Byers, J.A. 1991. Simulation of mate-finding behaviour of pine shoot beetles, Tomicus piniperda. Animal Behaviour 41:649-660.

Abstract-- An algorithm for animal searching behaviour was constructed that allows independent variation of movement parameters such as speed, duration, step size, and maximum right or left turn angle. The area, radius of the object searched for, and number of objects and searchers can also be varied. A capture algorithm is presented that can determine whether a searching animal intercepts a circle (object) irrespective of the step size. These two algorithms were incorporated in a computer simulation model of mate finding for walking male bark beetles, Tomicus piniperda, in search of females that remained stationary when boring into the bark of Scots pine, Pinus sylvestris. The model showed that, for realistically chosen parameters, males were able to find `females' beyond the size of an actual female. This indicates that there may be little, if any, advantage in the evolution of a long-range pheromone, for which no chemical and behavioural evidence has yet been found.
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