Volatiles of High-Elevation Five-Needle Pines: Chemical Signatures through Ratios and Insight into Insect and Pathogen Resistance

High-elevation five-needle pine trees are a group of Pinus species in the subgenus Strobus that occur at the edges of plant growth near the alpine tree line. These species are ecologically very important and are also threatened by climate-driven insect outbreaks and an exotic pathogen. Volatile organic compounds (VOCs) play central roles in the environmental adaptation of plants and in their defense against insects and pathogens. For example, the VOCs emitted by some high-elevation five-needle pine species attract female, tree-killing mountain pine beetles (MPB, Dendroctonus ponderosae) in the pioneering phase whereas VOCs from other species strongly repel this foremost herbivore, but the mechanism is unknown. We collected and compared headspace VOCs from foliage of eight species of high-elevation five-needle pines in Europe and North America. Overall, VOCs differed quantitatively among species with few qualitative differences. Despite species emitting essentially the same compounds, Random Forest analysis correctly classified 117 of the 126 trees sampled by using VOCs and identified the most important compounds for species classification and for separating species resistant from those susceptible to MPB or white pine blister rust (Cronartium ribicola). These VOC 'fingerprints' resulted largely from species emitting distinctive ratios of compounds, rather than through presence of species-specific compounds. Importantly, these Pinus species vary greatly in resistance to the main herbivore (MPB) and pathogen (white pine blister rust) causing tree mortality. Thus, these findings provide insights and should guide research into understanding resistance and in developing tools to manage these important trees. For instance, studies into the functions of five-needle pine VOCs in defense against abiotic or biotic stressors should focus on blend ratios rather than on individual compounds.