José F. Negrón

Climate Change and Bark Beetles of the Western United States and Canada: Direct and Indirect Effects

Climatic changes are predicted to significantly affect the frequency and severity of disturbances that shape forest ecosystems. We provide a synthesis of climate change effects on native bark beetles, important mortality agents of conifers in western North America. Because of differences in temperature-dependent life-history strategies, including cold-induced mortality and developmental timing, responses to warming will differ among and within bark beetle species. The success of bark beetle populations will also be influenced indirectly by the effects of climate on community associates and host-tree vigor, although little information is available to quantify these relationships. We used available population models and climate forecasts to explore the responses of two eruptive bark beetle species. Based on projected warming, increases in thermal regimes conducive to population success are predicted for Dendroctonus rufipennis (Kirby) and Dendroctonus ponderosae Hopkins, although there is considerable spatial and temporal variability. These predictions from population models suggest a movement of temperature suitability to higher latitudes and elevations and identify regions with a high potential for bark beetle outbreaks and associated tree mortality in the coming century.

Probability of infestation and extent of mortality associated with the Douglas-fir beetle in the Colorado Front Range

Infested and uninfested areas within Douglas fir, Pseudotsuga menziesii (Mirb.) Franco, stands affected by the Douglas-fir beetle, Dendroctonus pseudotsugae Hopk. were sampled in the Colorado Front Range, CO. Classification tree models were built to predict probabilities of infestation. Regression trees and linear regression analysis were used to model amount of tree mortality in terms of basal area killed in infested stands. Classification trees had cross-validation estimates of classification accuracy ranging from 0.55 to 0.63. The data suggests that Douglas-fir beetle-attacked stands contain a high percentage of the basal area represented by Douglas-fir, high tree densities, and poor growth during the last 5 years prior to attack. Trees prone to attack by the Douglas-fir beetle within infested points also exhibited reduced growth rates. Tree and linear regression analysis indicate that initial amount of Douglas-fir basal area can be used as a predictor variable for the amount of basal area affected.

Persistent reduced ecosystem respiration after insect disturbance in high elevation forests

Amid a worldwide increase in tree mortality, mountain pine beetles (Dendroctonus ponderosae Hopkins) have led to the death of billions of trees from Mexico to Alaska since 2000. This is predicted to have important carbon, water and energy balance feedbacks on the Earth system. Counter to current projections, we show that on a decadal scale, tree mortality causes no increase in ecosystem respiration from scales of several square metres up to an 84 km2 valley. Rather, we found comparable declines in both gross primary productivity and respiration suggesting little change in net flux, with a transitory recovery of respiration 6–7 years after mortality associated with increased incorporation of leaf litter C into soil organic matter, followed by further decline in years 8–10. The mechanism of the impact of tree mortality caused by these biotic disturbances is consistent with reduced input rather than increased output of carbon.

Effects of Mountain Pine Beetle on Fuels and Expected Fire Behavior in Lodgepole Pine Forests, Colorado, USA

In Colorado and southern Wyoming, mountain pine beetle (MPB) has affected over 1.6 million ha of predominantly lodgepole pine forests, raising concerns about effects of MPB-caused mortality on subsequent wildfire risk and behavior. Using empirical data we modeled potential fire behavior across a gradient of wind speeds and moisture scenarios in Green stands compared three stages since MPB attack (Red [1–3 yrs], Grey [4–10 yrs], and Old-MPB [∼30 yrs]). MPB killed 50% of the trees and 70% of the basal area in Red and Grey stages. Across moisture scenarios, canopy fuel moisture was one-third lower in Red and Grey stages compared to the Green stage, making active crown fire possible at lower wind speeds and less extreme moisture conditions. More-open canopies and high loads of large surface fuels due to treefall in Grey and Old-MPB stages significantly increased surface fireline intensities, facilitating active crown fire at lower wind speeds (>30–55 km/hr) across all moisture scenarios. Not accounting for low foliar moistures in Red and Grey stages, and large surface fuels in Grey and Old-MPB stages, underestimates the occurrence of active crown fire. Under extreme burning conditions, minimum wind speeds for active crown fire were 25–35 km/hr lower for Red, Grey and Old-MPB stands compared to Green. However, if transition to crown fire occurs (outside the stand, or within the stand via ladder fuels or wind gusts >65 km/hr), active crown fire would be sustained at similar wind speeds, suggesting observed fire behavior may not be qualitatively different among MPB stages under extreme burning conditions. Overall, the risk (probability) of active crown fire appears elevated in MPB-affected stands, but the predominant fire hazard (crown fire) is similar across MPB stages and is characteristic of lodgepole pine forests where extremely dry, gusty weather conditions are key factors in determining fire behavior.

Ecosystem CO2/H2O fluxes are explained by hydraulically limited gas exchange during tree mortality from spruce bark beetles

Disturbances are increasing globally due to anthropogenic changes in land use and climate. This study determines whether a disturbance that affects the physiology of individual trees can be used to predict the response of the ecosystem by weighing two competing hypothesis at annual time scales: (a) changes in ecosystem fluxes are proportional to observable patterns of mortality or (b) to explain ecosystem fluxes the physiology of dying trees must also be incorporated. We evaluate these hypotheses by analyzing 6 years of eddy covariance flux data collected throughout the progression of a spruce beetle (Dendroctonus rufipennis) epidemic in a Wyoming Engelmann spruce (Picea engelmannii)–subalpine fir (Abies lasiocarpa) forest and testing for changes in canopy conductance (gc), evapotranspiration (ET), and net ecosystem exchange (NEE) of CO2. We predict from these hypotheses that (a) gc, ET, and NEE all diminish (decrease in absolute magnitude) as trees die or (b) that (1) gc and ET decline as trees are attacked (hydraulic failure from beetle-associated blue-stain fungi) and (2) NEE diminishes both as trees are attacked (restricted gas exchange) and when they die. Ecosystem fluxes declined as the outbreak progressed and the epidemic was best described as two phases: (I) hydraulic failure caused restricted gc, ET (28 ± 4% decline, Bayesian posterior mean ± standard deviation), and gas exchange (NEE diminished 13 ± 6%) and (II) trees died (NEE diminished 51 ± 3% with minimal further change in ET to 36 ± 4%). These results support hypothesis b and suggest that model predictions of ecosystem fluxes following massive disturbances must be modified to account for changes in tree physiological controls and not simply observed mortality.

Traps and Attractants for Wood-Boring Insects in Ponderosa Pine Stands in the Black Hills, South Dakota

Abstract Recent large-scale wildfires have increased populations of wood-boring insects in the Black Hills of South Dakota. Because little is known about possible impacts of wood-boring insects in the Black Hills, land managers are interested in developing monitoring techniques such as flight trapping with semiochemical baits. Two trap designs and four semiochemical attractants were tested in a recently burned ponderosa pine, Pinus ponderosa Dougl. ex Laws., forest in the Black Hills. Modified panel and funnel traps were tested in combination with the attractants, which included a woodborer standard (ethanol and α-pinene), standard plus 3-carene, standard plus ipsenol, and standard plus ipsdienol. We found that funnel traps were equally efficient or more efficient in capturing wood-boring insects than modified panel traps. Trap catches of cerambycids increased when we added the Ips spp. pheromone components (ipsenol or ipsdienol) or the host monoterpene (3-carene) to the woodborer standard. During the summers of 2003 and 2004, 18 cerambycid, 14 buprestid, and five siricid species were collected. One species of cerambycid, Monochamus clamator (LeConte), composed 49 and 40% of the 2003 and 2004 trap catches, respectively. Two other cerambycids, Acanthocinus obliquus (LeConte) and Acmaeops proteus (Kirby), also were frequently collected. Flight trap data indicated that some species were present throughout the summer, whereas others were caught only at the beginning or end of the summer.

Stand Conditions Associated with Roundheaded Pine Beetle (Coleoptera: Scolytidae) Infestations in Arizona and Utah

Abstract Stand conditions associated with outbreak populations of the roundheaded pine beetle, Dendroctonus adjunctus Blandford, in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests were studied in the Pinaleno Mountains, AZ, and the Pine Valley Mountains, UT. Classification tree models to estimate the probability of infestation based on stand attributes were built for both study areas using growth rate and ponderosa pine basal area information. Cross-validation estimates of correct classification were 0.60 for the Pinaleno Mountains and 0.58 and 0.78 for the Pine Valley Mountains. Regression tree and linear regression models to estimate amount of mortality caused by the beetles were also built for both sites using growth rate, ponderosa pine basal area, and trees per hectare information. The occurrence and mortality levels caused by the roundheaded pine beetle are positively related at both the stand and tree scale with reduced growth rates caused by high stocking densities.

Within-stand spatial distribution of tree mortality caused by the Douglas-Fir beetle (Coleoptera: Scolytidae)

Abstract The Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins, causes considerable mortality in Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, forests. Within-stand distribution of mortality was examined in affected stands using geostatistical techniques. A 10 × 10 m grid was established in two 4-ha study sites. Live and beetle-killed host basal area was measured at each node. In a 16-ha stand, a variable-resolution grid was established and the same information collected. The relationship between Douglas-fir basal area and Douglas-fir basal area killed was examined using non-spatially explicit and spatially explicit linear regression models. A positive linear relationship was observed between the variables. Significant spatially explicit models suggest that the relationship is also true at fine scales. Relative variograms were constructed for Douglas-fir basal area before and after the Douglas-fir beetle outbreaks. For the 4-ha sites, increased spatial dependency in the distribution of Douglas-fir basal area was observed as a result of the Douglas-fir beetle outbreak. For the 16-ha site, kriging was used to estimate live Douglas-fir basal area before and after the outbreak to a 10-m resolution and the stand rated for potential mortality illustrating the potential applicability of geostatistical techniques to rating a stand for potential mortality. Cross-validation analysis indicated that although the potential exists for large estimation errors, the majority of the estimates were within acceptable ranges. The study suggests that geostatistical approaches may be suitable to extend our understanding bark beetle ecology and improving the application of extent of mortality models.

Semiochemical-Mediated Flight Strategies of Two Invasive Elm Bark Beetles: A Potential Factor in Competitive Displacement

ABSTRACT A seven-state survey showed that the recently detected invasive Asian banded elm bark beetle, Scolytus schevyrewi Semenov, was abundant in areas of Colorado and Wyoming, whereas the long-established European elm bark beetle, S. multistriatus (Marsham), was not as abundant. In one of a series of studies to evaluate whether S. schevyrewi is competitively displacing S. multistriatus in their North American zone of sympatry, we characterized long-range flight responses infested or uninfested to small cut logs of American, Chinese, and Siberian elm, Ulmus americana, U. parvifolia, and U. pumila. Trials were conducted in Colorado and Wyoming to test the flight response of S. schevyrewi; in California to test the response of S. multistriatus; and in Nevada to test the responses of both species simultaneously. Studies with S. schevyrewi showed that males and females aggregated toward Ulmus spp. host volatiles but provided no evidence of a putative aggregation pheromone during the 0- to 48- or 48- to 96-h period of infestation. In contrast, S. multistriatus was attracted to U. pumila over unbaited controls, more to U. pumila infested with conspecific females than without, and more during the 48- to 96- versus 0- to 48-h period of infestation. This confirmed that male and female S. multistriatus aggregated toward host volatiles and that females produced an aggregation pheromone. In a cross-attraction study, S. schevyrewi displayed neither flight preference nor interruption to 17. pumila infested with conspecifics, heterospecifics, or a mix of both species. Response of S. multistriatus was too low to draw conclusions. Although S. multistriatus aggregates moderately to host volatiles and strongly to female-derived pheromones emitted after a few days, S. multistriatus may have a relative disadvantage by selecting elm hosts more slowly than S. schevyrewi, which aggregates very strongly to host volatiles. The differential long-range host location strategy may be one factor in a chain of behavioral events that leads to advantageous host colonization and development by S. schevyrewi.

The flight periodicity, attack patterns, and life history of Dryocoetes confusus Swaine (Coleoptera: Curculionidae: Scolytinae), the western balsam bark beetle, in north central Colorado.

ABSTRACT. Dryocoetes confusus Swaine, the western balsam bark beetle, is an important bark beetle associated with Abies lasiocarpa (Hook.) Nutt. (subalpine fir) in western North America. Little information is available on the life cycle and ecology of this insect in Colorado. In this study in north central Colorado, we examined its flight periodicity, attack patterns, and life cycle in downed trees. Flight season, as indicated by pheromone-trap catches, began in early June. Peak flight was observed during early to mid-July. A second peak in flight was observed in some years, but its occurrence was not consistent. Preferred attack sites on downed trees were the underside or side of the log and through branch stubs. The life cycle for this beetle was completed in 2 years, consistent with earlier observations from British Columbia. Sex ratio was 1:1, but a higher percentage of males fly earlier in the season.