shoals in the wild. In our laboratory study, we found that single-species groups were more polarized than mixed-species groups, while single-species groups of threespine sticklebacks and roach were more cohesive than mixed shoals of these species. Furthermore, while there was no difference between the inter-individual distances between threespine and ninespine sticklebacks within mixed-species groups, there was some evidence of segregation by species in mixed groups of threespine sticklebacks and roach. There were differences between treatments in mean pairwise transfer entropy, and in particular we identify species-differences in information use within the mixed-species groups, and, similarly, differences in responses to conspecifics and heterospecifics in mixed-species groups. We speculate that differences in the patterns of interactions between species in mixed-species groups may determine patterns of fission and fusion in such groups.

Crimson Spotted Rainbowfish (Melanotaenia duboulayi) Change Their Spatial Position according to Nutritional Requirement.

Decision making in moving animal groups has been shown to be disproportionately influenced by individuals at the front of groups. Therefore, an explanation of state-dependent positioning of individuals within animal groups may provide a mechanism for group movement decisions. Nutritional state is dynamic and can differ between members of the same group. It is also known to drive animal movement decisions. Therefore, we assayed 6 groups of 8 rainbowfish foraging in a flow tank. Half of the fish had been starved for 24h and half had been fed 1h prior to experimental start. Groups were assayed again one week later but individuals were allocated to the opposite nutritional treatment. During the assay the positions of individually identified fish were recorded as were the number of food items they each ate and the position within the group they acquired them from. Food-deprived fish were more often found towards the front of the shoal; the mean weighted positional score of food-deprived fish was significantly larger than that of well-fed fish. Individuals were not consistent in their position within a shoal between treatments. There was a significant positive correlation between mean weighted positional score and number of food items acquired which displays an obvious benefit to front positions. These results suggest that positional preferences are based on nutritional state and provide a mechanism for state-dependent influence on group decision-making as well as increasing our understanding of what factors are important for group functioning.

Do small swarms have an advantage when house hunting? The effect of swarm size on nest-site selection by Apis mellifera.

Reproductive swarms of honeybees are faced with the problem of finding a good site to establish a new colony. We examined the potential effects of swarm size on the quality of nest-site choice through a combination of modelling and field experiments. We used an individual-based model to examine the effects of swarm size on decision accuracy under the assumption that the number of bees actively involved in the decision-making process (scouts) is an increasing function of swarm size. We found that the ability of a swarm to choose the best of two nest sites decreases as swarm size increases when there is some time-lag between discovering the sites, consistent with Janson & Beekman (Janson & Beekman 2007 Proceedings of European Conference on Complex Systems, pp. 204-211.). However, when simulated swarms were faced with a realistic problem of choosing between many nest sites discoverable at all times, larger swarms were more accurate in their decisions than smaller swarms owing to their ability to discover nest sites more rapidly. Our experimental fieldwork showed that large swarms invest a larger number of scouts into the decision-making process than smaller swarms. Preliminary analysis of waggle dances from experimental swarms also suggested that large swarms could indeed discover and advertise nest sites at a faster rate than small swarms.

The role of individuality in collective group movement.

How different levels of biological organization interact to shape each other's function is a central question in biology. One particularly important topic in this context is how individuals' variation in behaviour shapes group-level characteristics. We investigated how fish that express different locomotory behaviour in an asocial context move collectively when in groups. First, we established that individual fish have characteristic, repeatable locomotion behaviours (i.e. median speeds, variance in speeds and median turning speeds) when tested on their own. When tested in groups of two, four or eight fish, we found individuals partly maintained their asocial median speed and median turning speed preferences, while their variance in speed preference was lost. The strength of this individuality decreased as group size increased, with individuals conforming to the speed of the group, while also decreasing the variability in their own speed. Further, individuals adopted movement characteristics that were dependent on what group size they were in. This study therefore shows the influence of social context on individual behaviour. If the results found here can be generalized across species and contexts, then although individuality is not entirely lost in groups, social conformity and group-size-dependent effects drive how individuals will adjust their behaviour in groups.