Schlyter, F., Byers, J.A. & Lfqvist, J. 1987a. Attraction to pheromone sources of different quantity, quality, and spacing: Density-regulation mechanisms in bark beetle Ips typographus. Journal of Chemical Ecology 13:1503-1523.

Abstract-- The density of bark-beetle colonization of a tree could be regulated by a quantitative effect of the pheromone signal from beetles in the tree (cessation of release of attractive pheromone) or by a qualitative effect (production of pheromone components inhibiting attraction). The quantitative hypothesis was tested on Ips typographus by varying the release rate of the two known attractive compounds, 2-methyl-3-buten-2-ol (MB) and (4S)-cis-verbenol (cV). The highest number of beetles were captured at traps with the highest release rates. The catch was nearly proportional to the release of MB and cV at a distance between traps of 12 m or more. At 6-, 3-, and 1.5-m distances between traps deployed in a triangular arrangement there was still a good discrimination between release rates, but relatively more beetles, especially males, were caught on the blank. The lower release rates caught an equal sex ratio while the highest release rate caught only about 30% males. The qualitative hypothesis was tested by releasing the suspected inhibitors ipsdienol (Id) and ipsenol (Ie), from traps in the same amounts as cV. Only small effects were noted for I. typographus. However, the competitor I. duplicatus was attracted to Id and inhibited by Ie, while the predator Thanasimus formicarius was attracted to both compounds. On the other hand, when the ratio of Id or Ie to cV was 10:1 or 0.1:1 rather than 1:1, they affected the numbers of I. typographus attracted. A small amount of Id combined with the attractants increased trap catch, while large amounts of Id or Ie decreased attraction, especially when combined. Attack density regulation is modeled as an effect of both quantitative and qualitative mechanisms acting in sequence.
Chemical Ecology