To test the hypothesis that drought predisposes trees to insect attacks, we quantified the effects of water availability on insect attacks, tree resistance mechanisms, and mortality of mature piñon ...pine (Pinus edulis) and one-seed juniper (Juniperus monosperma) using an experimental drought study in New Mexico, USA.
The study had four replicated treatments (40 × 40 m plot/replicate): removal of 45% of ambient annual precipitation (H2O−); irrigation to produce 125% of ambient annual precipitation (H2O+); a drought control (C) to quantify the impact of the drought infrastructure; and ambient precipitation (A).
Piñon began dying 1 yr after drought initiation, with higher mortality in the H2O− treatment relative to other treatments. Beetles (bark/twig) were present in 92% of dead trees. Resin duct density and area were more strongly affected by treatments and more strongly associated with piñon mortality than direct measurements of resin flow. For juniper, treatments had no effect on insect resistance or attacks, but needle browning was highest in the H2O− treatment.
Our results provide strong evidence that ≥ 1 yr of severe drought predisposes piñon to insect attacks and increases mortality, whereas 3 yr of the same drought causes partial canopy loss in juniper.
Conifers have incurred high mortality during recent global-change-type drought(s) in the western USA. Mechanisms of drought-related tree mortality need to be resolved to support predictions of the ...impacts of future increases in aridity on vegetation. Hydraulic failure, carbon starvation and lethal biotic agents are three potentially interrelated mechanisms of tree mortality during drought. Our study compared a suite of measurements related to these mechanisms between 49 mature piñon pine (Pinus edulis Engelm.) trees that survived severe drought in 2002 (live trees) and 49 trees that died during the drought (dead trees) over three sites in Arizona and New Mexico. Results were consistent over all sites indicating common mortality mechanisms over a wide region rather than site-specific mechanisms. We found evidence for an interactive role of hydraulic failure, carbon starvation and biotic agents in tree death. For the decade prior to the mortality event, dead trees had twofold greater sapwood cavitation based on frequency of aspirated tracheid pits observed with scanning electron microscopy (SEM), smaller inter-tracheid pit diameter measured by SEM, greater diffusional constraints to photosynthesis based on higher wood δ(13)C, smaller xylem resin ducts, lower radial growth and more bark beetle (Coleoptera: Curculionidae) attacks than live trees. Results suggest that sapwood cavitation, low carbon assimilation and low resin defense predispose piñon pine trees to bark beetle attacks and mortality during severe drought. Our novel approach is an important step forward to yield new insights into how trees die via retrospective analysis.
We assessed attraction of pine engraver, Ips pini (Say) (Coleoptera: Curculionidae; Scolytinae), to pheromone-baited funnel traps treated with repellent semiochemicals in ponderosa pine, Pinus ...ponderosa var. scopulorum Engelm., forests in northern Arizona. Treatments included: 1) baited control (B, ipsdienol + lanierone), 2) 70 g of SPLAT Verb (a flowable, biodegradable formulation containing 10% verbenone, ISCA Technologies Inc., Riverside, CA, USA) + B, 3) 70 g of SPLAT Verb + (E)-2-hexen-1-ol+(Z)-2-hexen-1-ol + acetophenone + B, 4) 7.84-g verbenone pouch (Product #3413, Synergy Semiochemicals Corp., Delta, British Columbia, Canada) + B, and 5) 7.84-g verbenone pouch + (E)-2-hexen-1-ol+(Z)-2-hexen-1-ol + acetophenone + B. In total, 472 I. pini were collected. Trap catches were highest in baited traps and declined significantly with the addition of both formulations of verbenone. Traps treated with SPLAT Verb caught significantly fewer I. pini and male I. pini than those treated with verbenone pouches. The addition of (E)-2-hexen-1-ol+(Z)-2-hexen-1-ol + acetophenone to SPLAT Verb and the verbenone pouch had no effect on trap catch. Verbenone has potential as an effective tool for protecting P. ponderosa trees and slash from I. pini in northern Arizona, but the addition of (E)-2-hexen-1-ol+(Z)-2-hexen-1-ol + acetophenone to verbenone is unwarranted.
Lure attraction strength for red turpentine beetle, Dendroctonus valens (Coleoptera: Curculionidae: Scolytinae) observed previously in US Pacific Northwest ponderosa pine forests is ...(−)‐β‐pinene+ethanol > (+)‐3‐carene+ethanol, but untested elsewhere in its western US range. Thus, both were tested with (−)‐β‐pinene, (+)‐3‐carene, ethanol, and a blank in Oregon and California sites burned by wildfire, whereas in Arizona the first four lures were tested in a thinned‐unburned site.
The D. valens responses in burned Oregon and California sites were similar, (−)‐β‐pinene+ethanol > (−)‐β‐pinene > 3‐carene = 3‐carene+ethanol > ethanol > blank, whereas in the cut‐unburned Arizona site it was 3‐carene+ethanol > 3‐carene = (−)‐β‐pinene+ethanol > (−)‐β‐pinene. Whether this variation was influenced by beetle genetic differences, or chemical and physical parameters in the different environments and remaining stressed host resources 1‐year post disturbance warrants additional study.
Responses to (−)‐β‐pinene varied, from a stronger attractant than (+)‐3‐carene in Oregon and California, to a weaker lure than (+)‐3‐carene in Arizona. This (−)‐β‐pinene variability was minimized when released in combination with ethanol, making (−)‐β‐pinene+ethanol the most consistent attractant of those tested across the three states, and a reliable lure for detection, monitoring, and management projects for D. valens in western US pine forests.
Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications ...for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.
Predictions of warmer droughts causing increasing forest mortality are becoming abundant, yet few studies have investigated the mechanisms of forest persistence. To examine the resistance of forests ...to warmer droughts, we used a five-year precipitation reduction (∼45% removal), heat (+4 °C above ambient) and combined drought and heat experiment in an isolated stand of mature Pinus edulis-Juniperus monosperma. Despite severe experimental drought and heating, no trees died, and we observed only minor evidence of hydraulic failure or carbon starvation. Two mechanisms promoting survival were supported. First, access to bedrock water, or 'hydraulic refugia' aided trees in their resistance to the experimental conditions. Second, the isolation of this stand amongst a landscape of dead trees precluded ingress by Ips confusus, frequently the ultimate biotic mortality agent of piñon. These combined abiotic and biotic landscape-scale processes can moderate the impacts of future droughts on tree mortality by enabling tree avoidance of hydraulic failure, carbon starvation, and exposure to attacking abiotic agents.
Recent outbreaks of engraver beetles, Ips spp. De Geer (Coleoptera: Curculionidae; Scolytinae), in ponderosa pine, Pinus ponderosa var. scopulorum Engelm. (Pinales: Pinaceae), forests of northern ...Arizona have resulted in widespread tree mortality. Current treatment options, such as spraying individual P. ponderosa with insecticides or deep watering of P. ponderosa in urban and periurban settings, are limited in applicability and scale. Thinning stands to increase tree vigor is also recommended, but appropriate timing is crucial. Antiaggregation pheromones, widely used to protect high-value trees or areas against attacks by several species of Dendroctonus Erichson (Coleoptera: Curculionidae; Scolytinae), would provide a feasible alternative with less environmental impacts than current treatments. We evaluated the efficacy of the antiaggregation pheromone verbenone (4,6,6-trimethylbicyclo3.1.1hept-3-en-2-one) in reducing attraction of pine engraver, I. pini (Say), to funnel traps baited with their aggregation pheromone in two trapping assays. Treatments included 1) unbaited control, 2) aggregation pheromone (bait), 3) bait with verbenone deployed from a pouch, and 4) bait with verbenone deployed from a flowable and biodegradable formulation (SPLAT Verb, ISCA Technologies Inc., Riverside, CA). Unbaited traps caught no beetles. In both assays, baited traps caught significantly more I. pini than traps with either formulation of verbenone, and no significant difference was observed between the verbenone pouch and SPLAT Verb. In the second assay, we also examined responses of Temnochila chlorodia (Mannerheim) (Coleoptera: Trogositidae), a common bark beetle predator. Traps containing verbenone pouches caught significantly fewer T. chlorodia than the baited control and SPLAT Verb treatments. We conclude that verbenone shows promise for reducing tree mortality from I. pini.
Abstract
Thinning is a necessary silvicultural activity for restoring the long-term sustainability of pine forests in much of the southwestern United States. In northern Arizona, a landscape-scale ...restoration effort, called the Four Forest Restoration Initiative, has been implemented to recover the long-term sustainability of 2.4 million acres on four national forests. Cost-effective and efficient thinning methods are needed due to the scale of the project to help improve habitat, conserve biodiversity, protect old growth, reduce risk of severe wildfire, and restore natural forest structure and function. Mechanical cutting using a feller-buncher is the primary method of thinning in these forests due to the extreme high number of small-diameter ponderosa pine trees. A feller-buncher places harvested trees into small piles known as “bunches.” In this review, we highlight advantages and disadvantages of bunching tree materials in restoration programs and review published studies on transpirational drying of bunches of various tree species in different forest habitats across the United States, including ponderosa pine in Arizona. Studies show that transpirational drying of trees in bunches can be an effective method to allow for wood drying, but this process can be affected by abiotic factors associated with seasonal climate and stand characteristics.
Study Implications
The Four Forest Restoration Initiative makes up the largest landscape-level collaborative project in the history of the USDA Forest Service with more than 2.4 million acres of forest habitat. Thinning is a necessary silvicultural activity for restoring the long-term sustainability of these forests in northern Arizona. Because of the extremely high number (i.e., average of 720 trees per acre) of small-diameter ponderosa pine (Pinus ponderosa) trees, mechanical cutting is more cost effective than individual saw cutting, which is why a feller-buncher machine is the primary thinning tool. During feller-buncher thinning, small piles of trees known as “bunches” are created. Bunching trees can allow for wood drying in the field that translates into lower operational costs because drier trees are lighter in weight, which reduces transportation costs. In northern Arizona, a 60-day time frame is allotted for transpirational drying before tree materials must be removed from the forest. However, because the drying process through bunching is affected by abiotic conditions, there may be a need to adjust this time frame to account for seasonal weather patterns. For example, during spring, when weather in northern Arizona is hot and dry, less time might be necessary for tree bunches to lose enough moisture while still rendering the thinning operation profitable and simultaneously avoiding bark beetle proliferation given that their life cycle consists of about 40 days. Furthermore, bunching studies should be developed to look at individual tree species in their respective locations and to investigate the effects of the presence, partial presence, or absence of branches and bark on trees within bunches. Studies should also be conducted to look at the effects of bunches on insect communities, particularly those that can cause extensive tree mortality.
Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, trees and stands can be protected from Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins (DFB)-caused mortality by application of synthetic ...formulations of the beetle's antiaggregation pheromone, 3-methylcyclohex-2-en-1-one (MCH). A biodegradable formulation of MCH, SPLAT MCH, was developed and evaluated for protecting individual Douglas-fir trees and small stands from colonization and mortality by DFB. In an individual-tree experiment in Idaho, both MCH bubble capsules and SPLAT MCH significantly reduced the proportion of treated trees colonized and killed by DFB compared to untreated controls. SPLAT MCH was as effective as MCH bubble capsules for protecting individual trees. Both MCH bubble capsules and SPLAT MCH significantly reduced the proportion of trees colonized and killed by DFB within 0.04-ha circular plots surrounding each treated tree compared to untreated controls. In 0.41 ha stands in New Mexico, both MCH bubble capsules and SPLAT MCH significantly reduced the proportion of trees colonized and killed by DFB compared to untreated controls, again with no differences observed between MCH treatments. In a similar stand level trial in Idaho, neither MCH treatment significantly reduced the proportion of trees colonized by DFB, and only MCH bubble capsules significantly reduced levels of tree mortality compared to untreated controls, but no significant difference was observed between SPLAT MCH and MCH bubble capsules. Overall, the results indicate that SPLAT MCH is as effective as MCH bubble capsules for protecting individual trees and small stands of Douglas-fir from DFB-caused mortality.
1
Bark beetles are significant mortality agents of conifers. Four beetle species, the pine engraver Ips pini, the six‐spined pine engraver Ips calligraphus sub. ponderosae, the southern pine beetle ...Dendroctonus frontalis, and the western pine beetle Dendroctonus brevicomis, cohabitate pines in Arizona.
2
A pheromone trapping study in ponderosa forests of Arizona determined the attraction of beetles to conspecific and heterospecific pheromone components in the presence and absence of host volatiles, and tested whether predators differ in their attraction to combinations of pheromone components and tree monoterpenes.
3
All four bark beetle species differed in their responses to heterospecific lures and monoterpenes. Ips calligraphus was the only species that increased in trap catches when heterospecific lures were added. Heterospecific lures did not inhibit the attraction of either Dendroctonus or Ips species. The replacement of myrcene with α‐pinene increased the attraction of Dendroctonus, whereas the addition of α‐pinene had mixed results for Ips. The prominent predators Temnochila chlorodia and Enoclerus lecontei were more attracted to the I. pini lure than the D. brevicomis lure, and the combination of the two lures with α‐pinene was most attractive to both predator species.
4
Cross attraction and limited inhibition of bark beetles to heterospecific pheromones suggest that some of these species might use heterospecific compounds to increase successful location and colonization of trees. Predator responses to treatments suggest that tree volatiles are used to locate potential prey and predators are more responsive to Ips than to Dendroctonus pheromone components in Arizona.
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