Random sequential addition simulations of animal aggregations provide null models of group structure

Abstract

Apparent structure in animal aggregations such as fish and Antarctic krill schools may result from the tight packing of these elongated animals. This geometrical structure may be difficult to differentiate from behavior-induced structure resulting from individuals preferentially taking up certain positions relative to conspecifics to gain an adaptive advantage such as reduced locomotive cost. Here we use random sequential addition (RSA) simulations to quantify the effect of animal shape, aggregation organization, and aggregation density on 2D school structure. This technique allows for the generation of a null model for nearest neighbor distance and nearest neighbor position angle for a specific body shape and aggregation density, thus isolating the effect of geometry from that of behavior. We further identify a shape-specific aggregation density threshold above which the animal shape affects the spatial distribution of nearest neighbors. Nearest neighbor distance data of fish schools with densities above and below the threshold are found to agree well with nearest neighbor statistics found from RSA-generated schools.