Does aggregation benefit bark beetles by diluting predation? Links between a group-colonisation strategy and the absence of emergent multiple predator effects

1. Aggregation in bark beetles (Coleoptera: Scolytidae) aids in mate attraction and resource procurement when colonising well-defended plants; however, some species colonise primarily poorly defended plants, and intraspecific competition increases mortality. The hypothesis that decreased risk of predation was a potential benefit to aggregation in such circumstances was tested, using the pine engraver, Ips pini (Say) and its two major predators Thanasimus dubius (F.) (Coleoptera: Cleridae) and Platysoma cylindrica (Paykull) (Coleoptera: Histeridae). Both single- and multiple-predator effects, across a range of prey densities, were tested. 2. Both male and female colonisation events increased with herbivore density, in an asymptotic fashion. 3. Predators decreased the number of colonisers in a density-dependent manner, consistent with a type II functional response. 4. The proportional impact of predators decreased with increased herbivore colonisation densities. These findings indicate that predator dilution may be a viable benefit to aggregation. 5. Total emergence of the herbivore also increased with density, although the net replacement rate during one generation was independent of initial arrival density. This was likely due to larval predation, which negates potential relationships between per capita reproductive success and establishment density. 6. Each predator species decreased I. pini's net replacement rate by approximately 42%, and their combined effect was approximately 70%. 7. Overall, these predators modified their prey's establishment and adult mortality relationships in additive manners. This is somewhat surprising, given the potential for emergent effects due to interactions between multiple predators foraging within a common habitat. The persistence of additivity, rather than risk reduction or enhancement to the prey, may increase the predator-swamping benefit to aggregation for this herbivore. 8. The effects of these predators are substitutable, and likely exert equivalent selective pressures to mask signals at the whole-plant level.