Whitebark pine (Pinus albicaulis) is a critical forest species of Northern Rocky Mountain upper subalpine ecosystems, yet little is known about the physiological response of whitebark pine to disturbance (e.g. fire, bark beetles, and pathogens) across a range of diverse environmental gradients. Resin-based defenses have long been recognized as the primary mechanism by which conifers respond to attack by bark beetles and pathogens and several studies have linked resin duct properties to survivorship during periods of increased beetle activity. However, to our knowledge, no studies have compared axial resin ducts in the secondary xylem of whitebark pine across pairs of living and dead whitebark pine trees to better understand survivorship following multiple disturbances including mountain pine beetle and white pine blister rust. 144 whitebark pine trees were sampled as part of this research (72 live and 72 dead) based on size (< 3 cm difference in diameter) and distance (< 20 m apart) to control for potential microsite differences. We analyzed growth (ring widths) and defense metrics (resin duct properties) across all years of overlapping lifespan for each live / dead pair.
Results/Conclusions
We found a clear distinction in growth and defense characteristics between live and dead whitebark pine. Across our study sites on the Flathead Indian Reservation in northwestern Montana, live whitebark pine produced larger resin ducts with a greater annual investment in resin-based defenses than whitebark pine that died. Resin duct size, duct area, and relative duct area were all greater in live whitebark pine (by 56%, 48%, and 57%, respectively) and these were the most important variables influencing whitebark pine survivorship. In contrast, whitebark pine that had died grew faster over time (22% larger ring widths) than their live counterparts and also produced more resin duct structures (20% more ducts on average). Whitebark pine at our study sites exhibit differing strategies in the allocation of resources toward growth and defense, with the majority of survivors of recent disturbance investing more in defensive structures than growth. Our results support the idea that maintaining genetic variability and the associated suite of differing physiological traits promotes diverse response strategies to a complex array of biophysical and biological stressors that might leave a species vulnerable to extinction across its range.