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.     

Site condition and predation influence a bark beetle's success: a spatially realistic approach

Abstract 1 Spatial pattern in abundance of Dendroctonus micans was studied in a 600‐ha spruce stand in the Massif Central (Lozère, France). The proportion of trees attacked was measured in 38 plots and these data were used to estimate spatial pattern of attack density in the stand and to identify a transect of decreasing attack density (80% to 30%) over less than 1000 m. 2 Spatial variation in attack density was analysed in relation to (i) data on site and stand characteristics (altitude, slope, tree density, tree average height, yield class and average age) collected from 63 points in the stand and (ii) the releases of the predator Rhizophagus grandis (localization and number of beetles released). 3 The proportion of attacked trees was analysed using geostatistics and showed a strong spatial structure reflecting the spatial scale of interaction of D. micans with its environment. The spatial structure was modelled in order to estimate the spatial distribution of attack density at unsampled locations. 4 A linear model relating interpolated attack density to the number of predators released 6–10 years before the survey in a 300‐m radius and to the average slope over a 250‐m radius explained 67% of the observed variability. Spatial autocorrelation was taken into account in a spatial regression model.     

Inducibility of chemical defenses in Norway spruce bark is correlated with unsuccessful mass attacks by the spruce bark beetle

Secondary attraction to aggregation pheromones plays a central role in the host colonization behavior of the European spruce bark beetle Ips typographus. However, it is largely unknown how the beetles pioneering an attack locate suitable host trees, and eventually accept or reject them. To find possible biomarkers for host choice by I. typographus, we analyzed the chemistry of 58 Norway spruce (Picea abies) trees that were subsequently either (1) successfully attacked and killed, (2) unsuccessfully attacked, or (3) left unattacked. The trees were sampled before the main beetle flight in a natural Norway spruce-dominated forest. No pheromones were used to attract beetles to the experimental trees. To test the trees' defense potential, each tree was treated in a local area with the defense hormone methyl jasmonate (MeJ), and treated and untreated bark were analyzed for 66 different compounds, including terpenes, phenolics and alkaloids. The chemistry of MeJ-treated bark correlated strongly with the success of I. typographus attack, revealing major chemical differences between killed trees and unsuccessfully attacked trees. Surviving trees produced significantly higher amounts of most of the 39 analyzed mono-, sesqui-, and diterpenes and of 4 of 20 phenolics. Alkaloids showed no clear pattern. Differences in untreated bark were less pronounced, where only 1,8-cineole and (-)-limonene were significantly higher in unsuccessfully attacked trees. Our results show that the potential of individual P. abies trees for inducing defense compounds upon I. typographus attack may partly determine tree resistance to this bark beetle by inhibiting its mass attack.     

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

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Characteristics of Norway spruce trees (Picea abies) surviving a spruce bark beetle (Ips typographus L.) outbreak

The characteristics of spruce individuals, which survived a massive bark beetle outbreak, were compared with the characteristics of neighbouring attacked trees in Šumava National Park (Czech Republic). Selected parameters related to crown geometry, stand conditions and distances between trees were measured or estimated. Significant differences were found between the surviving trees and the neighbouring trees attacked by I. typographus. Trees with a higher level of stem shading (longer crown length) tended to survive. The attacked trees were usually located in areas with larger basal area, especially southwards from them. A shorter distance to a previously attacked tree increased the probability of additional attack. Spruce trees with more progressive crown structure transformation (primary structure defoliation) were significantly more frequently attacked by spruce bark beetle. Superior and taller trees had a clearly longer life expectancy than dominant ones. These results show that the attack of trees by bark beetle can be predicted to a certain degree, which can be used in management of endangered spruce forests.     

Impacts of silvicultural thinning treatments on beetle trap captures and tree attacks during low bark beetle populations in ponderosa pine forests of northern Arizona

Our research used a combination of passive traps, funnel traps with lures, baited trees, and surveys of long-term thinning plots to assess the impacts of different levels of stand basal area (BA) on bark beetle tree attack and on trap captures of Ips spp., Dendroctonus spp., and their predators. The study occurred at two sites in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests, from 2004 to 2007 during low bark beetle populations. Residual stand BA ranged from 9.0 to 37.0 m2/ha. More predators and bark beetles were collected in passive traps in stands of lower BA than in stands of higher BA; however, significance varied by species and site, and total number of beetles collected was low. Height of the clear panel passive traps affected trap catches for some species at some sites and years. When pheromone lures were used with funnel traps [Ips pini (Say) lure: lanierone, +03/-97 ipsdienol], we found no significant difference in trap catches among basal area treatments for bark beetles and their predators. Similarly, when trees were baited (Dendroctonus brevicomis LeConte lure: myrcene, exo-brevicomin and frontalin), we found no significant difference for days to first bark beetle attack. Surveys of long-term thinning treatments found evidence of bark beetle attacks only in unthinned plots (approximately 37 m2/ha basal area). We discuss our results in terms of management implications for bark beetle trapping and control.     

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.

     

Olfactory recognition and behavioural avoidance of angiosperm nonhost volatiles by conifer-inhabiting bark beetles

Abstract 1 When searching for suitable hosts in flight, especially in mixed forests, conifer‐inhabiting bark beetles will encounter not only suitable host trees and their odours, but also unsuitable hosts and nonhost trees. Rejection of these trees could be based on an imbalance of certain host characteristics and/or a negative response to some nonhost stimuli, such as nonhost volatiles (NHV). 2 Recent electrophysiological and behavioural studies clearly indicate that conifer‐inhabiting bark beetles are not only able to recognize, but also to avoid, nonhost habitats or trees by olfactory means. Green leaf volatiles (GLV), especially C₆‐alcohols, from the leaves (and partly from bark) of nonhost angiosperm trees, may represent nonhost odour signals at the habitat level. Specific bark volatiles such as trans‐conophthorin, C₈‐alcohols, and some aromatic compounds, may indicate nonhosts at the tree species level. Flying bark beetles are also capable of determining whether a possible host is unsuitable by reacting to signals from conspecifics or sympatric heterospecifics that indicate old or colonized host tree individuals. 3 Combined NHV signals in blends showed both redundancy and synergism in their inhibitory effects. The coexistence of redundancy and synergism in negative NHV signals may indicate different functional levels (nonhost habitats, species, and unsuitable hosts) in the host selection process. Combinations of NHV and verbenone significantly reduced the number of mass attacked host trees or logs on several economically important species (e.g. Dendroctonus ponderosae, Ips typographus, and I. sexdentatus). 4 We suggest a semiochemical‐diversity hypothesis, based on the inhibition by NHV of bark beetle host‐location, which might partly explain the lower outbreak rates of forest insects in mixed forests. This ‘semiochemical‐diversity hypothesis’ would provide new support to the general ‘stability‐diversity hypothesis’. 5 Natural selection appears to have caused conifer‐inhabiting bark beetles to evolve several olfactory mechanisms for finding their hosts and avoiding unsuitable hosts and nonhost species. NHV and unsuitable host signals have potential for use in protecting trees from attack. The use of these signals may be facilitated by the fact that their combination has an active inhibition radius of several metres in trap test, and by the observation of area effects for several trees near inhibitor soruces in tree protection experiments. Furthermore, incorporation of negative signals (such as NHV and verbenone) and pheromone‐based mass‐trapping in a ‘push–pull’ fashion may significantly increase the options for control against outbreaks of conifer‐inhabiting bark beetles, especially in high risk areas.     

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.

     

Induced defence reaction in Scots pine following stem attacks by Tomicus piniperda

The induced defence reaction of Scots pine Pinus sylvestris to stem attacks by the common pine shoot beetle, Tomicus piniperda (Col, Scolytidae), was studied by inducing natural attacks on young Scots pine trees of different vigour Pruned trees were more heavily attacked by T piniperda than unpruned ones Vigorous trees successfully contained the attacking beetles within resinous lesions, whereas less vigorous trees failed to defend themselves The content of resin acids increased dramatically in the lesions on all trees, and the resin acid composition differed somewhat between some treatments Starch accumulation in the inner bark of the main stem was lower in pruned trees than in unpruned ones, and starch reserves were depleted in the most heavily attacked trees All but one of the unpruned trees survived the attack, whereas most of the severely pruned ones died following attack by beetles at densities exceeding c 300 egg galleries m⁻²     

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.     

The role of host tree condition in attack of white oaks by the twolined chestnut borer,Agrilus bilineatus (Weber) (Coleoptera: Buprestidae)

Summary The twolined chestnut borer, Agrilus bilineatus (Weber) (Coleoptera: Buprestidae), attacks stressed oaks (Quercus spp.) and is associated with extensive mortality of trees in the eastern deciduous forests of North America. We examined host location by the insect and subsequent host mortality in experimentally stressed trees. A. bilineatus adults were able to rapidly and specifically locate stressed oak trees. Up to 160 beetles per week were captured on sticky band traps on the trunks of stressed trees, while beetles rarely landed on unstressed control trees. This suggests that adult borers have an acute perception of host tree quality, and that this perception is from a distance. One mechanism of host location may be detection of volatile compounds produced by stressed trees.The condition of the host tree appears to regulate both beetle attraction and successful colonization. Mortally wounded (xylem-girdled) trees attracted beetles only until the cambium died. Xylem-girdled trees were attacked early in the beetle flight season, but larvae did not survive to emerge as adults from these trees. In contrast, phloemgirdled trees continued to attract beetles throughout the flight period. Phloem-girdled trees which were heavily attacked by A. bilineatus died late in the season in which they were attacked. Lightly attacked trees survived until the following growing season, and were then heavily attacked and killed. In one stand, phloem-girdled trees were not attacked, healed over the girdling wounds and were still alive three years after girdling. These results indicate that oak trees are only attractive to A. bilineatus within a narrow range of physiological conditions following stress but prior to mortality. A. bilineatus appears to be a proximate agent of mortality in stressed oaks in eastern North America.This paper (85-7-8-216) resulted from a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director     

Density related plasticity in stand-level spatial distribution of the ambrosia beetle, Platypus koryoensis (Coleoptera: Curculionidae)

We tested the hypothesis that the population density of ambrosia beetles at the stand level influences the spatial distribution of infested trees. We evaluated the spatial distribution of the ambrosia beetle, Platypus koryoensis (Murayama) in three oak forest stands that varied in beetle population density using a multi-year trapping survey. We used these data to inform a clustering analysis based on aggregation indices using the SADIE software. Four important findings emerged: (1) the spatial distribution pattern of P. koryoensis at the stand level changed as the population density of the beetle varied; (2) at low population densities, beetle distribution was contagious at the stand level; (3) as beetle population densities increased, the spatial distribution of infested trees became random, potentially due to beetle avoidance of mass attacked trees; and (4) at high beetle population densities, the spatial distribution of infested trees became contagious, possibly due to temporal changes in location of the attack epicenter within the stand. Our results support the hypothesis that beetle population density has consequences for the spatial distribution of infested trees at the within-stand scale. We conclude that the spatial distribution of infested trees is flexible in response to beetle population density, suggesting that beetle attack behaviors are mediated by one or more density-dependent effects.     

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

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Spatial and temporal dynamics of bark beetles in Chinese white pine in Qinling Mountains of Shaanxi Province, China

Spatial and temporal dynamics of bark beetles in single tree trunks of Pinus armandi were studied in Qinling Mountains, Shaanxi Province, China. Ten species of engraver bark beetles attacked from healthy to withered Chinese white pines, but seven species were commonly detected every year in Qinling forest ecosystem. Dendroctonus armandi and Hylurgops longipilis were common species at the lower of trunks, and Ips acuminatus, Polygraphus sinensis, and Pityogenes japonicus primary distributed in the middle of trunks, whereas population densities of Cryphalus lipingensis and C. chinlingensis centralized at the upper trunks and branches of Chinese white pines. On the time series, D. armandi, as a dominant species in Chinese white pines of Qinling forest ecosystem, mainly attacked healthy and weakened trees and cooperated with blue stain fungus that resulted in the declining abruptly resistance and triggered the secondary bark beetles to attack the infected or withered host trees. Attacking and colonizing phenology of bark beetles in Qinling forest ecosystem are caused by complex interactions among spatial and trophic competition and cooperation and exhibit particular spatial and temporal patterns. Our results support the view that competition and cooperation within bark beetles are a critical factor to influence bark beetles spatial and temporal distribution, and stability of bark beetles' ecosystem, D. armandi, I. acuminatus, P. japonicus, P. sinensis, C. lipingensis, C. chinlingensis, and H. longipilis in Chinese white pine of Qinling forest ecosystem.     

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.     

Host specificity of ambrosia and bark beetles (Col., Curculionidae: Scolytinae and Platypodinae) in a New Guinea rainforest

Abstract.  1. Bark and ambrosia beetles are crucial for woody biomass decomposition in tropical forests worldwide. Despite that, quantitative data on their host specificity are scarce.
2. Bark and ambrosia beetles (Scolytinae and Platypodinae) were reared from 13 species of tropical trees representing 11 families from all major lineages of dicotyledonous plants. Standardised samples of beetle-infested twigs, branches, trunks, and roots were taken from three individuals of each tree species growing in a lowland tropical rainforest in Papua New Guinea.
3. A total of 81 742 beetles from 74 species were reared, 67 of them identified. Local species richness of bark and ambrosia beetles was estimated at 80–92 species.
4. Ambrosia beetles were broad generalists as 95% of species did not show any preference for a particular host species or clade. Similarity of ambrosia beetle communities from different tree species was not correlated with phylogenetic distances between tree species. Similarity of ambrosia beetle communities from individual conspecific trees was not higher than that from heterospecific trees and different parts of the trees hosted similar ambrosia beetle communities, as only a few species preferred particular tree parts.
5. In contrast, phloeophagous bark beetles showed strict specificity to host plant genus or family. However, this guild was poor in species (12 species) and restricted to only three plant families (Moraceae, Myristicaceae, Sapindaceae).
6. Local diversity of both bark and ambrosia beetles is not driven by the local diversity of trees in tropical forests, since ambrosia beetles display no host specificity and bark beetles are species poor and restricted to a few plant families.     

Colonization of disturbed Scots pine trees by bark‐ and wood‐boring beetles

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Mass attack of trees by Ips typographus induced by sex-specific pheromone: a model of attack dynamics

A model of attack dynamics on a single tree under mass attack by the spruce bark beetle Ips typographus is developed from earlier studies of pheromone production and response of the flying population to the pheromone signal. The model is explored for different numbers and sex-ratios of pioneer beetles and responding beetles. Both numbers of pioneers and responders were important for the dynamics, while the sex-ratio of pioneers was less important. A decreased proportion of males among the responding beetles was important for the result of the model. In the early stage of the attack the model predictions of both numbers and sex-ratio of beetles landing were similar to examples from naturally mass-attacked trees, although the attack proceeded more slowly than in the field. Several aspects of the attack dynamics, such as density-regulation of the number of attacks on the tree, as observed in the field, were not predicted by the model. Gaps in the knowledge of the chemical ecology of the tree-insect relation, such as qualitative change in odours from the tree, spacing mechanisms on the bark surface, and mortality due to host resistence, are pointed out with help of the model.