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(Dendroctonus Ponderosae Hopkins) Under Climate Change Utah State University DigitalCommons@USU All Graduate Theses and Dissertations Graduate Studies 8-2021 Susceptibility of High-Elevation Forests to Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) Under Climate Change David N. Soderberg Utah State University Follow this and additional works at: https://digitalcommons.usu.edu/etd Part of the Ecology and Evolutionary Biology Commons Recommended Citation Soderberg, David N., "Susceptibility of High-Elevation Forests to Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) Under Climate Change" (2021). All Graduate Theses and Dissertations. 8139. https://digitalcommons.usu.edu/etd/8139 This Dissertation is brought to you for free and open access by the Graduate Studies at DigitalCommons@USU. It has been accepted for inclusion in All Graduate Theses and Dissertations by an authorized administrator of DigitalCommons@USU. For more information, please contact [email protected]. SUSCEPTIBILITY OF HIGH-ELEVATION FORESTS TO MOUNTAIN PINE BEETLE (DENDROCTONUS PONDEROSAE HOPKINS) UNDER CLIMATE CHANGE by David N. Soderberg A dissertation submitted in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY in Ecology Approved: ______________________ ______________________ Karen E. Mock, Ph.D. Barbara J. Bentz, Ph.D. Major Professor Committee Member ______________________ ______________________ R. Justin DeRose, Ph.D. Zachariah Gompert, Ph.D. Committee Member Committee Member ______________________ ______________________ Justin B. Runyon, Ph.D. D. Richard Cutler, Ph.D. Committee Member Interim Vice Provost of Graduate Studies UTAH STATE UNIVERSITY Logan, UT 2021 ii Copyright © David N. Soderberg 2021 All Rights Reserved iii ABSTRACT Susceptibility of High-elevation Forests to Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) Under Climate Change by David N. Soderberg, Doctor of Philosophy Utah State University, 2021 Major Professor: Dr. Karen Mock Department: Wildland Resources High-elevation Pinus species act as keystone species in subalpine communities through providing habitat and food sources that foster biodiversity. The mountain pine beetle (Dendroctonus ponderosae Hopkins Coleoptera: Curculionidae, Scolytinae; MPB) affects the majority of Pinus species including many high-elevation, five-needle Pinus species, and has recently expanded its distribution further northward and increased persistence at higher elevations. Future distribution of MPB will be dictated by climate and the ability to adapt to novel environments, in addition to the availability of susceptible Pinus hosts. Considerable variability exists not only in regional adaptation among MPB populations, but also in the resistance conferred by defense strategies employed among and within Pinus species. Defenses take the form of chemical secondary metabolites (e.g., monoterpenoids) or anatomical structures (e.g., resin ducts), both of which can be expressed constitutively, or upregulated as needed to maximize the economy of available resources. Among Pinus, the concentration and composition of iv constitutive and inducible secondary metabolites has been shown to confer defense against MPB. In addition, lignification within bark beetle feeding tissues (e.g., bark, phloem) has been shown to reduce brood fitness within related genera, but its defensive efficacy has yet to be assessed within Pinus. I employed a variety of experimental approaches to assess the role of climate on MPB persistence and southern range expansion, in addition to resource allocation strategies of growth and both chemical (i.e., secondary metabolites) and anatomical (i.e., lignin) defenses among high-elevation Pinus. The results from this work suggest that due to genetic variability and extensive plasticity in multiple fitness traits, MPB populations will not only persist, but increase reproductive success in a warming climate. In addition, the MPB southern range boundary is likely limited by biotic interactions, rather than direct temperature effects. Among Pinus that differ in susceptibility to MPB, the concentration and composition of chemical secondary metabolites, as well as concentrations of lignin within the phloem were inversely correlated, with less MPB-susceptible Pinus species (e.g., P. longaeva) displaying higher concentrations of secondary metabolites, but lower concentrations of phloem lignin, relative to more MPB-susceptible species (e.g. P. flexilis). These findings provide supporting evidence for evolved differences among Pinus species in resource allocation to growth and defenses, where SM concentration and composition, but not phloem or bark lignification, are adaptive traits for resisting MPB attack and brood success. My dissertation research advances our understanding of the interactions between MPB and its high-elevation, five-needle Pinus hosts, contributing to the adaptive management of high-elevation forests. (225 pages) v PUBLIC ABSTRACT Susceptibility of High-elevation Forests to Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) Under Climate Change David N. Soderberg Across western North America, pine forests are important for timber, wildlife habitat, and at high elevations are important for water retention and yield from rain and snowmelt. The mountain pine beetle (MPB) is one of the most significant disturbance agents shaping pine forests, and like all insects, temperature is a major driver of its population success and the dynamics of the landscapes that they inhabit. Changing temperature regimes can therefore directly influence MPB population persistence at a particular location, in addition to potential shifts in the range boundaries that they inhabit. MPB is currently expanding its range northward in British Columbia and Alberta, Canada in parallel with warming climates, however, the potential impact of climate change on southern populations of mountain pine beetle is unknown. As the climate warms, the future distribution of MPB will be dictated by the ability to adapt to new and changing environments, in addition the availability and susceptibility of the pine trees that they feed upon. Pine species are known to vary in susceptibility to MPB, which is largely attributed to differences in the production of chemical (e.g., terpenes and their derivatives) and physical (e.g., resin ducts) defenses. Among pines, chemical defenses have been shown to confer defense against MPB, however, the nature of these defenses following biotic incitation has not been evaluated in many pine species. Moreover, lignification within bark beetle feeding tissues (e.g., bark, phloem) has been shown to vi confer defense within related conifers, but its defensive efficacy has yet to be assessed within pines. To assess MPB response to a changing climate and the relative susceptibility of their pine hosts, I employed a variety of experimental approaches to assess the role of climate on MPB persistence and southern range expansion, in addition to the growth and defense strategies employed within and among high-elevation pine hosts that vary in resistance to MPB. The results from this work suggests that in a warming climate, MPB populations will not only persist, but increase in population. In addition, the MPB southern range boundary is likely limited by biotic interactions, rather than direct temperature effects. Among pines that differ in susceptibility to MPB, the concentration and composition of chemical defenses, as well as concentrations of lignin within the phloem were inversely correlated, with less MPB-susceptible pine species (e.g., Great Basin bristlecone pine) displaying higher concentrations of chemical defenses, but lower concentrations of phloem lignin, relative to more MPB-susceptible species (e.g., limber pine). These findings provide supporting evidence for evolved differences among pine species in investment between growth and defenses, where the concentration and composition of various chemical defenses, but not phloem or bark lignification, are adaptive traits for resisting MPB attack and brood development. My dissertation research advances our understanding of the interactions between MPB and its high-elevation, five- needle Pinus hosts, contributing to the adaptive management of high-elevation forests. vii ACKNOWLEDGMENTS I would first and foremost like to thank my two major advisors, Dr. Barbara Bentz and Dr. Karen Mock. You both exemplify the telos of a scientist and mentor and I am overwhelmingly thankful for your persistence and everlasting effort and patience in guiding me through this process. It has been a genuine privilege to be under your tutelage and I cannot overstate the positive impact that you have had on my maturation as a scientist and person. I would like to thank my committee members, Drs. Justin DeRose, Zach Gompert, and Justin Runyon, as well as my coauthors, Dr. Enrico Bonello, Dr. Richard Hofstetter, Bethany Kyre, and Dr. Sharon Hood, for sharing your insight and work spaces with me. I would like to thank my fellow graduate students and university collaborators for their friendship. In particular, but in no specific order, Kendall Becker, Martin Holdredge, Erika Eidson, Tucker Furniss, Sara Germain, Alex Howe, Stephen Pfeiler, Jim Walton, Torrey Rodgers, and most of all, my partner Erika Blomdahl. I have and will continue to enjoy our discussions on the state of the snowpack, philosophy, logic, bike and home maintenance, and reason – usually in that order. You all (and many more!) provided the scaffolding for the growth and joy that I feel so lucky to have experienced
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