Drought and stand susceptibility to attacks by the European spruce bark beetle: A remote sensing approach

1. Several time-series analyses have demonstrated that after extreme summer drought bark beetle damage increased. However, studies predicting stand susceptibility over large spatial extents are limited by technical constraints in obtaining detailed, spatially-explicit data on infestation spot occurrence.
2. Using a unique dataset of georeferenced bark beetle infestation data, we tested whether the spatial variation of local growing conditions of forest stands, topography, and landscape variables modified the local occurrence of Ips typographus infestations after a severe hot drought in Central Europe.
3. Bark beetle infestation occurrence depended on soil-related aridity intensity, elevation, slope, and soil conditions. We showed that elevation interacted with growing conditions and topography. At low elevations, spruce forests growing on flat areas and wetter soils were more sensitive to the infestations. On the contrary, forests on steep slopes and soils with low water availability were rarely attacked. At the landscape scale, bark beetle damage increased with host tree cover but decreased with compositional diversity.
4. Our findings are generally consistent with the growth-differentiation balance hypothesis that predicts that trees growing under chronic dry conditions tend to be more resistant against biotic disturbances.
5. Spruce stands at low elevations located in homogeneous landscapes dominated by spruce were those more exposed to bark beetles in the initial phase of a drought-induced outbreak.

     

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.

     

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

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Evaluation of traps used to monitor southern pine beetle aerial populations and sex ratios

1 Various kinds of traps have been employed to monitor and forecast population trends of the southern pine beetle (Dendroctonus frontalis Zimmermann; Coleoptera: Scolytidae), but their accuracy in assessing pine‐beetle abundance and sex ratio in the field has not been evaluated directly. 2 In this study, we used fluorescent powder to mark pine beetles emerging from six isolated infestations. We then compared estimates of total abundances and proportions of males emerging from within each infestation to the estimates from three types of traps: passive sticky traps (2, 5, 10 and 20 m away from the source of beetles), multi‐funnel traps baited with pine beetle attractants (100 m away) and pine trees baited with attractants (also 100 m away). 3 We found that the proportion of males captured in traps was significantly affected by the type of trap used. 4 Within an infestation, equal proportions of males and females were marked (0.53 ± 0.02 males; mean ± SE), but the proportions captured in trap trees and passive traps were more female biased (0.42 ± 0.03 and 0.46 ± 0.01 males, respectively). On the other hand, funnel traps provided an estimate of the proportion of males that was nearly identical to the proportion from within infestations (0.51 ± 0.03). 5 Numbers of marked beetles captured in traps were uncorrelated with the numbers of marked beetles emerging from the focal infestations. This suggests that traps positioned around an infestation may not be effective at estimating relative abundances of beetles within the infestation.     

Reducing spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) emergence for hibernation in central British Columbia by felling infested trees

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Continuous population estimates forDendroctonus frontalis Zimm. (Coleoptera: Scolytidae) occurring in infestations

Infestations of Dendroctonus frontalisZimm. are often observed to enlarge continuously by the colonization of new hosts in a pattern similar to a forest fire. This pattern of infestation growth presents unique problems in quantitatively estimating populations of D. frontalis. Beetle populations on each infested tree in an infestation go through five processes: attack, oviposition, reemergence, survivorship, and emergence. These processes, which have been described mathematically in the literature, each take several days for completion. In order to follow the distribution and abundance of D. frontalis throughout the course of development of a spot, we need a daily estimate of the number of beetles involved in each process on every tree. Since it is not practical to sample each tree daily, we developed a procedure whereby quantitative estimation procedures for within-tree populations were used in combination with the mathematical models for the life processes to produce a daily record of the number of adults successfully attacking trees, the number of eggs oviposited, the number of beetles reemerging, number of beetles surviving within the trees, and the number of beetles emerging. These daily estimates were then summarized for all trees in the spot for the duration of the infestation. The daily record of populations of D. frontalis, used with information on infestation geometry, were suggested to be of value in describing and elucidating several important facets of population dynamics including dispersal patterns within infestations, between tree beetle loss (mortality), and time lags among the various population processes. The information reported can be used to develop simulation models of population dynamics or to validate existing models.     

Ability of stress‐related volatiles to attract and induce attacks by Xylosandrus germanus and other ambrosia beetles

• 1 Xylosandrus germanus typically colonizes physiologically‐stressed deciduous hosts but it is increasingly being recognized as a key pest of ornamental nursery stock. We tested the attractiveness of common plant stress‐related volatiles to ambrosia beetles occupying the nursery agroecosystem, as well as their ability to induce attacks on selected trees. Experiments were conducted in Ohio, U.S.A.
• 2 Stress volatile attractiveness was first assessed by positioning traps baited with acetaldehyde, acetone, ethanol and methanol in ornamental nurseries. Cumulative trap counts confirmed that ethanol was the most attractive stress‐related volatile to X. germanus. Methanol‐baited traps were slightly attractive to X. germanus, whereas traps baited with acetaldehyde and acetone were not attractive to any ambrosia beetle.
• 3 A series of tree injection experiments were also conducted to determine the ability of these volatiles to induce attacks by ambrosia beetles under field conditions. Injection of ethanol into Magnolia virginiana induced the largest number of attacks, whereas injection of acetaldehyde induced more attacks than methanol or acetone. Xylosandrus germanus was the most predominant species emerging from M. virginiana injected with each of the stress‐related volatiles. No attacks by wood‐boring beetles were observed on water injected or uninjected control trees.
• 4 Solid‐phase microextraction–gas chromatography–mass spectrometry confirmed the emission of acetaldehyde, acetone, ethanol and methanol after their injection into M. virginiana.
• 5 Xylosandrus germanus has an efficient olfactory‐based mechanism for differentiating among host volatile cues. Injecting select trees with stress‐related volatiles, particularly ethanol, shows promise as a trap tree strategy for X. germanus and other ambrosia beetles.

     

Tree species and position matter: the role of pests for survival of other insects

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Intraspecific variation in host plant quality: the aphid Cinara cupressi on the Mexican cypress, Cupressus lusitanica

Abstract.

• 1 In Kenya, the aphid Cinara cupressi (Buckton) has an aggregated distribution across individuals of its host plant, the Mexican cypress, Cupressus lusitanica (Mill).
• 2 There are significant differences in host plant quality: C.cupressi nymphs transplanted onto cypress trees with low aphid infestations are significantly less likely to establish than aphids transplanted onto trees with high aphid infestations.
• 3 Given that some transplanted nymphs establish, the previous level of infestation does not influence aphid survival.
• 4 Evidence for local adaptation to individual plants was not found: nymphs transplanted to the trees from which their parents were collected neither established more readily nor showed lower mortality than those transplanted to other trees with similar levels of infestation. Therefore the variation in host plant quality has not led to the development of local adaptation to particular host plants.

     

Early responses of bark and wood boring beetles to an outbreak of the hemlock looper Lambdina fiscellaria (Guenée) (Lepidoptera: Geometridae) in a boreal balsam fir forest of North America

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Effects of the emerald ash borer invasion on the community composition of arthropods associated with ash tree boles in Maryland,U.S.A.

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Sequential patterns of colonization of coarse woody debris by Ips pini (Say) (Coleoptera: Scolytidae) following a major ice storm in Ontario

Abstract

• 1 It is widely known that many bark and wood‐boring beetle species use nonresistant coarse woody debris (CWD) created by disturbances; however, the ability of these secondary species to cause mortality in healthy trees following a build‐up of their populations remains unclear. We characterized the pattern of colonization by Ips pini (Say) following a major ice storm that created large amounts of CWD varying in resistance to colonization (i.e. ranging from snapped tops with no resistance to heavily damaged trees with intact root systems). A major question was how the beetles responded to the different types of storm‐damaged material and whether healthy undamaged trees were colonized and killed following increases in beetle populations.
• 2 Six red pine, Pinus resinosa Ait., plantations in eastern Ontario were monitored from 1998 to 2001 inclusive: three with high storm damage (approximately 120 m³/ha CWD) and three with minimal damage (approximately 20 m³/ha CWD). Transects (200 × 2 m) were sampled yearly in each plantation to assess the type and amount of damaged pine brood material colonized by the pine engraver beetle, I. pini.
• 3 Beetles preferentially infested the most nonresistant material available each year (i.e. all snapped tops in year 1, all standing snags, up‐rooted trees and many bent trees by year 2, but still less than 50% of trees blown over but with intact root systems by year 3). By years 3 and 4, the majority (approximately 75%) of severely damaged trees (with > 50% crown loss) died prior to beetle colonization.
• 4 The size of the beetle population tracked the abundance of available woody material from year‐to‐year within a plantation; populations were very large in the first 2 years, and declined significantly in the last 2 years.
• 5 Healthy standing red pines were apparently resistant to colonization by the beetles, despite the significant build‐up in their populations. Hence, the results of the present study suggest that native bark beetle populations will not cause further tree mortality following such a disturbance in this region.

     

Freeze stress of deciduous trees induces attacks by opportunistic ambrosia beetles

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Attraction of ambrosia and bark beetles to coast live oaks infected by Phytophthora ramorum

1 Sudden oak death is caused by the apparently introduced oomycete, Phytophthora ramorum. We investigated the role of bark and ambrosia beetles in disease progression in coast live oaks Quercus agrifolia. 2 In two Marin County, California sites, 80 trees were inoculated in July 2002 with P. ramorum and 40 were wounded without inoculation. Half of the trees in each group were sprayed with the insecticide permethrin [cyclopropanecarboxylic acid, 3‐(2,2‐dichloroethenyl)‐2,2‐dimethyl‐(3‐phenoxyphenyl) methyl ester] to prevent ambrosia and bark beetle attacks, and then were sprayed twice per year thereafter. After each treatment, sticky traps were placed on only the permethrin‐treated trees. Beetles were collected periodically in 2003. 3 Inoculated trees accounted for 95% of all beetles trapped. The ambrosia beetles Monarthrum scutellare and Xyleborinus saxeseni and the western oak bark beetle Pseudopityophthorus pubipennis were the most abundant of the seven species trapped. 4 Permethrin treatment delayed initiation of beetle attacks and significantly reduced the mean number of attacks per tree. Beetles did not attack any wounded or noncankered inoculated trees. 5 Trees with larger cankers trapped more beetles early in the disease. Once permethrin lost effectiveness, the number of beetle entrance tunnels was a more reliable predictor of subsequent trap catch than was canker size. 6 Beetles were initially attracted to P. ramorum cankers in response to kairomones generated in the host‐pathogen interaction. After beetles attacked the permethrin‐treated trees, aggregation pheromones most probably were the principal factor in beetle colonization behaviour.     

Random movement pattern of fed and unfed adult Colorado potato beetles in bare‐ground habitat

• 1 Although the successful management of the Colorado potato beetle Leptinotarsa decemlineata (Say) depends on the prevention of its dispersal, its walking pattern in the landscape remains poorly understood. In the present study, post‐diapause, early summer, late summer and colony adult beetles, both fed and unfed before release, were tracked with a harmonic radar to establish their walking movement pattern in a bare‐ground field.
• 2 The random walk model successfully described the dispersal of all beetle types, whether fed or unfed.
• 3 The diverse life history of this species was manifested by an increased distance travelled and deviations of individual paths from the random model. Starved post‐diapause beetles travelled furthest and individual paths deviating from random were both local and directed, probably aiming to maximize opportunities for host colonization. Starved early summer beetles also travelled further than fed beetles but relied more on random movement to disperse in the habitat. Starving had little impact on the distance travelled or the path deviations of late summer beetles that are searching for overwintering site rather than hosts.
• 4 The increased displacement of starving beetles over fed beetles corresponded with an increased walking step and index of straightness.
• 5 The impact of starvation on travel distance was greater than expected from laboratory tests.
• 6 In conclusion, the results obtained in the present study suggest a random walking pattern to search arable land until host volatile or visual impulses trigger a more directed walk or flight.

     

Relationships among wood‐boring beetles,fungi, and the decomposition of forest biomass

A prevailing paradigm in forest ecology is that wood‐boring beetles facilitate wood decay and carbon cycling, but empirical tests have yielded mixed results. We experimentally determined the effects of wood borers on fungal community assembly and wood decay within pine trunks in the southeastern United States. Pine trunks were made either beetle‐accessible or inaccessible. Fungal communities were compared using culturing and high‐throughput amplicon sequencing (HTAS) of DNA and RNA. Prior to beetle infestation, living pines had diverse fungal endophyte communities. Endophytes were displaced by beetle‐associated fungi in beetle‐accessible trees, whereas some endophytes persisted as saprotrophs in beetle‐excluded trees. Beetles increased fungal diversity several fold. Over forty taxa of Ascomycota were significantly associated with beetles, but beetles were not consistently associated with any known wood‐decaying fungi. Instead, increasing ambrosia beetle infestations caused reduced decay, consistent with previous in vitro experiments that showed beetle‐associated fungi reduce decay rates by competing with decay fungi. No effect of bark‐inhabiting beetles on decay was detected. Platypodines carried significantly more fungal taxa than scolytines. Molecular results were validated by synthetic and biological mock communities and were consistent across methodologies. RNA sequencing confirmed that beetle‐associated fungi were biologically active in the wood. Metabarcode sequencing of the LSU/28S marker recovered important fungal symbionts that were missed by ITS2, though community‐level effects were similar between markers. In contrast to the current paradigm, our results indicate ambrosia beetles introduce diverse fungal communities that do not extensively decay wood, but instead reduce decay rates by competing with wood decay fungi.     

Observations and modeling of aboveground tree carbon stocks and fluxes following a bark beetle outbreak in the western United States

Bark beetle epidemics result in tree mortality across millions of hectares in North America. However, few studies have quantified impacts on carbon (C) cycling. In this study, we quantified the immediate response and subsequent trajectories of stand‐level aboveground tree C stocks and fluxes using field measurements and modeling for a location in central Idaho, USA that experienced an outbreak of mountain pine beetle (Dendroctonus ponderosae Hopkins). We measured tree characteristics in lodgepole pine (Pinus contorta) plots spanning a range of structure and mortality conditions. We then initialized the forest vegetation simulator, an individual tree‐based model, with these measurements and simulated the response of aboveground production of C fluxes as well as trajectories of C stocks and fluxes in the coming decades. Mountain pine beetles killed up to 52% of the trees within plots, with more larger trees killed. C stocks in lodgepole pine were reduced by 31–83% following the outbreak, and plot‐level C fluxes decreased 28–73%. Modeled C stocks increased nearly continuously following the infestation, recovering to preoutbreak levels in 25 years or less. Simulated aboveground tree C fluxes increased following the immediate postoutbreak decrease, then subsequently declined. Substantial variability of C stocks and fluxes among plots resulted from the number and size of killed and surviving trees. Our study illustrates that bark beetle epidemics alter forest C cycling unlike stand‐replacement wildfires or clear‐cut harvests, due in part to incomplete mortality coupled with the preference by beetles for larger trees. The dependency of postoutbreak C stocks and fluxes on stand structure suggests that C budget models and studies in areas experiencing mountain pine beetle disturbances need to include size distribution of trees for the most accurate results.     

A predictive framework to assess spatio‐temporal variability of infestations by the European spruce bark beetle

Natural disturbances are key factors for the development of forest ecosystems. In forests of central Europe and Scandinavia, the European spruce bark beetle Ips typographus is the most devastating biotic disturbance agent in Norway spruce Picea abies, but our understanding of the factors determining its spatio‐temporal dynamics is still quite limited. To quantify the drivers of bark beetle dynamics, we analyzed a survey dataset with annual resolution that covers 17 yr and 469 forest districts (10 860 km² of forest area) all over Switzerland. We used Poisson log‐normal models in a Bayesian framework to analyze the spatio‐temporal dynamics of bark beetle infestation spots at the forest district level. Bark beetle infestations increased with increasing heat sum (> 8.3°C), volume of standing Norway spruce stock, and the number of infestation spots of the previous year. Precipitation tended to slightly affect the risk of bark beetle infestations. Two major storm events further increased the spatio‐temporal variability of bark beetle infestations. Spruce abundance, storm damage and temperature are known to be important factors influencing the population dynamics of the European spruce bark beetle. Our study is the first to quantify the combined effects of spruce abundance and heat sum, whereby the heat sum turned out to be the most important and consistent predictor. Because our study area encompasses large ecological and climatological gradients, our model is likely to be applicable to Norway spruce forests in other regions of central Europe and Scandinavia.     

Negative Feedbacks on Bark Beetle Outbreaks: Widespread and Severe Spruce Beetle Infestation Restricts Subsequent Infestation

Understanding disturbance interactions and their ecological consequences remains a major challenge for research on the response of forests to a changing climate. When, where, and how one disturbance may alter the severity, extent, or occurrence probability of a subsequent disturbance is encapsulated by the concept of linked disturbances. Here, we evaluated 1) how climate and forest habitat variables, including disturbance history, interact to drive 2000s spruce beetle (Dendroctonus rufipennis) infestation of Engelmann spruce (Picea engelmannii) across the Southern Rocky Mountains; and 2) how previous spruce beetle infestation affects subsequent infestation across the Flat Tops Wilderness in northwestern Colorado, which experienced a severe landscape-scale spruce beetle infestation in the 1940s. We hypothesized that drought and warm temperatures would promote infestation, whereas small diameter and non-host trees, which may reflect past disturbance by spruce beetles, would inhibit infestation. Across the Southern Rocky Mountains, we found that climate and forest structure interacted to drive the 2000s infestation. Within the Flat Tops study area we found that stands infested in the 1940s were composed of higher proportions of small diameter and non-host trees ca. 60 years later. In this area, the 2000s infestation was constrained by a paucity of large diameter host trees (> 23 cm at diameter breast height), not climate. This suggests that there has not been sufficient time for trees to grow large enough to become susceptible to infestation. Concordantly, we found no overlap between areas affected by the 1940s infestation and the current infestation. These results show a severe spruce beetle infestation, which results in the depletion of susceptible hosts, can create a landscape template reducing the potential for future infestations.     

Behaviours of phoretic mites (Acari) associated with Ips pini and Ips grandicollis (Coleoptera: Curculionidae) during host‐tree colonization

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