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Two recent studies using tree aggregations to analyze forest age-structure stability and past forest structure are flawed. A better understanding of aggregation dynamics is needed before aggregation analysis is used in forest management.
The 7th North American Forest Ecology Workshop, consisting of 149 presentations in 16 oral sessions and a poster session, reflected a broad range of topical areas currently under investigation in forest ecology and management. There was an overarching emphasis on the role of disturbance, both natural and anthropogenic, in the dynamics of forest ecosystems, and the recognition that legacies from past disturbances strongly influence future trajectories. Climate was invoked as a major driver of ecosystem change. An emphasis was placed on application of research findings for predicting system responses to changing forest management initiatives. Several “needs” emerged from the discussions regarding approaches to the study of forest ecosystems, including (1) consideration of variable spatial and temporal scales, (2) long-term monitoring, (3) development of universal databases more encompassing of time and space to facilitate meta-analyses, (4) combining field studies and modeling approaches, (5) standardizing methods of measurement and assessment, (6) guarding against oversimplification or overgeneralization from limited site-specific results, (7) greater emphasis on plant-animal interactions, and (8) better alignment of needs and communication of results between researchers and managers.
Studies of relationships between seasons and Neotropical carnivore distributions tend to focus on water and prey availability without considering other habitat components such as escape, foraging, and resting cover. Our goal was to evaluate habitat characteristics that may be important for predicting the seasonal (dry or rainy) relative abundance of four commonly captured Neotropical carnivores (i.e., jaguar [ Panthera onca ], puma [ Puma concolor ], ocelot [ Leopardus pardalis ], and grey fox [ Urocyon cinereoargenteus ]) in Chiquibul Forest Reserve in Belize, Central America. We used trail camera data and random-effect Poisson models to investigate how prey ratios (number of prey detections/total detections), cover (e.g., logs and stumps used for hiding cover from predators), vegetation structure, and environmental site characteristics (e.g., site harvest-history, slope, aspect) were related to carnivore relative abundance. Both prey ratios and vegetation structure appeared in supported models more frequently than other environmental site characteristics and were negatively correlated with carnivore relative abundance. Supported models differed for each season for all species except jaguars for which mammalian prey ratios and prey cover at sites was always negatively correlated with jaguar relative abundance. Carnivores appeared to avoid sites where vegetation created ideal escape and hiding cover for prey even though prey may be less abundant. Our data suggest that vegetation structure and composition can create conditions conducive to carnivore foraging and that these characteristics can differ by season in the tropics.
The complex mix of economic and ecological objectives facing today's forest managers necessitates the development of growth models with a capacity for simulating a wide range of forest conditions while producing outputs useful for economic analyses. We calibrated the gap model ZELIG to simulate stand-level forest development in the Oregon Coast Range as part of a landscape-scale assessment of different forest management strategies. Our goal was to incorporate the predictive ability of an empirical model with the flexibility of a forest succession model. We emphasized the development of commercial-aged stands of Douglas-fir, the dominant tree species in the study area and primary source of timber. In addition, we judged that the ecological approach of ZELIG would be robust to the variety of other forest conditions and practices encountered in the Coast Range, including mixed-species stands, small-scale gap formation, innovative silvicultural methods, and reserve areas where forests grow unmanaged for long periods of time. We parameterized the model to distinguish forest development among two ecoregions, three forest types and two site productivity classes using three data sources: chronosequences of forest inventory data, long-term research data, and simulations from an empirical growth-and-yield model. The calibrated model was tested with independent, long-term measurements from 11 Douglas-fir plots (6 unthinned, 5 thinned), 3 spruce-hemlock plots, and 1 red alder plot. ZELIG closely approximated developmental trajectories of basal area and large trees in the Douglas-fir plots. Differences between simulated and observed conifer basal area for these plots ranged from -2.6 to 2.4 m2/ha; differences in the number of trees/ha ???50 cm dbh ranged from -8.8 to 7.3 tph. Achieving these results required the use of a diameter-growth multiplier, suggesting some underlying constraints on tree growth such as the temperature response function. ZELIG also tended to overestimate regeneration of shade-tolerant trees and underestimate total tree density (i.e., higher rates of tree mortality). However, comparisons with the chronosequences of forest inventory data indicated that the simulated data are within the range of variability observed in the Coast Range. Further exploration and improvement of ZELIG is warranted in three key areas: (1) modeling rapid rates of conifer tree growth without the need for a diameter-growth multiplier; (2) understanding and remedying rates of tree mortality that were higher than those observed in the independent data; and (3) improving the tree regeneration module to account for competition with understory vegetation. ?? 2008 Elsevier B.V.
The future of dry forests around the world is uncertain given predictions that rising temperatures and enhanced aridity will increase drought-induced tree mortality. Using forest management and ecological restoration to reduce density and competition for water offers one of the few pathways that forests managers can potentially minimize drought-induced tree mortality. Competition for water during drought leads to elevated tree mortality in dense stands, although the influence of density on heat-induced stress, and the durations of hot or dry conditions that most impact mortality, remain unclear. Understanding how competition interacts with hot-drought stress is essential to recognize how, where, and how much reducing density can help sustain dry forests in a rapidly changing world. Here, we integrated repeat measurements of 28,881 ponderosa pine trees across the western US (2000-2017) with soil moisture estimates from a water balance model to examine how annual mortality responds to competition, temperature and soil moisture conditions. Tree mortality responded most strongly to basal area, and was elevated in places with high mean temperatures, unusually hot 7-year high temperature anomalies, and unusually dry 8-year low soil moisture anomalies. Mortality was also lower in places that experienced unusually wet 3-year soil moisture anomalies between measurements. Importantly, we found that basal area interacts with temperature and soil moisture, exacerbating mortality during times of stress imposed by high temperature or low moisture. Synthesis and Applications: Our results imply that a 50% reduction in forest basal area could reduce drought-driven tree mortality by 20-80%. The largest impacts of density reduction are seen in areas with high current basal area and places that experience high temperatures and/or severe multiyear droughts. These interactions between competition and drought are critical to understand past and future patterns of tree mortality in the context of climate change, and provide information for resource managers seeking to enhance dry forest drought resistance.
The implications of cumulative land-use decisions and shifting climate on forests, require us to integrate our understanding of ecosystems, markets, policy, and resource management into a social-ecological system. Humans play a central role in macrosystem dynamics, which complicates ecological theories that do not explicitly include human interactions. These dynamics also impact ecological services and related markets, which challenges economic theory. Here, we use two forest macroscale management initiatives to develop a theoretical understanding of how management interacts with ecological functions and services at these scales and how the multiple large-scale management goals work either in consort or conflict with other forest functions and services. We suggest that calling upon theories developed for organismal ecology, ecosystem ecology, and ecological economics adds to our understanding of social-ecological macrosystems. To initiate progress, we propose future research questions to add rigor to macrosystem-scale studies: (1) What are the ecosystem functions that operate at macroscales, their necessary structural components, and how do we observe them? (2) How do systems at one scale respond if altered at another scale? (3) How do we both effectively measure these components and interactions, and communicate that information in a meaningful manner for policy and management across different scales?
Sugar maple ( Acer saccharum ) is among the most ecologically and economically important tree species in North America, and its growth and regeneration is often the focus of silvicultural practices in northern hardwood forests. A key stressor for sugar maple (SM) is acid rain, which depletes base cations from poorly-buffered forest soils and has been associated with much lower SM vigor, growth, and recruitment. However, the potential interactions between forest management and soil acidification – and their implications for the sustainability of SM and its economic and cultural benefits – have not been investigated. In this study, we simulated the development of 50 extant SM stands in the western Adirondack region of NY (USA) for 100 years under different soil chemical conditions and silvicultural prescriptions. We found that interactions between management prescription and soil base saturation will strongly shape the ability to maintain SM in managed forests. Below 12% base saturation, SM did not regenerate sufficiently after harvest and was replaced mainly by red maple ( Acer rubrum ) and American beech ( Fagus grandifolia ). Loss of SM on acid-impaired sites was predicted regardless of whether the shelterwood or diameter-limit prescriptions were used. On soils with sufficient base saturation, models predicted that SM will regenerate after harvest and be sustained for future rotations. We then estimated how these different post-harvest outcomes, mediated by acid impairment of forest soils, would affect the potential monetary value of ecosystem services provided by SM forests. Model simulations indicated that a management strategy focused on syrup production – although not feasible across the vast areas where acid impairment has occurred – may generate the greatest economic return. Although pollution from acid rain is declining, its long-term legacy in forest soils will shape future options for sustainable forestry and ecosystem stewardship in the northern hardwood forests of North America.
Timber harvesting has been proposed as a management tool to enhance breeding habitat for the Cerulean Warbler ( Setophaga cerulea ), a declining Neotropical–Nearctic migratory songbird that nests in the canopy of mature eastern deciduous forests. To evaluate how this single-species management focus might fit within an ecologically based management approach for multiple forest birds, we performed a manipulative experiment using four treatments (three intensities of timber harvests and an unharvested control) at each of seven study areas within the core Cerulean Warbler breeding range. We collected pre-harvest (one year) and post-harvest (four years) data on the territory density of Cerulean Warblers and six additional focal species, avian community relative abundance, and several key habitat variables. We evaluated the avian and habitat responses across the 3–32 m 2 ha −1 residual basal area (RBA) range of the treatments. Cerulean Warbler territory density peaked with medium RBA (∼16 m 2 ha −1 ). In contrast, territory densities of the other focal species were negatively related to RBA (e.g., Hooded Warbler [ Setophaga citrina ]), were positively related to RBA (e.g., Ovenbird [ Seiurus aurocapilla ]), or were not sensitive to this measure (Scarlet Tanager [ Piranga olivacea ]). Some species (e.g., Hooded Warbler) increased with time post-treatment and were likely tied to a developing understory, whereas declines (e.g., Ovenbird) were immediate. Relative abundance responses of additional species were consistent with the territory density responses of the focal species. Across the RBA gradient, greatest separation in the avian community was between early successional forest species (e.g., Yellow-breasted Chat [ Icteria virens ]) and closed-canopy mature forest species (e.g., Ovenbird), with the Cerulean Warbler and other species located intermediate to these two extremes. Overall, our results suggest that harvests within 10–20 m 2 ha −1 RBA yield the largest increases in Cerulean Warblers, benefit additional disturbance-dependent species, and may retain closed-canopy species but at reduced levels. Harvests outside the optimum RBA range for Cerulean Warblers can support bird assemblages specifically associated with early or late (closed-canopy) successional stages.
Pinyon-juniper (PJ) woodlands are a diverse ecosystem type providing a wealth of ecosystem services across western North America. Managing PJ woodlands in the 21st century entails balancing multiple conservation objectives, and resource managers and policy-makers working to sustain PJ woodlands need spatially explicit information about current PJ woodland conditions and how they may be impacted in coming decades in the context of wildfire risk and changing climate. Here, we address knowledge gaps and provide information that improves the long-term value of conservation and restoration actions in PJ woodlands. To this end, we merged projections of future environmental suitability for nine PJ species with wildfire risk and locations of mature and old-growth (MOG) woodlands to assess spatial variation in PJ woodlands with differing threats and management opportunities. We identified potential climate refugia with enduring high community suitability and low burn probability (3 % of study area ) that may persist with relatively little management. We found promising locations of PJ-MOG forest type with high future suitability (12 % of areas) that could be prioritized for fire risk reduction to maintain high-value woodlands. Despite a 38 % mean community suitability decline under future climate conditions, some locations (7 % of areas) may act as climate refugia where future climate conditions can support current PJ woodland composition and structure. We conclude by demonstrating how this information can be integrated into a conceptual framework to help prioritize conservation and climate adaptation in PJ woodlands.
Introduction: Climate change is impacting forest-based agricultural systems with implications for producer decision-making and livelihoods. This article presents a case study on the observations, perceptions, knowledge, and adaptation strategies of maple syrup producers in the United States to climate change. Methods: We carried out two semi-structured surveys with maple producers on: (1) climate change and its impacts on the maple system ( n = 106 participants); and (2) responses to climate adaptation scenarios ( n = 98 participants). Additionally, we carried out two focus groups and key informant interviews ( n = 70+) to understand barriers and opportunities for climate adaptation. One of these focus groups and follow up key informant interviews was with tribally affiliated community members with the intention to acknowledge Indigenous Peoples’ voices, history, and relationships to the land. Results: Findings highlight that most of the surveyed producers (89%) have experienced the negative impacts of climate on maple syrup production. While 40% of participants feel concerned regarding the future of the maple system, 39% feel hopeful, with significant differences based on the age of the surveyed producers. The majority of producers have adapted their harvesting practices to climate effects. Producers shared knowledge of multiple adaptation strategies in response to climate scenarios comprised of: (1) stand management practices such as diversification of sap species tapped; (2) harvesting practices such as changing the type and number of taps; (3) sap processing practices focused on the integration of technology such as the use of an evaporator and reverse osmosis; and (4) marketing practices such as innovation of products and marketing different maple syrup characteristics. Responses shared by tribally affiliated producers highlight knowledge of multiple adaptation strategies that focus on long-term ecological management of forests rather than technological solutions. Discussion: Overall, findings emphasize the importance of cooperation and diversification at every level and dimension of the maple system for its long-term resilience.
Unraveling the effects of climate and land-use on historical fire regimes provides important insights into broader human-fire-climate dynamics, which are necessary for ecologically-based forest management. We developed a spatial human land-use model for Navajo Nation forests across which we sampled a network of tree-ring fire history sites to reflect contrasting historical land-use intensity: high human use, primarily in the Chuska Mountains, and low human use, primarily on the central Defiance Plateau. We tested for and compared human- and climate-driven changes in the fire regimes by applying change point detection, regression, and superposed epoch analyses. The historical fire regimes and fire-climate relationships reflect those of similar forests regionally, and are similar between the two Navajo landscapes until the early 1800s. We then determined that a previously identified, localized, early (1830s) decline in fire activity was geographically widespread across higher human use sites. In contrast, fires continued to burn uninterrupted through this period at the lower use sites. Though the 1830s included significantly wet and cold periods that could have contributed to fire regime decline, human factors pose a more spatio-temporally consistent explanation. A rise in Navajo pastoralism in the 1820s-1830s was concentrated seasonally in the heavy use sites. By the 1880s, livestock numbers more than doubled, grazing became far more spatially widespread, and frequent fire regimes of Navajo forests collapsed. The last widespread fire recorded on either landscape was in 1886. In the Chuska Mountains, livestock and fire co-existed for over 50 years between the initial 1832 fire decline and the end of frequent fires after 1886, an exceptional pattern in the western US. Though unique in its timing, character, and spatial dynamics, the collapse of historical fire regimes in Navajo forests contributed to now over a century without frequent surface fire, leaving Navajo forests at risk for large, uncharacteristic high-severity fires.
Laboratory and field tests were performed to examine the feasibility of using time domain reflectometry (TDR) to monitor changes in the moisture storage of the woody parts of trees. To serve as wave guides for the TDR signal, pairs of stainless steel rods (13 cm long, 0.32 cm in diameter, and 2.5 cm separation) were driven into parallel pilot holes drilled into the woody parts of trees, and a cable testing oscilloscope was used to determine the apparent dielectric constant. A laboratory calibration test was performed on two sapwood samples, so that the relation between the volumetric water content and the apparent dielectric constant of the sapwood could be determined over a range of water contents. The resulting calibration curve for these sapwood samples was significantly different than the general calibration curve used for soils, showing a smaller change in the apparent dielectric constant for a given change in the volumetric water content than is typical for soils. The calibration curve was used to estimate the average volumetric water content to a depth of 13 cm in living trees. One field experiment was conducted on an English walnut tree ( Juglans regia ) with a diameter of 40 cm, growing in a flood-irrigated orchard on a Hanford sandy loam near Modesto, California (U.S.A.). Rods were driven into the tree at about 50 cm above the soil surface and monitored hourly for the month of August, 1988. The moisture content determined by TDR showed a gradual decrease from 0.44 to 0.42 cm 3 cm −3 over a two week period prior to flood irrigation, followed by a rapid rise to 0.47 cm 3 cm −3 over a four day period after irrigation, then again a gradual decline approaching the next irrigation. A second field experiment was made on ten evergreen and deciduous trees with diameters ranging from 30 to 120 cm, growing in the foothills of the Coast Range of central California. Rods were driven into each tree at 50 to 100 cm above the soil surface and monitored on a biweekly to monthly basis for over a year. Most trees showed an early spring maximum in moisture content determined by TDR associated with leaf growth, and a late summer minimum in moisture content associated with the end of the dry season. Moisture contents ranged from 0.20 to 0.70 cm 3 cm −3 , with an annual percentage change in moisture of 15% to 70% depending on species and environmental conditions. A final field test was performed in northern New Mexico (U.S.A.) to examine the effect of trunk freezing on TDR measurements. This test confirmed that freezing conditions were recorded as a total loss of liquid water by the TDR method. These results suggest that further TDR calibration for wood, plus some understanding of the relation between tree moisture and physiological stress could be useful to several disciplines, ranging from irrigation scheduling to watershed management to forest ecology.
The diverse forest types of the southwestern US are inseparable from fire. Across climate zones in California, Nevada, Arizona, and New Mexico, fire suppression has left many forest types out of sync with their historic fire regimes. As a result, high fuel loads place them at risk of severe fire, particularly as fire activity increases due to climate change. A legacy of fire exclusion coupled with a warming climate has led to increasingly large and severe wildfires in many southwest forest types. Climate change projections include an extended fire season length due to earlier snowmelt and a general drying trend due to rising temperatures. This suggests the future will be warmer and drier regardless of changes in precipitation. Hotter, drier conditions are likely to increase forest flammability, at least initially. Changes in climate alone have the potential to alter the distribution of vegetation types within the region, and climate-driven shifts in vegetation distribution are likely to be accelerated when coupled with stand-replacing fire. Regardless of the rate of change, the interaction of climate and fire and their effects on Southwest ecosystems will alter the provisioning of ecosystem services, including carbon storage and biodiversity. Interactions between climate, fire, and vegetation growth provide a source of great uncertainty in projecting future fire activity in the region, as post-fire forest recovery is strongly influenced by climate and subsequent fire frequency. Severe fire can be mitigated with fuels management including prescribed fire, thinning, and wildfire management, but new strategies are needed to ensure the effectiveness of treatments across landscapes. We review the current understanding of the relationship between fire and climate in the Southwest, both historical and projected. We then discuss the potential implications of climate change for fire management and examine the potential effects of climate change and fire on ecosystem services. We conclude with an assessment of the role of fire management in an increasingly flammable Southwest.
Review of: Ne'eman, G. & Trabaud, L. Ecology, Biogeography and Management of Pinus halepensis and P. brutia Forest Ecosystems in the Mediterranean Basin. xii + 412 pp. Backhuys Publishers, Leiden. ISBN 90?5782-055-2 (hardcover). Price: USD 120.00.
Rockland pine forests of south Florida dominated by Pinus elliottii var. densa characteristically have poor soil development in relation to neighboring hardwood hammocks. This has led to the hypothesis that Everglades hammock trees are more reliant on soil moisture derived from local precipitation whereas pineland plants must depend more on groundwater linked to broader regional hydrologic patterns. Because soil moisture sources are likely to vary more than groundwater sources, we hypothesized that hammock plants would exhibit correspondingly higher levels of dry season water stress. This was examined by measuring predawn water potentials, and by analyzing water uptake in representative hammock and pineland woody species using stable isotopes of plant water and that of potential sources during wet and dry seasons. Two species typical of each of the two communities were selected; a fifth species which was found in both communities, Lysiloma latisiliqua Benth., was also analyzed. Water content of soils in both communities decreased from wet to dry season. Consistent with our hypothesis, the change in predawn water potentials between the wet and dry season was less in pineland species than that of hammock species. Water potential changes in L. latisiliqua in both communities resembled that of hammock species more than pineland plants. Isotopic data showed that pineland species rely proportionately more on groundwater than hammock species. Nevertheless, unlike hammock species in the Florida Keys, mainland hammock species utilized a substantial amount of groundwater during the dry season.
The structure and developmental dynamics of old-growth forests often serve as important baselines for restoration prescriptions aimed at promoting more complex structural conditions in managed forest landscapes. Nonetheless, long-term information on natural patterns of development is rare for many commercially important and ecologically widespread forest types. Moreover, the effectiveness of approaches recommended for restoring old-growth structural conditions to managed forests, such as the application of extended rotation forestry, has been little studied. This study uses several long-term datasets from old growth, extended rotation, and unmanaged second growth Pinus resinosa (red pine) forests in northern Minnesota, USA, to quantify the range of variation in structural conditions for this forest type and to evaluate the effectiveness of extended rotation forestry at promoting the development of late-successional structural conditions. Long-term tree population data from permanent plots for one of the old-growth stands and the extended rotation stands (87 and 61 years, respectively) also allowed for an examination of the long-term structural dynamics of these systems. Old-growth forests were more structurally complex than unmanaged second-growth and extended rotation red pine stands, due in large part to the significantly higher volumes of coarse woody debris (70.7 vs. 11.5 and 4.7 m 3 /ha, respectively) and higher snag basal area (6.9 vs. 2.9 and 0.5 m 2 /ha, respectively). In addition, old-growth forests, although red pine-dominated, contained a greater abundance of other species, including Pinus strobus , Abies balsamea , and Picea glauca relative to the other stand types examined. These differences between stand types largely reflect historic gap-scale disturbances within the old-growth systems and their corresponding structural and compositional legacies. Nonetheless, extended rotation thinning treatments, by accelerating advancement to larger tree diameter classes, generated diameter distributions more closely approximating those found in old growth within a shorter time frame than depicted in long-term examinations of old-growth structural development. These results suggest that extended rotation treatments may accelerate the development of old-growth structural characteristics, provided that coarse woody debris and snags are deliberately retained and created on site. These and other developmental characteristics of old-growth systems can inform forest management when objectives include the restoration of structural conditions found in late-successional forests.
Fire exclusion has led to an unnatural accumulation and greater spatial continuity of organic material on the ground in many forests. This material serves both as potential fuel for forest fires and habitat for a large array of forest species. Managers must balance fuel reduction to reduce wildfire hazard with fuel retention targets to maintain other forest functions. This study reports fuel consumption and changes to coarse woody debris attributes with prescribed burns ignited under different fuel moisture conditions. Replicated early season burn, late season burn, and unburned control plots were established in old-growth mixed conifer forest in Sequoia National Park that had not experienced fire for more than 120 years. Early season burns were ignited during June 2002 when fuels were relatively moist, and late season burns were ignited during September/October 2001 when fuels were dry. Fuel loading and coarse woody debris abundance, cover, volume, and mass were evaluated prior to and after the burns. While both types of burns reduced fuel loading, early season burns consumed significantly less of the total dead and down organic matter than late season burns (67% versus 88%). This difference in fuel consumption between burning treatments was significant for most all woody fuel components evaluated, plus the litter and duff layers. Many logs were not entirely consumed – therefore the number of logs was not significantly changed by fire – but burning did reduce log length, cover, volume, and mass. Log cover, volume, and mass were reduced to a lesser extent by early season burns than late season burns, as a result of higher wood moisture levels. Early season burns also spread over less of the ground surface within the burn perimeter (73%) than late season burns (88%), and were significantly patchier. Organic material remaining after a fire can dam sediments and reduce erosion, while unburned patches may help mitigate the impact of fire on fire-sensitive species by creating refugia from which these species can recolonize burned areas. Early season burns may be an effective means of moderating potential ecosystem damage when treating heavy and/or continuous fuels resulting from long periods of fire exclusion, if burning during this season is not detrimental to other forest functions.
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.