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Richard G. Everett

Publications and source records attributed to Richard G. Everett.

4 recordsLinked to original sources

Increased whitebark pine (Pinus albicaulis) growth and defense under a warmer and regionally drier climate

Introduction: Tree defense characteristics play a crucial role in modulating conifer bark beetle interactions, and there is a growing body of literature investigating factors mediating tree growth and resin-based defenses in conifers. A subset of studies have looked at relationships between tree growth, resin duct morphology and climate; however, these studies are almost exclusively from lower-elevation, moisture-limited systems. The relationship between resin ducts and climate in higher-elevation, energy-limited ecosystems is currently poorly understood. Methods: In this study, we: (1) evaluated the relationship between biological trends in tree growth, resin duct anatomy, and climatic variability and (2) determined if tree growth and resin duct morphology of whitebark pine, a high-elevation conifer of management concern, is constrained by climate and/or regional drought conditions. Results: We found that high-elevation whitebark pine trees growing in an energy-limited system experienced increased growth and defense under warmer and regionally drier conditions, with climate variables explaining a substantive proportion of variation (∼20–31%) in tree diameter growth and resin duct anatomy. Discussion: Our results suggest that whitebark pine growth and defense was historically limited by short growing seasons in high-elevation environments; however, this relationship may change in the future with prolonged warming conditions.

Montana

Growth and defense characteristics of whitebark pine (Pinus albicaulis) and lodgepole pine (Pinus contorta var latifolia) in a high-elevation, disturbance-prone mixed-conifer forest in northwestern Montana, USA

Recent, widespread tree mortality in the western U.S. resulting from changes in climate, pathogens, insect activity, and forest management practices has led to concerns for many ecologically and culturally important species. Within conifers, resin-based defenses have long been recognized as a primary defense mechanism against a variety of insects and pathogens. Oleoresin produced by trees contain complex mixtures of terpenoids that have numerous insecticidal and fungicidal properties. Research has also identified links between resin duct characteristics and increased probability of survival during bark beetle outbreaks. Whitebark pine ( Pinus albicaulis ) is a culturally significant high elevation species that provides numerous ecological services within subalpine and alpine ecosystems. Whitebark pine has co-evolved with a suite of biotic and abiotic disturbances. Individual trees allocate resources towards growth and resin-based defenses, making it a good candidate species to evaluate growth and defense relationships and tradeoffs. In this study we compared constitutive resin chemistry, tree growth and resin duct anatomy between similarly aged whitebark and lodgepole pine ( P. contorta var latifolia ) growing in proximity within a disturbance-prone, mixed-conifer forest in northwestern Montana. These two host species have varying degrees of historical exposure to mountain pine beetle. Our research yields four important findings. First, we did not find evidence of a tradeoff between tree growth and tree defenses (resin duct morphology and resin chemistry). This suggests that trees growing under favorable field conditions can experience high growth rates and still allocate ample resources towards defense. Second, we found that resin ducts and constitutive mono- and sesqui- terpenes were not correlated in lodgepole pine while duct production and area were positively related to constitutive monoterpenes, and duct size and area were positively related to constitutive sesquiterpenes, in whitebark pine. The lack of distinct, consistent relationships between these defensive features suggests that both whitebark and lodgepole pine trees present beetles with numerous, complex combinations of resin-based defenses. Third, based on constitutive terpene profiles, bark beetles are more likely to enter lodgepole pine but more likely to successfully elicit mass attacks in whitebark pine, which agrees with beetle attack and success patterns observed in the field. Fourth, overstory competition, particularly by Engelmann spruce ( Picea engelmannii ), can influence tree defenses, specifically by reducing constitutive terpene concentrations in lodgepole and whitebark pine. Competitive tree interactions could lead to altered bark beetle-conifer interactions as host and nonhost species migrate in response to changing climate. Our results suggest that strategies designed to support whitebark pine populations can benefit from better understanding interactions among growth, competition and physical and chemical defenses in response to multiple disturbance.

Montana

Millennial-scale climate and human drivers of environmental change and fire activity in a dry, mixed-conifer forest of northwestern Montana

Warm summer temperatures and longer fire seasons are promoting larger, and in some cases, more fires that are severe in low- and mid-elevation, dry mixed-conifer forests of the Northern Rocky Mountains (NRM). Long-term historical fire conditions and human influence on past fire activity are not well understood for these topographically and biophysically heterogeneous forests. We developed reconstructions of millennial-scale fire activity, vegetation change, and human presence at Black Lake, a small closed-basin lake on the Flathead Indian Reservation in the Mission Valley, Northwestern Montana, United States. Fossil pollen, charcoal, and biomarkers associated with human presence were used to evaluate the interaction between climate variability, fire activity, vegetation change and human activity for the past 7000 years. Comparisons among multiple proxies suggest climate variability acted as the primary control on fire activity and vegetation change from the early Holocene until the late Holocene when records suggest fire activity and climate variability decoupled. Specific biomarkers (5β-stanols including coprostanol and epi-coprostanol) associated with human presence indicate humans were present within the Black Lake watershed for thousands of years, although the inferred intensity of human presence is highly variable. A strong relationship between climate variability and fire activity during the early and mid-Holocene weakens during the last few thousand years, suggesting possible increased influence of humans in mediating fire activity in recent millennia, and/or a shift in the interaction between the distribution and abundance of woody fuel and fire severity. Human-set fires during the cooler and wetter late Holocene may have been aimed at maintaining important cultural resources associated with the heterogeneous mosaic of mixed conifer forests within the Black Lake watershed. The paleoenvironmental reconstruction at Black Lake corroborates archeological records that show humans were present within the Black Lake watershed for over 7000 years. Further research is needed to evaluate the evidence for this continuous presence and the possible role that people played in shaping fire regimes and vegetation within low- to mid-elevation mixed-conifer ecosystems of the NRM.

Montana

Whitebark pine (Pinus albicaulis) growth and defense in response to mountain pine beetle outbreaks

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. 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.

Northern Rocky Mountain