Effects of tree phytochemistry on the interactions among endophloedic fungi associated with the southern pine beetle

We examined the interaction between host trees and fungi associated with a tree-killing bark beetle, Dendroctonus frontalis. We evaluated (1) the response of four Pinus species to fungal invasion and (2) the effects of plant secondary metabolites on primary growth of and secondary colonization of three consistent fungal associates. Two of these fungi, Entomocorticium sp. A and Ophiostoma ranaculosum, are obligate mutualists with D. frontalis, and the third associate is a blue-staining fungus, O. minus, that is commonly introduced by beetles and phoretic mites. O. minus negatively affects beetle larvae and in high abundance can impact D. frontalispopulation dynamics. Size of lesions formed and quantity of secondary metabolites produced in response to fungal inoculations varied significantly among Pinus species. However, monoterpene composition within infected tissue did not significantly vary across treatments. While all eight tested metabolites negatively affected the growth rate of O. minus, only 4-allylanisole, p-cymene, and terpinene reduced the growth of the mycangial fungi. Surprisingly, growth rates of mycangial fungi increased in the presence of several secondary metabolite volatiles. O. minus out-competed both mycangial fungi, but the presence of secondary metabolites altered the outcome slightly. O. ranaculosum out-performed E. sp. A in the presence of dominant conifer monoterpenes, such as α- and β-pinene. Volatiles from the mycangial fungi, particularly E. sp. A, had a negative effect on O. minus growth. In general, phloem phytochemistry of particular Pinus species appeared to alter the relative growth and competitiveness of mutualistic and non-mutualistic fungi associated with D. frontalis. The outcome of interactions among these fungi likely has important consequences for the population dynamics of D. frontalis.