Ja. Byers, AN ENCOUNTER RATE MODEL OF BARK BEETLE POPULATIONS SEARCHING AT RANDOM FOR SUSCEPTIBLE HOST TREES, Ecological modelling, 91(1-3), 1996, pp. 57-66
Iterative equations were developed that predict the encounter rate bet
ween a population of moving animals and a population of stationary obj
ects, where the animals cease to search upon finding an object. The en
counter rate through time depends on the number of searching animals (
bark beetles), number of stationary objects (host trees), average spee
d of the animals, average radius of the object, and area of the search
arena. The iterative equations were used in a computer program to var
y these parameters with regard to flight dispersal of the bark beetle
Ips typographus (Coleoptera: Scolytidae) during their search for susce
ptible host trees of Norway spruce (Picea abies). Realistic parameters
of flight speed, numbers of beetles and susceptible host trees, tree
diameters, density of healthy trees, search area, and time searching w
ere held constant while certain of these parameters were varied in com
puter model runs. In most cases, significant proportions of the modele
d bark beetle population (of which individuals fly forward with a rand
om component) found the relatively few susceptible host trees suitable
for colonization. Only at very low effective flight speeds (due to lo
nger stays on trees) or with widely distributed hosts of smaller diame
ter, did relatively few beetles find suitable hosts. Once the 'pioneer
' beetles find susceptible hosts, release of aggregation pheromone wou
ld greatly increase the effective radius of the host and allow rapid c
oncentration of the population on these trees. The model suggests that
primary attraction to host tree volatiles is not mandatory for host f
inding and selection in many species of bark beetle. Evolution of an o
lfactory response to host volatiles is more probable in species with l
ow population densities or widely dispersed host plants, or both. Also
, there would be little selection pressure on a bark beetle species to
evolve aggregation pheromones if they can respond over some meters to
plant volatiles that reveal the location of a susceptible host.