Bayesian Predictions of Bark Beetle Attack and Mortality of Three Conifer Species During Epidemic and Endemic Population Stages

Bark beetles naturally inhabit forests and can cause large-scale tree mortality when they reach epidemic population numbers. A recent epidemic (1990s–2010s), primarily driven by mountain pine beetles (Dendroctonus ponderosae), was a leading mortality agent in western United States forests. Predictive models of beetle populations and their impact on forests largely depend on host related parameters, such as stand age, basal area, and density. We hypothesized that bark beetle attack patterns are also dependent on inferred beetle population densities: large epidemic populations of beetles will preferentially attack large-diameter trees, and successfully kill them with overwhelming numbers. Conversely, small endemic beetle populations will opportunistically attack stressed and small trees. We tested this hypothesis using 12 years of repeated field observations of three dominant forest species (lodgepole pine Pinus contorta, Engelmann spruce Picea engelmannii, and subalpine fir Abies lasiocarpa) in subalpine forests of southeastern Wyoming paired with a Bayesian modeling approach. The models provide probabilistic predictions of beetle attack patterns that are free of assumptions required by frequentist models that are often violated in these data sets. Furthermore, we assessed seedling/sapling regeneration in response to overstory mortality and hypothesized that higher seedling/sapling establishment occurs in areas with highest overstory mortality because resources are freed from competing trees. Our results indicate that large-diameter trees were more likely to be attacked and killed by bark beetles than small-diameter trees during epidemic years for all species, but there was no shift toward preferentially attacking small-diameter trees in post-epidemic years. However, probabilities of bark beetle attack and mortality increased for small diameter lodgepole pine and Engelmann spruce trees in post-epidemic years compared to epidemic years. We also show an increase in overall understory growth (graminoids, forbs, and shrubs) and seedling/sapling establishment in response to beetle-caused overstory mortality, especially in lodgepole pine dominated stands. Our observations provide evidence of the trajectories of attack and mortality as well as early forest regrowth of three common tree species during the transition from epidemic to post-epidemic stages of bark beetle populations in the field.

Climate-induced treeline mortality during the termination of the Little Ice Age in the Greater Yellowstone Ecoregion, USA

Paleoclimate reconstructions for the western US show spatial variability in the timing, duration, and magnitude of climate changes within the Medieval Climate Anomaly (MCA, ca. 900–1350 CE) and Little Ice Age (LIA, ca. 1350–1850 CE), indicating that additional data are needed to more completely characterize late-Holocene climate change in the region. Here, we use dendrochronology to investigate how climate changes during the MCA and LIA affected a treeline, whitebark pine ( Pinus albicaulis Engelm.) ecosystem in the Greater Yellowstone Ecoregion (GYE). We present two new millennial-length tree-ring chronologies and multiple lines of tree-ring evidence from living and remnant whitebark pine and Engelmann spruce ( Picea engelmannii Parry ex. Engelm.) trees, including patterns of establishment and mortality; changes in tree growth; frost rings; and blue-intensity-based, reconstructed summer temperatures, to highlight the terminus of the LIA as one of the coldest periods of the last millennium for the GYE. Patterns of tree establishment and mortality indicate conditions favorable to recruitment during the latter half of the MCA and climate-induced mortality of trees during the middle-to-late LIA. These patterns correspond with decreased growth, frost damage, and reconstructed cooler temperature anomalies for the 1800–1850 CE period. Results provide important insight into how past climate change affected important GYE ecosystems and highlight the value of using multiple lines of proxy evidence, along with climate reconstructions of high spatial resolution, to better describe spatial and temporal variability in MCA and LIA climate and the ecological influence of climate change.

Toxicity of two Engelmann spruce (Pinaceae) monoterpene chemotypes from the southern Rocky Mountains to North American spruce beetle (Coleoptera: Scolytidae)

Engelmann spruce, Picea engelmannii Parry ex Engelm. (Pinaceae), in the southern Rocky Mountains is composed of two distinct phloem monoterpene chemotypes that differ in relative abundances of multiple monoterpenes, particularly α-pinene and Δ3-carene (hereafter, the “α-pinene chemotype” and the “Δ3-carene chemotype”). Here, relative toxicity of these chemotypes is tested on spruce beetle (Dendroctonus rufipennis Kirby) (Coleoptera: Scolytinae), a phloeophagous herbivore that colonises trees of both types. Synthetic monoterpene blends representing each chemotype were tested across a range of concentrations (0, 10, 50, 100, 200, and 500 µg/L) in the lab, and probability of survival of adult beetles exposed to each blend was modelled using a logit function. Logit curves were solved to determine LC25, LC50, and LC75 of each monoterpene blend. On average, probability of beetle survival was lower when exposed to the Δ3-carene chemotype than when exposed to the α-pinene chemotype. However, both chemotypes were completely lethal to beetles at concentrations exceeding 100 µg/L. Adult body mass did not affect survival probability. It is concluded that spruce phloem chemotypes may differ in their toxicity to spruce beetles, with potential consequences for patterns of host-tree colonisation by spruce beetle.

Post-Wildfire Regeneration in a Sky-Island Mixed- Conifer Ecosystem of the North American Great Basin

Information on wildfire impacts and ecosystem responses is relatively sparse in the Great Basin of North America, where subalpine ecosystems are generally dominated by five-needle pines. We analyzed existing vegetation, with an emphasis on regeneration following the year 2000 Phillips Ranch Fire, at a sky-island site in the Snake Range of eastern Nevada. Our main objective was to compare bristlecone pine (Pinus longaeva; PILO) post-fire establishment and survival to that of the co-occurring dominant conifers limber pine (Pinus flexilis; PIFL) and Engelmann spruce (Picea engelmannii; PIEN) in connection with site characteristics. Field data were collected in 40 circular 0.1 ha plots (17.8 m radius) randomly located using GIS so that half of them were inside (“burned”) and half were outside (“unburned”) the 2000 fire boundary. While evidence of previous burns was also found, we focused on impacts from the Phillips Ranch Fire. Mean total basal area, including live and dead stems, was not significantly different between plots inside the burn and plots outside the fire perimeter, but the live basal area was significantly less in the former than in the latter. Wildfire impacts did not limit regeneration, and indeed bristlecone seedlings and saplings were more abundant in plots inside the 2000 fire perimeter than in those outside of it. PILO regeneration, especially saplings, was more abundant than PIFL and PCEN combined, indicating that PILO can competitively regenerate under modern climatic conditions. Surviving PILO regeneration in burned plots was also taller than that of PIFL. By contrast, PCEN was nearly absent in the plots that had been impacted by fire. Additional research should explicitly address how climatic changes and disturbance processes may interact in shaping future vegetation dynamics.

Topographic control of forest species distributions in the Interior West USA

Background The geographic distribution of forest and woodland ecosystems in the Interior West United States is strongly influenced by topographic gradients that, in part, control moisture availability through their effect on insolation, and precipitation capture and retention. Through an empirical approach, we use unique, plot-level data from the Forest Inventory and Analysis Program ( n = 13,437) over eight ecoregions within eight Interior West states to characterize the distribution of the 12 most abundant tree species with respect to the effects of elevation, slope aspect, and slope steepness. Results Across species, elevation, and aspect, most plots occurred on gentle slopes and the number decreased with increasing slope. Species-specific differences to microenvironmental conditions were evident in the variation between observed (plots containing a subject tree) and expected (all forest plots from the systematic sample) numbers of plots across the gradient combinations. Species groups, broadly defined as woodland, montane, and subalpine, generally exhibited similar responses and revealed more generality than hypothesized. Only Douglas-fir, white fir, subalpine fir, and Engelmann spruce exhibited significant patterns of affinity for particular aspects—most often on north and least often on south—with the relative importance of south aspects decreasing with increasing elevation. Limber pine showed unique, unimodal patterns of affinity for moderately steep slopes, with no consistent patterns by aspect or elevation. Although not significant, at high elevations woodland species exhibited a tendency to occur more often on south aspects on gentle to intermediate slopes, and less often on north aspects. Conclusions Unique microenvironments created by interactions between aspect, slope, and elevation create some predictability in patterns of geographic distribution. However, the general lack of species-specific response suggests that patterns of occurrence in relation to physiographic gradients is much broader than in common generalizations.

Evaluations of Bole Injections for Protecting Engelmann Spruce from Mortality Attributed to Spruce Beetle (Coleoptera: Curculionidae) in the Intermountain West

Bark beetles are important disturbance agents in coniferous forests, and spruce beetle, Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae), is one of the more notable species causing landscape-level tree mortality in western North America. We evaluated the efficacy of bole injections of emamectin benzoate (TREE-äge®; Arborjet Inc., Woburn, MA) alone and combined with propiconazole (Alamo®; Syngenta Crop Protection Inc., Wilmington, DE) for protecting Engelmann spruce, Picea engelmannii Parry ex Engelmann (Pinales: Pinaceae), from mortality attributed to colonization by D. rufipennis. Two injection periods in 2013 (the spring and fall of the year prior to trees first being challenged by D. rufipennis in 2014) and distributions of injection points (7.6- and 15.2-cm spacings) were evaluated. Tree mortality was monitored over a 3-yr period (2014–2017). Emamectin benzoate injected in spring at a narrow spacing (7.6 cm) was the only effective treatment. Two (but not three) field seasons of protection can be expected with a single injection of this treatment. We discuss the implications of these and other results regarding the use of emamectin benzoate and propiconazole for protecting western conifers from mortality attributed to bark beetles, and provide suggestions for future research. A table summarizing the appropriate timing of treatments in different bark beetle/host systems is provided.

Responses of Engelmann spruce to inoculation with Leptographium abietinum, a symbiotic fungus of the North American spruce beetle

Symbiotic fungi associated with tree-killing bark beetles can alter host-tree physiology with consequences for tree survival, and symbiont genetic variation and environmental variability may impact these interactions. Here, we test whether multiple genetically distinct isolates of a symbiotic fungus (Leptographium abietinum (Peck) M.J. Wingf.) associated with North American spruce beetle (Dendroctonus rufipennis (Kirby, 1837)) vary in their ability to manifest defensive responses consistent with disease symptoms in seedlings of Engelmann spruce (Picea engelmannii Parry ex Engelm.), a primary host-tree species for the beetle–fungus complex in North America. Our experiments incorporate variation in both host-tree water availability and host defenses (phloem monoterpene concentration). Three central findings emerged: (i) isolates varied considerably in their effects on host trees — inoculation with L. abietinum isolates from Colorado caused significantly larger phloem lesions than isolates from Wyoming, though all isolates caused phloem oxidation; (ii) neither water availability nor spruce phloem monoterpene concentrations impacted lesion formation; and (iii) both inoculation with L. abietinum and water deficit inhibited the formation of callus tissue at wound sites. We conclude that L. abietinum isolates vary in their virulence and that inoculation is not lethal but may benefit beetles by altering tree defensive responses.