Beetle Attack Research Articles

Lethal Trap Trees and Semiochemical Repellents as Area Host Protection Strategies for Spruce Beetle (Coleoptera: Curculionidae, Scolytinae) in Utah.

We tested lethal trap trees and repellent semiochemicals as area treatments to protect host trees from spruce beetle (Dendroctonus rufipennis Kirby) attacks. Lethal trap tree treatments ("spray treatment") combined a spruce beetle bait with carbaryl treatment of the baited spruce. Repellent treatments ("spray-repellent") combined a baited lethal trap tree within a 16-m grid of MCH (3-methylcyclohex-2-en-1-one) and two novel spruce beetle repellents. After beetle flight, we surveyed all trees within 50 m of plot center, stratified by 10-m radius subplots, and compared attack rates to those from baited and unbaited control plots. Compared to the baited controls, spruce in the spray treatment had significantly reduced likelihood of a more severe attack classification (e.g., mass-attacked over strip-attacked or unsuccessful-attacked over unattacked). Because spruce in the spray treatment also had significantly heightened probability of more severe attack classification than those in the unbaited controls, however, we do not recommend lethal trap trees as a stand-alone beetle suppression strategy for epidemic beetle populations. Spruce in the spray-repellent treatment were slightly more likely to be classified as more severely attacked within 30 m of plot center compared to unbaited controls but, overall, had reduced probabilities of beetle attack over the entire 50-m radius plots. The semiochemical repellents deployed in this study were effective at reducing attacks on spruce within treated plots despite the presence of a centrally located spruce beetle bait. Further testing will be required to clarify operational protocols such as dose, elution rate, and release device spacing.

Ethanol‐injection induces attacks by ambrosia beetles (Coleoptera: Curculionidae: Scolytinae) on a variety of tree species

Abstract Exotic ambrosia beetles have become serious pests in ornamental tree nurseries. However, research on their biology, ecology and behaviour to develop management tactics is hampered by the lack of reliable attacks on experimental trees. Injecting Magnolia virginiana L. with ethanol consistently induced attacks by exotic ambrosia beetles to facilitate research on their biology and management. The ability to attract ambrosia beetles to trap trees may be useful for developing nonchemical strategies and techniques for managing them in nurseries. However, to be useful in management techniques, ethanol‐injection has to reliably induce attacks on a vlariety of tree species. Experiments were conducted in Ohio, Tennessee and Virginia, where ethanol‐injection was tested on 16 species of deciduous trees from 11 families. Ethanol‐injection reliably induced attacks by exotic ambrosia beetles on all tree species tested. There were differences in the numbers of attacks by ambrosia beetles among species of trees in most experiments, although no species was identified consistently as the most or least attacked in repeated experiments. The positive association of attacks with ethanol concentrations in bark tissue indicate that the attack rates were most likely related to the uptake and emission of ethanol. The results of the present study indicate that ethanol‐injection reliably induces ambrosia beetle attacks on a wide range of deciduous tree species, which should further facilitate research on ambrosia beetles in nurseries and the development of behaviour‐based management tactics that divert ambrosia beetles away from ornamental trees.

Pre-treatment with salicylic acid induces phenolic responses of Norway spruce (Picea abies) bark to bark beetle (Ips typographus) attack

The temporal gradations of the investigated phenolics in Norway spruce bark after bark beetle (Ips typographus) attack followed the general eco-physiological concept. Treatment with salicylic acid inhibits bark beetle colonisation, alleviates the phenolic responses and activates the synthesis of condensed tannins on later sampling dates. Conifer bark is the target of numerous organisms due to its assimilated transport and nutrient storage functions. In the presented study, 100 mM salicylic acid (SA) was applied onto Norway spruce stems prior to being infested with bark beetles (Ips typographus L.), to study the temporal gradation of changes in condensed tannins (CT) and total phenolics (tPH) and their significance for mediating stress-tolerance. A significant accumulation of CT was monitored in untreated trees in response to progressive bark beetle infestation occurring from May onwards. In SA-treated infested trees, the CT values remained at control levels until May, but after the re-treatment of infested trees in June, the concentrations of CT rose significantly in comparison to the controls. The tPH values dropped 16 days after SA-treatment, independent of infestation, and later on remained at control level until July. In contrast, tPH contents accumulated in untreated infested trees in May, eased in June and increased again in July, when the trees were affected by the second generation of bark beetles. To sum up, in May and July when the highest beetle-flight activity was monitored the metabolic shift of phenolics within untreated infested trees differed significantly from the response of SA-treated trees. In addition, on SA-treated trees less entrance holes were monitored over the whole period of sampling when compared to untreated infested trees. These results provide evidence that SA-treatment alleviates the phenolic responses, activates the synthesis of condensed tannins and inhibits bark beetle colonisation.

Developing a Media Moisture Threshold for Nurseries to Reduce Tree Stress and Ambrosia Beetle Attacks.

Exotic ambrosia beetles are among the most damaging pests of trees grown in nurseries. The primary pests Xylosandrus crassiusculus Motschulsky and Xylosandrus germanus Blandford use ethanol to locate vulnerable trees. Research, primarily with X. germanus, has shown that flood-stressed trees emit ethanol and are preferentially attacked by ambrosia beetles. Our goal was to develop a media (also called potting soil) moisture threshold as an integrated pest management (IPM) tactic and assess grower practices that lead to ambrosia beetle attacks. Flooded Cornus florida L., Cornus kousa Burg., and Magnolia grandiflora L. trees incurred more attacks than unflooded trees that were not attacked. To determine optimal media moisture levels, we grew flood-tolerant Acer rubrum L. and flood-intolerant C. florida in containers with 10, 30, 50, 70, or 90% media moisture. No flooded or unflooded A. rubrum were attacked. However, C. florida grown in 70 or 90% moisture were attacked and died, whereas trees at 30 and 50% moisture were not attacked. Thus, we suggest an upper moisture threshold of 50% when growing C. florida and other flood-intolerant trees. However, during peak ambrosia beetle flight activity in spring 2013 and 2014, we found that media moisture levels in commercial nurseries were often between 50 and 90%. Implementing a media moisture threshold, as a new IPM tool, could reduce ambrosia beetle attacks and the need for insecticide applications, which is currently the only available management tactic. Future research should focus on how changes in substrates, irrigation, and other practices could help growers meet this threshold.

The impact of an invasive ambrosia beetle on the riparian habitats of the Tijuana River Valley, California.

The Tijuana River Valley is the first natural habitat in California to be substantially invaded by the Kuroshio Shot Hole Borer (KSHB, Euwallacea sp.), an ambrosia beetle native to Southeast Asia. This paper documents the distribution of the KSHB in the riparian vegetation in the valley and assesses the damage done to the vegetation as of early 2016, approximately six months after the beetle was first observed in the valley. I divided the riparian habitats into 29 survey units so that the vegetation within each unit was relatively homogenous in terms of plant species composition, age and density. From a random point within each unit, I examined approximately 60 individuals of the dominant plant species for evidence of KSHB infestation and evidence of major damage such as limb breakage. In the 22 forested units,I examined the dominant arroyo and black willows (Salix lasiolepis Benth. and S. gooddingii C.R. Ball), and in the seven scrub units, I examined mule fat (Baccharis salicifolia (Ruiz & Pav.) Pers.). Evidence of KSHB infestation was found in 25 of the 29 units. In the forest units, infestation rates ranged from 0 to 100% and were high (>60%) in 16 of the units. In the scrub units, infestation rates ranged from 0 to 33%. Infestation rates were significantly correlated with the wetness of a unit; wetter units had higher infestation rates. Evidence of major physical damage was found in 24 units, and dense stands of willows were reduced to broken trunks in several areas. Overall, I estimated that more than 280,000 (70%) of the willows in the valley were infested, and more than 140,000 had suffered major limb damage. In addition, I recorded evidence of KSHB infestation in the other common plant species in the valley; of the 23 species examined, 14 showed evidence of beetle attack. The four species with the highest rates of infestation were native trees in the Salicaceae family. The three species considered to be the worst invasive plants in the valley, Ricinus communis L., Tamarix ramosissima Ledeb. and Arundo donax L., had low rates of infestation. Several findings from this study have significance for resource managers: (1) the KSHB attack caused extensive mortality of trees soon after being first discovered so, if managers are to control the spread of the beetle, they will need to develop an effective early detection and rapid response program; (2) infestation rates were highest in units that were wet, so resource managers trying to detect the beetle in other areas should thoroughly search trees near water, particularly nutrient-enriched water; (3) the infestation appears to be a novel form of disturbance, and the affected forests may need special management actions in order to recover; and (4) the infestation has altered the structure of the forest canopy, and this is likely to promote the growth of invasive plant species that were relatively inconspicuous in the forests prior to the beetle attack but will now need more attention.

Volatile and Within-Needle Terpene Changes to Douglas-fir Trees Associated With Douglas-fir Beetle (Coleoptera: Curculionidae) Attack

Mass attack by tree-killing bark beetles (Curculionidae: Scolytinae) brings about large chemical changes in host trees that can have important ecological consequences. For example, mountain pine beetle (Dendroctonus ponderosae Hopkins) attack increases emission of terpenes by lodgepole pine (Pinus contorta Dougl. ex Loud.), affecting foliage flammability with consequences for wildfires. In this study, we measured chemical changes to Douglas-fir (Pseudotsuga menziesii var. glauca (Mirb.) Franco) foliage in response to attack by Douglas-fir beetles (Dendroctonus pseudotsugae Hopkins) as trees die and crowns transitioned from green/healthy, to green-infested (year of attack), to yellow (year after attack), and red (2 yr after attack). We found large differences in volatile and within-needle terpene concentrations among crown classes and variation across a growing season. In general, emissions and concentrations of total and individual terpenes were greater for yellow and red needles than green needles. Douglas-fir beetle attack increased emissions and concentrations of terpene compounds linked to increased tree flammability in other conifer species and compounds known to attract beetles (e.g., [Formula: see text]-pinene, camphene, and D-limonene). There was little relationship between air temperature or within-needle concentrations of terpenes and emission of terpenes, suggesting that passive emission of terpenes (e.g., from dead foliage) does not fully explain changes in volatile emissions. The potential physiological causes and ecological consequences of these bark beetle-associated chemical changes are discussed.

Intra-Annual Variation in Responses by Flying Southern Pine Beetles (Coleoptera: Curculionidae: Scolytinae) to Pheromone Component endo-Brevicomin.

The southern pine beetle Dendroctonus frontalis Zimmermann (Coleoptera: Curculionidae: Scolytinae) is attracted to an aggregation pheromone that includes the multifunctional pheromone component endo-brevicomin. The effect of endo-brevicomin on attractive lures varies from strong enhancement to reduction of beetle attraction depending upon release rate, lure component spacing, and proximity of beetle infestations. Anecdotal observations have further suggested that the effects of endo-brevicomin vary during the year. We investigated this possibility under nonoutbreak conditions in southwestern Mississippi where for two-and-a-half years we monitored traps baited with frontalin and the host odor alpha-pinene either (a) alone, or with an endo-brevicomin release device either (b) located directly on the trap, or (c) displaced 6 m away. The endo-brevicomin devices in our tests increased D. frontalis catches during all times of year, and 6 m displacement of the endo-brevicomin release device from the trap did not significantly alter responses except during the spring flight peak when displacement increased catches. Our data suggest that flying D. frontalis have a stronger tendency to avoid the immediate proximity of a release point of endo-brevicomin during their springtime dispersal flight when catches are greatest. Catches of Thanasimus dubius (F.) (Coleoptera: Cleridae), a major predator of D. frontalis, were not altered by endo-brevicomin, and ratios of D. frontalis to T. dubius changed over the course of the year. We discuss the possible effects of intra-annual variation in D. frontalis response to endo-brevicomin both on beetle attack behavior and use of endo-brevicomin as a lure adjuvant in D. frontalis population monitoring.

The Bark-Beetle-Associated Fungus, Endoconidiophora polonica, Utilizes the Phenolic Defense Compounds of Its Host as a Carbon Source.

Norway spruce (Picea abies) is periodically attacked by the bark beetle Ips typographus and its fungal associate, Endoconidiophora polonica, whose infection is thought to be required for successful beetle attack. Norway spruce produces terpenoid resins and phenolics in response to fungal and bark beetle invasion. However, how the fungal associate copes with these chemical defenses is still unclear. In this study, we investigated changes in the phenolic content of Norway spruce bark upon E. polonica infection and the biochemical factors mediating these changes. Although genes encoding the rate-limiting enzymes in Norway spruce stilbene and flavonoid biosynthesis were actively transcribed during fungal infection, there was a significant time-dependent decline of the corresponding metabolites in fungal lesions. In vitro feeding experiments with pure phenolics revealed that E. polonica transforms both stilbenes and flavonoids to muconoid-type ring-cleavage products, which are likely the first steps in the degradation of spruce defenses to substrates that can enter the tricarboxylic acid cycle. Four genes were identified in E. polonica that encode catechol dioxygenases carrying out these reactions. These enzymes catalyze the cleavage of phenolic rings with a vicinal dihydroxyl group to muconoid products accepting a wide range of Norway spruce-produced phenolics as substrates. The expression of these genes and E. polonica utilization of the most abundant spruce phenolics as carbon sources both correlated positively with fungal virulence in several strains. Thus, the pathways for the degradation of phenolic compounds in E. polonica, initiated by catechol dioxygenase action, are important to the infection, growth, and survival of this bark beetle-vectored fungus and may play a major role in the ability of I. typographus to colonize spruce trees.

Mortality, structure, and regeneration in whitebark pine stands impacted by mountain pine beetle in the southern Sierra Nevada

Whitebark pine (Pinus albicaulis Engelm.) is vulnerable to mountain pine beetle (Dendroctonus ponderosae Hopkins) attack throughout western North America, but beetle outbreaks in the southwestern portion of the range (i.e., Sierra Nevada) have been spatially limited until recently. We examined patterns of mortality, structure, and regeneration in whitebark pine stands impacted by mountain pine beetle in the southern Sierra Nevada. Mortality was greatest in medium to large diameter (>10–20 cm dbh) trees, resulting in declines in mean and maximum tree diameter and tree size class diversity following an outbreak. Severity of beetle attack was positively related to mean tree diameter and density. Density of young (<3 years old) whitebark pine seedling clusters was positively related to severity of beetle attack on mature stands. All sites showed a stable production of whitebark pine regeneration within at least the past 30–40 years, with a pulse of new seedlings in the past 3 years in beetle-impacted stands. Our results show that mountain pine beetle outbreaks in the southern Sierra Nevada result in substantial changes in whitebark pine stand structure and suggest low resistance but high resilience to initial attack, especially in the absence of white pine blister rust.

Analisi dell’infestazione di Ips typographus nella foresta dell’Alta Val Parma e strategie per la ricomposizione del bosco

Abstract: Analysis of the bark beetle outbreak in the forest “Alta Val Parma” (Corniglio, Parma, Italy) and strategies for its regeneration. Norway spruce plantations located in the Foresta Demaniale Alta Val Parma (Corniglio, province of Parma - Italy) experienced since 2004 a massive outbreak of Norway spruce bark beetle (Ips typographus). This outbreak has been triggered by the exceptionally warm and dry summer of 2003. In the following years bark beetle attacks repeated and spread, raising concern about the future of this kind of stands. A survey program has been carried out to help local administration to chose the correct managements strategies. Monitoring of Ips typographus population, carried out between 2007 and 2013, confirmed the presence of two generations per year, with values above the risk threshold in 2007 and just below though very high in 2011. In the affected area, six survey plots have been realized to test different management options with the aim of favoring a fast regeneration of the forest cover. The tested options showed the great difficulty in the establishment of natural generation either for the lack of mother plants in such pure stands or for competition with tall grasses. Sowing brought no significative results, while direct plantation of indigenous broadleaves was more effective, with almost half of the individuals still alive after three years. Our results confirm the great difficulty to rebuild the forest cover after strong ecological disturbances in these artificial forests. Hybrid management strategies and ad hoc silvicultural choices seem to be the only way to manage such kind of situations in a National park, where the priority is biodiversity conservation.

Effect of bark beetle (Ips typographus L.) attack on bark VOC emissions of Norway spruce (Picea abies Karst.) trees

Abstract Climate warming driven storms are evident causes for an outbreak of the European spruce bark beetle (Ips typographus L.) resulting in the serious destruction of mature Norway spruce (Picea abies Karst.) forests in northern Europe. Conifer species are major sources of biogenic volatile organic compounds (BVOCs) in the boreal zone. Climate relevant BVOC emissions are expected to increase when conifer trees defend against bark beetle attack by monoterpene (MT)-rich resin flow. In this study, BVOC emission rates from the bark surface of beetle-attacked and non-attacked spruce trees were measured from two outbreak areas, Iitti and Lahti in southern Finland, and from one control site at Kuopio in central Finland. Beetle attack increased emissions of total MTs 20-fold at Iitti compared to Kuopio, but decreased the emissions of several sesquiterpenes (SQTs) at Iitti. At the Lahti site, the emission rate of α-pinene was positively correlated with mean trap catch of bark beetles. The responsive individual MTs were tricyclene, α-pinene, camphene, myrcene, limonene, 1,8-cineole and bornyl acetate in both of the outbreak areas. Our results suggest that bark beetle outbreaks affect local BVOC emissions from conifer forests dominated by Norway spruce. Therefore, the impacts of insect outbreaks are worth of consideration to global BVOC emission models.

Prescribed fire does not promote outbreaks of a primary bark beetle at low‐density populations

Summary The causes of bark beetle outbreaks – particularly the role of disturbances – are poorly understood. Stand‐scale disturbances, like fires, can suddenly improve local host susceptibility and may attract beetles; however, whether such increases can lead to outbreaks in post‐disturbance stands is unclear. Using low‐density Dendroctonus ponderosae mountain pine beetle populations in Pinus contorta lodgepole pine forests in western Canada, we investigated whether prescribed fires promote outbreaks or provide only short‐term resources. Proportionally more burned than non‐burned trees were attacked. At one site, beetle attacks increased in response to a resource pulse, but the proportions of attacked trees and numbers of attacks per tree declined over four years after fire. Elsewhere, beetle attacks remained very low. As the resource (phloem) quality of burned trees remained high three years after fire, we propose that post‐fire mortality, resulting in fewer available host trees, can at least partially explain why D. ponderosae did not build up populations in burned stands. Synthesis and applications. Our study emphasizes the importance of examining long‐term trends in fire–bark beetle interactions, and of understanding low‐density beetle populations. Because fire does not seem to promote mountain pine beetle outbreaks, we recommend the continued use of prescribed fire for the general management of P. contorta forests with low‐density beetle populations.

Ponderosa pine resin defenses and growth: metrics matter.

Bark beetles (Coleoptera: Curculionidae, Scolytinae) cause widespread tree mortality in coniferous forests worldwide. Constitutive and induced host defenses are important factors in an individual tree's ability to survive an attack and in bottom-up regulation of bark beetle population dynamics, yet quantifying defense levels is often difficult. For example, in Pinus spp., resin flow is important for resistance to bark beetles but is extremely variable among individuals and within a season. While resin is produced and stored in resin ducts, the specific resin duct metrics that best correlate with resin flow remain unclear. The ability and timing of some pine species to produce induced resin is also not well understood. We investigated (i) the relationships between ponderosa pine (Pinus ponderosa Lawson & C. Lawson) resin flow and axial resin duct characteristics, tree growth and physiological variables, and (ii) if mechanical wounding induces ponderosa pine resin flow and resin ducts in the absence of bark beetles. Resin flow increased later in the growing season under moderate water stress and was highest in faster growing trees. The best predictors of resin flow were nonstandardized measures of resin ducts, resin duct size and total resin duct area, both of which increased with tree growth. However, while faster growing trees tended to produce more resin, models of resin flow using only tree growth were not statistically significant. Further, the standardized measures of resin ducts, density and duct area relative to xylem area, decreased with tree growth rate, indicating that slower growing trees invested more in resin duct defenses per unit area of radial growth, despite a tendency to produce less resin overall. We also found that mechanical wounding induced ponderosa pine defenses, but this response was slow. Resin flow increased after 28 days, and resin duct production did not increase until the following year. These slow induced responses may allow unsuccessfully attacked or wounded trees to resist future bark beetle attacks. Forest management that encourages healthy, vigorously growing trees will also favor larger resin ducts, thereby conferring increased constitutive resistance to bark beetle attacks.

Arboreal Squirrel Abundance in Response to a Gradient of Mountain Pine Beetle Attack in Sub-boreal Forests

To assist in evaluating habitat retention options, the abundance of northern flying squirrels (Glaucomys sabrinus) and North American red squirrels (Tamiasciurus hudsonicus) wer examined in 2005 and again in 2010 across a gradient of mountain pine beetle (Dendroctonus ponderosae) attack intensity in sub-boreal forests of west-central British Columbia. Among 30, 16-ha live-trapping grids, estimated mean abundance of both species increased non-linearly with remaining basal area of live overstorey (live trees ≥ 7.5 cm diameter at 1.3 m height). A weak (most likely positive but possibly negative) additional response of flying squirrels to dead overstorey (on average 1 m2/ha dead ≈ +0.11 m2/ha live basal area) was evident. The basal area of live spruce-fir (Picea spp., Abies spp.) overstorey, understorey tree density, and tree diameter covariates did not have substantive additional effects on estimated squirrel abundance. Whereas survey year affected overall abundance, it did not change the relationship with habitat attributes. The results suggest that dead lodgepole pine (Pinus contorta) has low habitat value for arboreal squirrels and thus is of lower impact on squirrels if salvage-harvested; however, retention or re-growth of sufficient live overstorey is necessary to maintain or recover squirrel abundance. The basal area oflive overstorey appears a simple yet useful management metric for prioritizing habitatvalue, at least as indicated by relative abundance for squirrels in beetle-affected forests.

Limber pine in the central and southern Rocky Mountains: Stand conditions and interactions with blister rust, mistletoe, and bark beetles

Combined effects of the recent mountain pine beetle (Dendroctonus ponderosae) outbreak and white pine blister rust (caused by Cronartium ribicola) (WPBR) are causing extensive crown dieback and mortality in limber pine (Pinus flexilis) in the central and southern Rocky Mountains. Additional stressors such as climate change and limber pine dwarf mistletoe (Aceuthobium cyanocarpum) may significantly alter stand structure and biodiversity of these ecosystems. The objectives of this study were to: (1) assess site, stand, and tree health conditions in limber pine stands of northern Colorado, Wyoming, and southeastern Montana, (2) quantify impacts of WPBR, bark beetles, and dwarf mistletoe, and (3) determine biotic and abiotic factors that influence the occurrence and incidence of these damage agents.We assessed 22,700 limber pines on 508 plots in limber pine-dominated stands in twenty-five study areas in northern Colorado, Wyoming, and southeastern Montana. Average live limber pine density was 311stemsha−1. Overall, 50% of limber pines were classified as healthy with over 50% of limber pine mortality attributed to bark beetle attacks. White pine blister rust damage was evident on over 60% of declining or dying limber pines. Blister rust was the primary damage agent, occurring on 73% of the plots and 26% of the trees. Bark beetle-caused mortality was found on 75% of plots and 18% of trees. Limber pine dwarf mistletoe was present on 29% of plots and 9% of trees. In study areas that were previously monitored, incidence of WPBR increased over 8–9years by 6%, bark beetle-caused mortality by 17%, while dwarf mistletoe incidence remained the same. We used statistical modeling to determine meteorological, macro and micro site factors, and stand factors that influenced the occurrence and incidence of WPBR, bark beetles, and dwarf mistletoe on limber pines.Declines in health and rise in mortality due to bark beetles and WPBR has left some study areas with low limber pine basal area and stand density. These stands may be at risk for localized extirpation if the minimum viable population thresholds are not maintained and are identified as areas of concern for restoration and conservation efforts. Current condition status and long-term monitoring of limber pine is needed for land manager decision making and facilitation of restoration goals.

Influence of selected factors on bark beetle outbreak dynamics in the Western Carpathians

Abstract In the period from 1992 to 2013, more than 3.8 million m3 of spruce wood from an area of 55 thousand ha of forests in the Kysuce region (Western Carpathians) was affected by bark beetles. This region has had the highest volume of salvage fellings in Slovakia. While before 1991, bark-beetle outbreak usually occurred after snow and wind disturbances, since 1992 they have occurred in the years with extremely warm and dry growing seasons and the years following them. These years were also characterised by high volumes of wood affected by honey fungus (Armillaria spp.), which only rarely took part in the calamities before 1992. Extreme deterioration of the situation occurred after 2003. In fragmented and sparse stands, the volume of wood damaged by wind increased. Artificial origin of spruce stands, their high occurrence, high age and even-agedness are likely pre-disposing factors of spruce forest decline. Bark beetles have become the most important factor of spruce decline. The most important factors driving the bark beetle attack on forest stands in the period 1973–2013 were the amount of unprocessed wood in the previous year; the amount of wood affected by honey fungus, precipitation total, and average temperature in the current growing season. Another important factor that complicated the situation was also the inferior quality of forest management.

Reduced growth sensitivity to climate in bark-beetle infested Aleppo pines: Connecting climatic and biotic drivers of forest dieback

Drought and bark-beetle infestation are major and often interconnected drivers of forest dieback and tree death. These two stressors may interact and accelerate forest mortality, since warmer and drier conditions boost beetle attacks and reduce tree growth. However, the way in which drought and bark-beetle infestation interact and affect declining or dying trees is still poorly understood. To disentangle the long-term interaction between the two stressors, we quantified radial growth (basal area increment), resin production, mortality and intrinsic water-use efficiency (iWUE) inferred from wood-carbon isotope discrimination. We compared trees infested and recently killed by two abundant bark beetle species (Orthotomicus erosus and Tomicus piniperda) with non-infested living trees in a drought-prone Aleppo pine plantation. Growth and iWUE showed similar values in infested and non-infested trees. Since bark-beetle-infested trees did not grow less than non-infested trees, our results did not support the hypothesis of higher costs of resin production at the expense of stem-wood formation. Radial growth was enhanced by cool and wet winter conditions prior to the growing season. However, infested trees showed lower growth responsiveness than did non-infested trees with respect to this climatic driver of growth. Infested trees also showed a lower resin-duct production two years prior to death than non-infested trees. The growth responsiveness to climate should be characterized in bark-beetle-infested trees, since a weak correlation between climate and growth can be regarded as a predisposing factor of infestation-induced tree death. Such reduced responsiveness to climate stress could be linked to the tree vulnerability to beetle attacks in drought-prone forests.

Nonstructural carbohydrate dynamics of lodgepole pine dying from mountain pine beetle attack.

Bark beetle outbreaks are an important cause of tree death, but the process by which trees die remains poorly understood. The effect of beetle attack on whole-tree nonstructural carbohydrate (NSC) dynamics is particularly unclear, despite the potential role of carbohydrates in plant defense and survival. We monitored NSC dynamics of all organs in attacked and protected lodgepole pines (Pinus contorta) during a mountain pine beetle (Dendroctonus ponderosae) outbreak in British Columbia, starting before beetle flight in June 2011 through October 2012, when most attacked trees had died. Following attack, NSC concentrations were first reduced in the attacked region of the bole. The first NSC reduction in a distant organ appeared in the needles at the end of 2011, while branch and root NSC did not decline until much later in 2012. Attacked trees that were still alive in October 2012 had less beetle damage, which was negatively correlated with initial bark sugar concentrations in the attack region. The NSC dynamics of dying trees indicate that trees were killed by a loss of water conduction and not girdling. Further, our results identify locally reduced carbohydrate availability as an important mechanism by which stressors like drought may increase tree susceptibility to biotic attack.

Tree response and mountain pine beetle attack preference, reproduction and emergence timing in mixed whitebark and lodgepole pine stands

AbstractMountain pine beetle (Dendroctonus ponderosae) is an important disturbance agent in Pinus ecosystems of western North America, historically causing significant tree mortality. Most recorded outbreaks have occurred in mid elevation lodgepole pine (Pinus contorta). In warm years, tree mortality also occurs at higher elevations in mixed species stands. Mountain pine beetle's relative preference for and performance in Pinus species that either commonly or less frequently encounter this insect has received little direct testing. Further, knowledge of the relative proportions of secondary compounds, which can differ among Pinus species and play important roles in attack rates and outcomes, is important to understanding host suitability. We monitored mountain pine beetle attacks, adult emergence timing and reproductive capacity in lodgepole and whitebark (Pinus albicaulis) pines growing in mixed stands at relatively high elevation. Phloem monoterpene chemistry of trees prior to and during attack was compared within and between species. Although beetles attacked lodgepole pine more frequently, lodgepole pines also resisted attacks more frequently. Overall, there were equal numbers of lethal attacks between species. Brood production and adult emergence timing did not differ between tree species. The relative composition of secondary compounds differed by tree species, although both species contained compounds that affect mountain pine beetle attack and reproductive success.

Pollinators, pests and soil properties interactively shape oilseed rape yield

Pollination, pest control, and soil properties are well known to affect agricultural production. These factors might interactively shape crop yield, but most studies focus on only one of these factors at a time. We used 15 winter oilseed rape (Brassica napus L.) fields in Sweden to study how variation among fields in pollinator visitation rates, pollen beetle attack rates and soil properties (soil texture, pH and organic carbon) interactively determined crop yield. The fields were embedded in a landscape gradient with contrasting proportions of arable and semi-natural land. In general, pollinator visitation and pest levels were negatively correlated and varied independently of soil properties. This may reflect that above- and below-ground processes react at landscape and local spatial scales, respectively. The above-ground biotic interactions and below-ground abiotic factors interactively affected crop yield. Pollinator visitation was the strongest predictor positively associated with yield. High soil pH also benefited yield, but only at lower pest loads. Surprisingly, high pest loads increased the pollinator benefits for yield. Implementing management plans at different spatial scales can create synergies among above- and below-ground ecosystem processes, but both scales are needed given that different processes react at different spatial scales.