Changes in Transpiration and Leaf Water Potential in Douglas-Fir Trees following Douglas-Fir Beetle Attack and Mechanical Girdling

Bark beetles and their associated fungi kill trees readily, but we often ignore which organism is the leading cause of tree mortality. While phloem feeding beetles inhibit photosynthate transport, their associated fungi block the tracheids disrupting transpiration. Within the family Pinaceae, knowledge of tree physiological decline following bark beetle and associated fungi colonization is limited to the genus Pinus. Here we investigate the physiological response of Pseudotsuga (P. menziesii) to bark beetles or its fungi. We hypothesized that fungi block water transport in Douglas-fir causing faster mortality than by bark beetle activity alone. We successfully lured Douglas-fir beetle to attack a subset of trees in our experimental area using pheromones and compared Beetle-Killed trees with mechanically Girdled, and Control trees. During spring snowmelt, nine months after treatments were applied, Control, Girdled, and five trees that Survived beetle attack had higher transpiration rates and less negative pre-dawn water potential than five Beetle-Killed trees. Declines in transpiration and leaf water potential in our Beetle-Killed trees occurred much earlier than those in studies of beetle-attacked lodgepole pines, suggesting stronger defensive traits in Douglas-fir. Our data suggest that, as in pines, bark beetle-associated fungi are the leading cause of mortality in Douglas-fir beetle-attacked trees.

Pathophysiological responses of pine defensive metabolites largely lack differences between pine species but vary with eliciting ophiostomatoid fungal species

Phytopathogenic ophiostomatoid fungi are common associates of bark beetles and contribute to beetle-associated mortality of trees. Mountain pine beetle outbreaks in Canada are facilitating novel associations between its vectored fungi (Grosmannia clavigera, Leptographium longiclavatum and Ophiostoma montium) and jack pine. How the induced defense-related metabolite responses of jack and lodgepole pines vary in response to the fungi is unknown. Understanding this variation is important to clarifying pine susceptibility to and the physiological impacts of infection. We used a comparative metabolite profiling approach to investigate the defense-related signaling, carbon utilization/mobilization, and synthesis responses of both pines to the fungi. Both pine species largely exhibited similar metabolite responses to the fungi. The magnitude of pine metabolite responses positively reflected pathogen virulence. Our findings indicate that pines can recognize and metabolomically respond to novel pathogens, likely due to signals common between the novel fungi and fungi coevolved with the pine. Thus, jack pine is likely as susceptible as lodgepole pine to infections by each of the MPB-vectored fungi. Furthermore, the magnitude of the metabolite responses of both pines varied by the eliciting fungal species, with the most virulent pathogen causing the greatest reduction in carbohydrates and the highest accumulation of defensive terpenes.

Gallery success, brood production, and condition of mountain pine beetles (Coleoptera: Curculionidae) reared in whitebark and lodgepole pine from Alberta, Canada

Breeding pairs of mountain pine beetle (Dendroctonus ponderosae Hopkins) were introduced into freshly cut bolts of whitebark pine (Pinus albicaulis Engelm.) and lodgepole pine (Pinus contorta var. latifolia Engelm. ex S. Watson) in the laboratory. Brood adults emerging from the bolts were collected and galleries were dissected to compare reproductive success, brood production, and adult condition between the two pines. Beetles were more likely to establish egg galleries that produced brood in lodgepole pine than in whitebark pine. Larval gallery density per centimetre of egg gallery was significantly higher in whitebark pine than in lodgepole pine; however, egg galleries also tended to be shorter in whitebark pine bolts, and consequently, brood adults emerging production per gallery did not differ between the two host species. Female body size, mass, and fat content of brood adults and survival from larva to adult did not differ between beetles reared in the two hosts. Though this no-choice assay did not simulate the sequence of events occurring during host selection, these results are consistent with other data suggesting that beetles could be less likely to attack whitebark pines in southwestern Alberta. Whitebark pines that are attacked will produce brood in similar numbers and condition as those from lodgepole pines.

Long-term efficacy of diameter-limit cutting to reduce mountain pine beetle-caused tree mortality in a lodgepole pine forest

Mountain pine beetle, Dendroctonus ponderosae Hopkins, is the most significant mortality agent in pine forests of western North America. Silvicultural treatments that reduce the number of susceptible host trees, alter age and size class distributions, and diversify species composition are considered viable, long-term options for reducing stand susceptibility to mountain pine beetle-caused tree mortality. Short-term efficacy of thinning treatments has been evaluated, but long-term efficacy has not. We evaluated mountain pine beetle-caused lodgepole pine mortality in 2008, ∼28 years after diameterlimit cutting from above that removed the largest diameter lodgepole pines in a Wyoming, USA forest. Following extensive recent mountain pine beetle activity, the partially-cut stands had significantly less mountain pine beetle-caused tree mortality compared to untreated reference stands. These results are similar to observations five years post-treatment, albeit using different reference stands because the original controls were lost to timber harvest. The original management objective was reduced mountain pine beetle-caused tree mortality, and this objective was achieved, lasting for up to 28 years. Despite the reduced mortality among partially-cut stands, however, untreated and treated stands had similar densities of residual live mature lodgepole pine and those in untreated stands had larger average diameters.

Species, elevation, and diameter affect whitebark pine and lodgepole pine stored resources in the sapwood and phloem: implications for bark beetle outbreaks

Stored resources in trees reflect physiological and environmental variables and affect life history traits, including growth, reproduction, resistance to abiotic stress, and defense. However, less attention has been paid to the fact that stored resources also determine tissue nutritional quality and may have direct consequences for the success of herbivores and pathogens. Here, we investigated whether stored resources differed between two hosts of the mountain pine beetle (Dendroctonus ponderosae Hopkins, 1902): lodgepole pine (Pinus contorta Douglas ex. Loudon), a common host, and whitebark pine (Pinus albicaulis Engelmann), a more naïve host that grows at higher altitudes. Phloem and sapwood were sampled in small- and large-diameter trees at two elevations, and nitrogen, phosphorus, nonstructural carbohydrates, and lipids were measured. We found that concentrations of stored resources increased with elevation and tree diameter for both species and that whitebark pine had thicker phloem than lodgepole pine. Overall, stored resources were higher in whitebark pine such that small-diameter whitebark pine trees often had resource concentrations higher than large-diameter lodgepole pines. These results suggest that whitebark pine is of higher nutritional quality than lodgepole pine, which could have implications for the current expansion of mountain pine beetles into higher altitude and latitude forests in response to climate warming.

Pollen and macrofossil evidence of Late Pleistocene and Holocene treeline fluctuations from an alpine lake in Colorado, USA

High-resolution pollen, plant macrofossil and magnetic susceptibility (MS) data are presented from an alpine lake sediment core from west-central Colorado, recording changes in vegetation and sedimentation for the latest Pleistocene and Holocene ( c. the last 12.5 ka; 1 ka = 1000 cal. yr BP). During the Younger Dryas chron ( c. 12.9–11.5 ka), Artemisia steppe or tundra grew around the lake, but by the earliest Holocene (10.7–9.5 ka) a subalpine Picea and Abies parkland was established there. Picea remained important through the early Holocene, but also bristlecone and lodgepole pines ( Pinus aristata and P. contorta) grew around the lake. Warming conditions are indicated from 9.5 ka, lasting until c. 4.5–3.5 ka, which may have been the warmest period, with greatest development of monsoonal conditions. Trees subsequently retreated downslope from Kite Lake c. 150–200 m during the last 3.5 ka, establishing their present treeline position. A decrease in total Pinus and increases in Artemisia and piñon ( P. edulis) indicate a trend toward progressive climate cooling and enhanced winter precipitation. These long-term climatic trends correlate with Holocene changes in summer insolation.

Efficacy of tree defense physiology varies with bark beetle population density: a basis for positive feedback in eruptive species

We evaluated the ability of constitutive and inducible defenses to protect trees and restrict herbivore reproduction across the endemic, incipient (i.e., transitory), and eruptive phases of a native bark beetle species. Host defenses were major constraints when mountain pine beetle (Dendroctonus ponderosae Hopkins) populations were low, but inconsequential after stand-level densities surpassed a critical threshold. We annually examined all lodgepole pines (Pinus contorta Douglas var. latifolia) in six 12–18 ha stands for 3–6 years for beetle attack and establishment as beetle densities progressed through various population phases. We also assayed a suite of tree physiological and chemical attributes and related them to subsequent attacks during that year. Rapidly inducible defenses appeared more important than constitutive defenses, and total monoterpenes were more important than particular constituents. Trees that exude more resin and accumulate higher monoterpene concentrations in response to simulated attack largely escaped natural attacks when populations were low. In stands where beetles had reached incipient densities, these defenses were ineffective. Larger diameter trees had more pronounced defenses than smaller diameter trees. As populations increased, beetles selected increasingly larger, more resource-rich trees, despite their better defenses. When populations were too low for cooperative attack, beetles exploited trees weakened by lower-stem insects. Behavioral plasticity allows beetles to persist at endemic levels until conditions shift, after which positive feedbacks predominate.