Influences of the biophysical environment on blister rust and mountain pine beetle,and their interactions,in whitebark pine forests

Aim To understand how the biophysical environment influences patterns of infection by non‐native blister rust (caused by Cronartium ribicola) and mortality caused by native mountain pine beetles (Dendroctonus ponderosae) in whitebark pine (Pinus albicaulis) communities, to determine how these disturbances interact, and to gain insight into how climate change may influence these patterns in the future. Location High‐elevation forests in south‐west Montana, central Idaho, eastern and western Oregon, USA. Methods Stand inventory and dendroecological methods were used to assess stand structure and composition and to reconstruct forest history at sixty 0.1‐ha plots. Patterns of blister rust infection and mountain pine beetle‐caused mortality in whitebark pine trees were examined using nonparametric Kruskal–Wallis ANOVA, Mann–Whitney U‐tests, and Kolmogorov–Smirnov two‐sample tests. Stepwise regression was used to build models of blister rust infection and mountain pine beetle‐related mortality rates based on a suite of biophysical site variables. Results Occurrence of blister rust infections was significantly different among the mountain ranges, with a general gradient of decreasing blister rust occurrence from east to west. Evidence of mountain pine beetle‐caused mortality was identified on 83% of all dead whitebark pine trees and was relatively homogenous across the study area. Blister rust infected trees of all ages and sizes uniformly, while mountain pine beetles infested older, larger trees at all sites. Stepwise regressions explained 64% and 58% of the variance in blister rust infection and beetle‐caused mortality, respectively, indicating that these processes are strongly influenced by the biophysical environment. More open stand structures produced by beetle outbreaks may increase the exposure of surviving whitebark pine trees to blister rust infection. Main conclusions Variability in the patterns of blister rust infection and mountain pine beetle‐caused mortality elucidated the fundamental dynamics of these disturbance agents and suggests that the effects of climate change will be complex in whitebark pine communities and vary across the species’ range. Interactions between blister rust and beetle outbreaks may accelerate declines or facilitate the rise of rust resistance in whitebark pine depending on forest conditions at the time of the outbreak.     

A test of high-dose verbenone for stand-level protection of lodgepole and whitebark pine from mountain pine beetle (Coleoptera: Curculionidae: Scolytinae) attacks

The efficacy of verbenone as a stand-level protectant against mountain pine beetle, Dendroctonus ponderosae Hopkins, attacks was tested in lodgepole and whitebark pine stands at five geographically separated sites, including three consecutive years at one site. Forty and 20 high-dose pouches, with a verbenone emission rate up to 50 mg/d per pouch, were spaced in a grid pattern throughout 0.40-ha plots, replicated up to six times at each site. Although the verbenone treatment did not prevent beetles from dispersing through treated stands, attacking large-diameter trees most frequently, the overall number of trees attacked was, on average, reduced significantly compared with nontreated stands. In a few blocks each year, verbenone-treated plots had more attacked trees than controls. These blocks tended to have a large emerging beetle population, exceeding 140 previously attacked trees within the hectare including and surrounding the treated area. Additional research is needed on the behavioral role of verbenone in mountain pine beetle population dynamics and quantification of the infestation level above which treatment efficacy tends to be reduced.     

Verbenone-releasing flakes protect individual Pinus contorta trees from attack by Dendroctonus ponderosae and Dendroctonus valens (Coleoptera: Curculionidae, Scolytinae)

1 In a study site in interior northern California, twenty individual lodgepole pines Pinus contorta were sprayed with a suspension of DISRUPT Micro‐Flake^® Verbenone (4,6,6‐trimethylbicyclo(3.1)hept‐3‐en‐2‐one) Bark Beetle Anti‐Aggregant flakes (Hercon Environmental, Emigsville, Pennsylvania) in water, with sticker and thickener, from ground level to a height of 7 m. Twenty trees sprayed with just water, sticker and thickener served as controls. All trees were baited immediately after spraying with mountain pine beetle Dendroctonus ponderosae aggregation pheromone lures, and lures were refreshed after 4 weeks. 2 Trees treated with verbenone had significantly lower attack density by D. ponderosae than controls at 2, 4, 6 and 8 weeks after application of flakes. 3 None of the treated trees was attacked by red turpentine beetle Dendroctonus valens, whereas control trees averaged nearly two D. valens attacks per tree, 8 weeks after treatment. 4 A dry frass index, used to predict ultimate tree mortality, was significantly higher in control trees than treated trees for all four sampling intervals. This index proved to be a significant predictor of ultimate tree mortality. 5 Ten months after application, treated trees showed significantly lower mortality than control trees.     

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

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Glyphosate-induced changes in the attack success and development of the mountain pine beetle and impact of its natural enemies

Roundup (active ingredient glyphosate) administered into the sapwood around the root collar of lodgepole pine trees, Pinus contorta var. latifolia Engelm., was investigated as a tool for creating trap trees for the mountain pine beetle (MPB), Dendroctonus ponderosae Hopk. MPB's attacked semiochemical-baited, glyphosate-treat trees before attacking baited control trees. Bark samples disclosed an increase in survival of MPB eggs and early instar larvae, enhanced larval development and increased attacks by insect parasites and predators in treated trees. The braconid wasp, Coeloides dendroctoni Cush., parasitized MPB larvae found at high densities in treated trees two months following attack. The dipteran predator, Medetera aldrichii Wh., was only found in treated trees. Increased competition by Ips pini (Say) had a negative impact on MPB survivorship. The enhanced impact of insectan agents apparently offset any glyphosate-induced gains in MPB survival or development, but did not cause a significant reduction in MPB emergence.     

Stand‐replacing fires reduce susceptibility of lodgepole pine to mountain pine beetle outbreaks in Colorado

Aim As climate change is increasing the frequency, severity and extent of wildfire and bark beetle outbreaks, it is important to understand how these disturbances interact to affect ecological patterns and processes, including susceptibility to subsequent disturbances. Stand‐replacing fires and outbreaks of mountain pine beetle (MPB), Dendroctonus ponderosae, are both important disturbances in the lodgepole pine, Pinus contorta, forests of the Rocky Mountains. In the current study we investigated how time since the last stand‐replacing fire affects the susceptibility of the stand to MPB outbreaks in these forests. We hypothesized that at a stand‐scale, young post‐fire stands (< c. 100–150 years old) are less susceptible to past and current MPB outbreaks than are older stands. Location Colorado, USA. Methods We used dendroecological methods to reconstruct stand‐origin dates and the history of outbreaks in 23 lodgepole pine stands. Results The relatively narrow range of establishment dates among the oldest trees in most sampled stands suggested that these stands originated after stand‐replacing or partially stand‐replacing fires over the past three centuries. Stands were affected by MPB outbreaks in the 1940s/1950s, 1980s and 2000s/2010s. Susceptibility to outbreaks generally increased with stand age (i.e. time since the last stand‐replacing fire). However, this reduced susceptibility of younger post‐fire stands was most pronounced for the 1940s/1950s outbreak, less so for the 1980s outbreak, and did not hold true for the 2000s/2010s outbreak. Main conclusions Younger stands may not have been less susceptible to the most recent outbreak because: (1) after stands reach a threshold age of > 100–150 years, stand age does not affect susceptibility to outbreaks, or (2) the high intensity of the most recent outbreak reduces the importance of pre‐disturbance conditions for susceptibility to disturbance. If the warm and dry conditions that contribute to MPB outbreaks concurrently increase the frequency and/or extent of severe fires, they may thereby mitigate the otherwise increased landscape‐scale susceptibility to outbreaks. Potential increases in severe fires driven by warm and dry climatic trends may lead to a negative feedback by making lodgepole pine stands less susceptible to future MPB outbreaks.     

Identification of further sex pheromone synergists in the codling moth, Cydia pomonella

A number of angiosperm nonhost volatiles (NHVs) and green leaf volatiles (GLVs) were tested alone and as supplements to the antiaggregation pheromone, verbenone, for their ability to disrupt attack by the mountain pine beetle (MPB), Dendroctonus ponderosae Hopkins (Coleoptera: Scolytidae), on lodgepole pine, Pinus contorta Dougl. ex Loud. var. latifolia Engel. Preliminary experiments led to a refined NHV blend [benzyl alcohol, guaiacol, benzaldehyde, nonanal, salicylaldehyde, and conophthorin] and a refined GLV blend [(Z)-3-hexen-1-ol and (E)-2-hexen-1-ol]. In a 20-replicate experiment, NHV and GLV groups both singly, and verbenone alone, significantly reduced MPB mass attack on pheromone-baited trees and on trees within 5 m of the pheromone-baited trees. Both blends in combination with verbenone reduced the number of mass attacked, baited trees to three out of 20 compared to 20 out of 20 of the baited controls. Each binary combination was also effective at reducing mass attack. In these experiments, all tested repellents were released from devices stapled to trees at the same point as the pheromone bait, suggesting that the repellency could have been to a point source, rather than to the whole tree. Therefore, in two further experiments bands of release devices were wrapped around the treated trees and the pheromone bait was removed from the treated trees. In one experiment, when the aggregation pheromone bait was suspended between pairs of trees treated with the NHV blend plus GLV blend plus verbenone, only three out of 25 treated pairs had mass attack on at least one member of the pair. In the other 60-replicate experiment, with no pheromone baits present, attack occurred on 13 untreated and 11 banded trees, all in the path of a large advancing infestation. However, the mean attack density on the banded trees was significantly reduced to a level below the 40 attacks m^–2 of bark surface required to kill a healthy lodgepole pine. As a result of these experiments, operational trials are recommended.     

Predators and competitors of the mountain pine beetle Dendroctonus ponderosae (Coleoptera: Curculionidae) in stands of changing forest composition associated with elevation

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Performance of the tree‐killing bark beetles Ips typographus and Pityogenes chalcographus in non‐indigenous lodgepole pine and their historical host Norway spruce

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Attack dynamics of the pine shoot beetle, Tomicus piniperda (Col.; Scolytinae) in Scots pine stands defoliated by Bupalus piniaria (Lep.; Geometridae)

Abstract 1 After a 1‐year, extensive pine looper (Bupalus piniaria) outbreak, plots were laid out to study tree susceptibility to attack, and performance of Tomicus piniperda in pine trees suffering from varying levels of defoliation. 2 Tomicus piniperda was the dominating stem‐attacking species among the primary stem colonizers, and 82% of all trees that died had been colonized by T. piniperda. 3 Beetle attacks primarily struck severely defoliated trees, i.e. trees that suffered from 90% to 100% defoliation. 4 Beetle attacks peaked in the second year after cessation of the outbreak, and suppressed trees were both more frequently attacked and more susceptible to beetle attack than intermediate and dominant trees. 5 Trees surviving beetle attacks carried more foliage than trees that did not survive the attacks. 6 A single year of severe defoliation is enough to render pine trees susceptible to secondary pests, such as T. piniperda.     

Nitrogen Cycling Responses to Mountain Pine Beetle Disturbance in a High Elevation Whitebark Pine Ecosystem

Ecological disturbances can significantly affect biogeochemical cycles in terrestrial ecosystems, but the biogeochemical consequences of the extensive mountain pine beetle outbreak in high elevation whitebark pine (WbP) (Pinus albicaulis) ecosystems of western North America have not been previously investigated. Mountain pine beetle attack has driven widespread WbP mortality, which could drive shifts in both the pools and fluxes of nitrogen (N) within these ecosystems. Because N availability can limit forest regrowth, understanding how beetle-induced mortality affects N cycling in WbP stands may be critical to understanding the trajectory of ecosystem recovery. Thus, we measured above- and belowground N pools and fluxes for trees representing three different times since beetle attack, including unattacked trees. Litterfall N inputs were more than ten times higher under recently attacked trees compared to unattacked trees. Soil inorganic N concentrations also increased following beetle attack, potentially driven by a more than two-fold increase in ammonium (NH₄ ⁺) concentrations in the surface soil organic horizon. However, there were no significant differences in mineral soil inorganic N or soil microbial biomass N concentrations between attacked and unattacked trees, implying that short-term changes in N cycling in response to the initial stages of WbP attack were restricted to the organic horizon. Our results suggest that while mountain pine beetle attack drives a pulse of N from the canopy to the forest floor, changes in litterfall quality and quantity do not have profound effects on soil biogeochemical cycling, at least in the short-term. However, continuous observation of these important ecosystems will be crucial to determining the long-term biogeochemical effects of mountain pine beetle outbreaks.     

Mountain pine beetle selectivity in old‐growth ponderosa pine forests,Montana, USA

A historically unprecedented mountain pine beetle (MPB) outbreak affected western Montana during the past decade. We examined radial growth rates (AD 1860–2007/8) of co‐occurring mature healthy and MPB‐infected ponderosa pine trees collected at two sites (Cabin Gulch and Kitchen Gulch) in western Montana and: (1) compared basal area increment (BAI) values within populations and between sites; (2) used carbon isotope analysis to calculate intrinsic water‐use efficiency (iWUE) at Cabin Gulch; and (3) compared climate‐growth responses using a suite of monthly climatic variables. BAI values within populations and between sites were similar until the last 20–30 years, at which point the visually healthy populations had consistently higher BAI values (22–34%) than the MPB‐infected trees. These results suggest that growth rates two–three decades prior to the current outbreak diverged between our selected populations, with the slower‐growing trees being more vulnerable to beetle infestation. Both samples from Cabin Gulch experienced upward trends in iWUE, with significant regime shifts toward higher iWUE beginning in 1955–59 for the visually healthy trees and 1960–64 for the MPB‐infected trees. Drought tolerance also varied between the two populations with the visually healthy trees having higher growth rates than MPB‐infected trees prior to infection during a multi‐decadal period of drying summertime conditions. Intrinsic water‐use efficiency significantly increased for both populations during the past 150 years, but there were no significant differences between the visually healthy and MPB‐infected chronologies.     

Tree‐mediated interactions between the jack pine budworm and a mountain pine beetle fungal associate

1. Coniferous trees deploy a combination of constitutive (pre‐existing) and induced (post‐invasion), structural and biochemical defences against invaders. Induced responses can also alter host suitability for other organisms sharing the same host, which may result in indirect, plant‐mediated interactions between different species of attacking organisms. 2. Current range and host expansion of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB) from lodgepole pine‐dominated forests to the jack pine‐dominated boreal forests provides a unique opportunity to investigate whether the colonisation of jack pine (Pinus banksiana Lamb.) by MPB will be affected by induced responses of jack pine to a native herbaceous insect species: the jack pine budworm (Choristoneura pinus pinus Freeman; JPBW). 3. We simulated MPB attacks with one of its fungal associates, Grosmannia clavigera Robinson‐Jeffrey & Davidson, and tested induction of either herbivory by JPBW or inoculation with the fungus followed by a challenge treatment with the other organism on jack pine seedlings and measured and compared monoterpene responses in needles. 4. There was clear evidence of an increase in jack pine resistance to G. clavigera with previous herbivory, indicated by smaller lesions in response to fungal inoculations. In contrast, although needle monoterpenes greatly increased after G. clavigera inoculation and continued to increase during the herbivory challenge, JPBW growth was not affected, but JPBW increased the feeding rate to possibly compensate for altered host quality. 5. Jack pine responses varied greatly and depended on whether seedlings were treated with single or multiple organisms, and their order of damage.     

The effect of stem density on the probability of attack by the ambrosia beetle Platypus quercivorus varies with spatial scale

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Mountain pine beetle host-range expansion threatens the boreal forest

The current epidemic of the mountain pine beetle (MPB), an indigenous pest of western North American pine, has resulted in significant losses of lodgepole pine. The leading edge has reached Alberta where forest composition shifts from lodgepole to jack pine through a hybrid zone. The susceptibility of jack pine to MPB is a major concern, but there has been no evidence of host-range expansion, in part due to the difficulty in distinguishing the parentals and their hybrids. We tested the utility of a panel of microsatellite loci optimized for both species to classify lodgepole pine, jack pine and their hybrids using simulated data. We were able to accurately classify simulated individuals, and hence applied these markers to identify the ancestry of attacked trees. Here we show for the first time successful MPB attack in natural jack pine stands at the leading edge of the epidemic. This once unsuitable habitat is now a novel environment for MPB to exploit, a potential risk which could be exacerbated by further climate change. The consequences of host-range expansion for the vast boreal ecosystem could be significant.     

Facilitation in bark beetles: endemic mountain pine beetle gets a helping hand

• 1 Endemic populations of the bark beetle Dendroctonus ponderosae attack weakened lodgepole pine (Pinus contorta var. latifolia) trees that are often previously infested by other bark beetle species, such as Pseudips mexicanus.
• 2 The effect of interactions on D. ponderosae was assessed by examining host selection and productivity of D. ponderosae in trees containing P. mexicanus and trees infested solely by D. ponderosae.
• 3 The findings obtained show that D. ponderosae attacked hosts previously occupied by P. mexicanus at greater densities, and offspring emerged earlier compared with hosts infested by D. ponderosae alone. Additionally, D. ponderosae larvae in P. mexicanus‐infested trees were found to require a significantly lower amount of resource to complete development with no loss in size.
• 4 The presence of P. mexicanus may affect host condition, improving the subcortical environment for endemic D. ponderosae, ultimately aiding in population maintenance at low levels. Hosts in this state should be preferentially attacked by D. ponderosae.

     

What explains landscape patterns of tree mortality caused by bark beetle outbreaks in Greater Yellowstone?

Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.     

Thinning Jeffrey pine stands to reduce susceptibility to bark beetle infestations in California,U.S.A.

• 1 Bark beetles (Coleoptera: Curculionidae, Scolytinae) are commonly recognized as important tree mortality agents in coniferous forests of the western U.S.A.
• 2 High stand density is consistently associated with bark beetle infestations in western coniferous forests, and therefore thinning has long been advocated as a preventive measure to alleviate or reduce the amount of bark beetle‐caused tree mortality.
• 3 The present study aimed to determine the effectiveness of thinning to reduce stand susceptibility to bark beetle infestations over a 10‐year period in Pinus jeffreyi forests on the Tahoe National Forest, California, U.S.A. Four treatments were replicated three times within 1‐ha square experimental plots. Treatments included thinning from below (i.e. initiating in the smallest diameter classes) to a residual target basal area (cross‐sectional area of trees at 1.37 m in height) of: (i) 18.4 m²/ha (low density thin); (ii) 27.6 m²/ha (medium density thin); (iii) 41.3 m²/ha (high density thin); and (iv) no stand manipulation (untreated control).
• 4 Throughout the present study, 107 trees died as a result of bark beetle attacks. Of these, 71% (75 trees) were Abies concolor killed by Scolytus ventralis; 20.6% (22 trees) were Pinus ponderosa killed by Dendroctonus ponderosae; 4.7% (five trees) were P. jeffreyi killed by Dendroctonus jeffreyi; 1.8% (two trees) were P. jeffreyi killed by Ips pini; 0.9% (one tree) were P. jeffreyi killed by Orthotomicus (= Ips) latidens; 0.9% (one tree) were P. ponderosa killed by both Dendroctonus brevicomis and D. ponderosae; and 0.9% (one tree) were P. jeffreyi killed by unknown causes.
• 5 In the low density thin, no pines were killed by bark beetles during the 10‐year period. Significantly fewer trees (per ha/year) were killed in the low density thin than the high density thin or untreated control. No significant treatment effect was observed for the percentage of trees (per year) killed by bark beetles.

     

Mass‐trapping trials for the control of pine processionary moth in a pine woodland recreational area

The pine processionary moth, Thaumetopoea pityocampa, causes serious defoliation to Cedrus, Pinus and Pseudotsuga trees, as well as health problems in humans, pets and farm animals due to their urticating hairs. Environmentally friendly strategies for the management of T. pityocampa include: removal of egg batches, removal of nests, trapping of migrant larvae, spraying microbial or Insect Growth Regulator (IGR) insecticides and biocontrol, as well as pheromone‐based adult trapping and mating‐disruption. In the present paper, results on innovative technology for the control of T. pityocampa infestation using pheromone mass‐trapping are reported. Two 1‐ha plots were identified in the study area (central‐south Italy), a pine woodland recreational site growing Pinus halepensis. In the experimental plot (MT‐plot), 10 G‐traps (funnel trap type) baited with (Z)‐13‐hexadecen‐11‐ynyl acetate sex pheromone component were placed for mass‐trapping of adults; the other plot was used as a control‐plot (C‐plot). The T. pityocampa population was monitored using the two central traps in the MT‐plot and two traps positioned in the C‐plot. In addition, the winter nests made by T. pityocampa larvae overwintering on pine trees were counted. After 2 years of mass‐trapping, the number of adults trapped by the monitoring pheromone traps decreased in the MT‐plot, but not in the C‐plot, whereas the number of nests decreased in both plots. Statistical results highlighted significant differences in trap catches between the two plots but not between years. In the case of nests, differences among plots were not significant before the mass‐trapping, but significant after 1‐year treatment. According to our results, the mass‐trapping technique is able to reduce T. pityocampa infestations. This pheromone method can be applied in combination with other control systems in the context of integrated pest management in recreational areas.