Colonization of weakened trees by mass-attacking bark beetles: no penalty for pioneers, scattered initial distributions and final regular patterns

Bark beetles use aggregation pheromones to promote group foraging, thus increasing the chances of an individual to find a host and, when relevant, to overwhelm the defences of healthy trees. When a male beetle finds a suitable host, it releases pheromones that attract potential mates as well as other 'spying' males, which result in aggregations on the new host. To date, most studies have been concerned with the use of aggregation pheromones by bark beetles to overcome the defences of living, well-protected trees. How insects behave when facing undefended or poorly defended hosts remains largely unknown. The spatio-temporal pattern of resource colonization by the European eight-toothed spruce bark beetle, Ips typographus, was quantified when weakly defended hosts (fallen trees) were attacked. In many of the replicates, colonization began with the insects rapidly scattering over the available surface and then randomly filling the gaps until a regular distribution was established, which resulted in a constant decrease in nearest-neighbour distances to a minimum below which attacks were not initiated. The scattered distribution of the first attacks suggested that the trees were only weakly defended. A minimal theoretical distance of 2.5 cm to the earlier settlers (corresponding to a density of 3.13 attacks dm(-2)) was calculated, but the attack density always remained lower, between 0.4 and 1.2 holes dm(-2), according to our observations.