Temporal clumping of bark beetle arrival at pheromone traps: Modeling anemotaxis in chaotic plumes

The sequence of arrival of the bark beetlesIps typographus andPityogenes chalcographus (Coleoptera: Scolytidae) at traps baited with their synthetic pheromones was monitored with a portable fraction collector. Histograms of the natural arrival patterns of both species were nonrandom and clumped at shorter time scales (1-, 2-, 4-, 5-, or 6-min cells) but appeared random at larger time scales (10, 20 or 30 min). Monte Carlo generation of similar histograms showed them to be random at all of these time scales. A stochastic computer model could graphically simulate insect orientation to odor sources based on present theories of odor-modulated anemotaxis and casting. Although this model was used throughout, since it assumes only that insects cast perpendicular to the current wind direction, a second model could slightly improve orientation success. However, the second model requires that the insect remember its ground path (upwind) prior to losing the plume (after an abrupt wind direction change). The effects of casting and flight parameters on orientation success and randomness of arrival sequence within various plumes were determined by simulation. Similarly, the effects of random walks in plume direction, plume width, and wind speed were explored. The results showed that dynamic random variations in plume direction and especially wind speed could cause an otherwise random arrival sequence (e.g., under constant wind) to become clumped and nonrandom. Therefore, the clumped arrival patterns of bark beetles and other insects, includingSpodoptera litura, at pheromone sources could result from random-walk fluctuations in wind speed and wind direction.