The American Naturalist
We consider the hypothesis that mountain pine beetles function as cybernetic regulators of primary productivity in ecosystems of lodgepole pine forest through their selective killing of dominant trees and the subsequent redistribution of resources. Following a recent major beetle outbreak in Yellowstone and Grand Teton national parks, surviving trees did grow significantly faster (P < .1); wood production was redistributed among canopy, subcanopy, and understory trees; and annual wood production per hectare usually returned to pre-attack levels or exceeded them within 10-15 yr. However, reconstructions of annual wood production over the last 70-80 yr indicate that the beetle outbreak did not reduce the variation in productivity; rather, the beetles introduced more variation than would have existed in their absence. Hence, our results do not support the hypothesis that the beetles function as cybernetic regulators (in the strict sense). Nevertheless, the beetle-pine system that we studied shows great resilience, and the effects of beetles on primary productivity do not appear to be as severe as conventional wisdom maintains. Annual wood production per hectare returned quickly to previous levels in the stands we studied, and associated ecological changes can be considered generally benign or even beneficial.
Romme, W.H., D.H. Knight, and J.B. Yavitt. 1986. Mountain pine beetle outbreaks in the Rocky Mountains: Regulators of primary productivity? Am. Nat. 127(4):484-494