Predator-prey pursuit-evasion games in structurally complex environments.

Pursuit and evasion behaviors in many predator-prey encounters occur in a geometrically structured environment. The physical structures in the environment impose strong constraints on the perception and behavioral responses of both antagonists. Nevertheless, no experimental or theoretical study has tackled the issue of quantifying the role of the habitat's architecture on the joint trajectories during a predator-prey encounter. In this study, we report the influence of microtopography of forest leaf litter on the pursuit-evasion trajectories of wolf spiders Pardosa sp. attacking the wood cricket Nemobius sylvestris. Fourteen intact leaf litter samples of 1 m × 0.5 m were extracted from an oak-beech forest floor in summer and winter, with later samples having the most recently fallen leaves. Elevation was mapped at a spatial resolution of 0.5 mm using a laser scanner. Litter structuring patterns were identified by height transects and experimental semi-variograms. Detailed analysis of all visible leaf-fragments of one sample enabled us to relate the observed statistical patterns to the underlying geometry of individual elements. Video recording of pursuit-evasion sequences in arenas with flat paper or leaf litter enabled us to estimate attack and fleeing distances as a function of substrate. The compaction index, the length of contiguous flat surfaces, and the experimental variograms showed that the leaf litter was smoother in summer than in winter. Thus, weathering as well as biotic activities compacted and flattened the litter over time. We found good agreement between the size of the structuring unit of leaf litter and the distance over which attack and escape behaviors both were initiated (both ∼3 cm). There was a four-fold topographical effect on pursuit-escape sequences; compared with a flat surface, leaf litter (1) greatly reduced the likelihood of launching a pursuit, (2) reduced pursuit and escape distances by half, (3) put prey and predator on par in terms of pursuit and escape distances, and (4) reduced the likelihood of secondary pursuits, after initial escape of the prey, to nearly zero. Thus, geometry of the habitat strongly modulates the rules of pursuit-evasion in predator-prey interactions in the wild.