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

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Attractant effect of citral on Platypus koryoensis (Coleoptera: Curculionidae)

An environmentally friendly control strategy for the ambrosia beetle Platypus koryoensis (Murayama), a vector of oak wilt disease, was investigated using the citral as a pheromone component. To develop an effective attractant trap, a semi‐field experiment (Yeoju) and a field experiment (Gunpo) were performed. Different ratios of four different attractant mixtures (ethanol 100 %, ethanol + citral (90:10, 95:5), ethanol + citral + hexane extract of Quercus mongolica (90:5:5)) were investigated to determine their effects on P. koryoensis. The 95:5 mixture of ethanol + citral had the greatest attractant effect on P. koryoensis. However, other ambrosia beetle species did not exhibit a specific response to citral. Our findings indicate that the pheromone citral was effective at attracting P. koryoensis and that certain concentrations of citral could be used to develop an effective and environmentally friendly control agent.     

Stand-level distribution and movement of Platypus quercivorus adults and patterns of incidence of new infestation

Abstract 1 Flying populations of an ambrosia beetle, Platypus quercivorus (Murayama), a vector of an ambrosia fungus Raffaelea quercivora, which causes deciduous oak diebacks in Japan, were determined by sticky screen traps. 2 Platypus quercivorus beetles tended to move upwards along slopes. The highest concentrations of flying beetles usually occured at the upper forest margins. 3 During the period when the number of flying beetles was increasing, the incidence of newly infested trees spread from the epicentre into the forest. During the period when the number of flying beetles was decreasing, the epicentre shrank into the upper forest edge. Newly infested trees did not occur in this period because most trees had already been infested. 4 Near the upper forest edge, where many beetles were highly concentrated throughout the season, the number of new entry holes decreased greatly after the initial attack early in the season, although many adults were present throughout the period.     

Suitability of California bay laurel and other species as hosts for the non‐native redbay ambrosia beetle and granulate ambrosia beetle

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Impacts of laurel wilt disease on redbay (Persea borbonia (L.) Spreng.) population structure and forest communities in the coastal plain of Georgia, USA

Laurel wilt disease (LWD), a fungal disease vectored by the non-native redbay ambrosia beetle (Xyleborus glabratus Eichhoff), has caused mortality of redbay (Persea borbonia (L.) Spreng.) in the coastal plain of Georgia since 2003. Despite its rapid spread, little research has evaluated its impacts on redbay population structure and forest communities. Diseased populations of redbay in five sites (2–4 years post infestation) were compared to healthy populations in three uninfested sites in five counties in Georgia. The results showed high redbay mortality, shifts in size structure, and changes in community composition. An average of 90 % of redbay trees ≥3 cm diameter at breast height (DBH) were dead in infested sites, compared to 0–35 % in control sites. Mortality was seen in individuals of the smallest stem diameter category (<1.00 cm diameter at ground height). DBH of live redbay trees in control sites was twice that of those in infested sites. Photosynthetically active radiation was 4.8 times greater at infested sites than control sites due to loss of redbay canopy. Community structure measurements showed redbay trees had the greatest mean importance value (IV) at control sites compared to the 8th mean IV at infested sites for live stems. Two species co-dominant to redbay, sweetbay (Magnolia virginiana L.) and loblolly bay (Gordonia lasianthus (L.) J. Ellis), were of higher importance at infested than control sites, suggesting they are increasing in dominance following the mortality of redbay. This study shows LWD has impacted redbay populations and altered associated forest communities in Georgia.     

Microstructure of the prothoracic mycangia in the ambrosia beetle Platypus koryoensis (Coleoptera: Curculionidae: Platypodinae)

Platypus koryoensis (Murayama) (Coleoptera: Curculionidae: Platypodinae) is a minute ambrosia beetle that can cause significant damage to live oak trees (Quercus mongolica) along with its symbiotic pathogenic fungus (Raffaelea sp.). Although both sexes of the beetle have a number of cuticular depressions for carrying microorganisms, only the females have mycangia that are typical in size on the pronotum. The mycangial cavities are filled with a mass of amorphous substances including yeast-like spores. In addition, mycangial cavities are equipped with cuticular pores which release substances to support symbiotic microorganisms during storage. Small depressions of both sexes can be subdivided into two clusters according to their relative sizes and locations. Although they have a typical cuticular peg (basiconic sensillum), they also share a common morphological characteristic with the typical mycangium of female: a round outlet and an invaginated reservoir containing a mass of secretory substances. Accordingly, our fine structural data suggest that both the female mycangia and the clusters of small cuticular depressions in both sexes also have a mycangial function in common regardless of their relative size and location.     

The influence of forest stand and site characteristics on the composition of exotic dominated ambrosia beetle communities (Coleoptera: Curculionidae: Scolytinae)

Economic and biological consequences are associated with exotic ambrosia beetles and their fungal associates. Despite this, knowledge of ambrosia beetles and their ecological interactions remain poorly understood, especially in the oak-hickory forest region. We examined how forest stand and site characteristics influenced ambrosia beetle habitat use as evaluated by species richness and abundance of ambrosia beetles, both the native component and individual exotic species. We documented the species composition of the ambrosia beetle community, flight activity, and habitat use over a 2-yr period by placing flight traps in regenerating clearcuts and older oak-hickory forest stands differing in topographic aspect. The ambrosia beetle community consisted of 20 species with exotic ambrosia beetle species dominating the community. Similar percentages of exotic ambrosia beetles occurred among the four forest habitats despite differences in stand age and aspect. Stand characteristics, such as stand age and forest structure, influenced ambrosia beetle richness and the abundances of a few exotic ambrosia beetle species and the native ambrosia beetle component. Topographic aspect had little influence on ambrosia beetle abundance or species richness. Older forests typically have more host material than younger forests and our results may be related to the amount of dead wood present. Different forms of forest management may not alter the percent contribution of exotic ambrosia beetles to the ambrosia beetle community.     

Spatial Pattern of Attacks of the Invasive Woodwasp Sirex noctilio,at Landscape and Stand Scales

Invasive insect pests are responsible for important damage to native and plantation forests, when population outbreaks occur. Understanding the spatial pattern of attacks by forest pest populations is essential to improve our understanding of insect population dynamics and for predicting attack risk by invasives or planning pest management strategies. The woodwasp Sirex noctilio is an invasive woodwasp that has become probably the most important pest of pine plantations in the Southern Hemisphere. Our aim was to study the spatial dynamics of S. noctilio populations in Southern Argentina. Specifically we describe: (1) the spatial patterns of S. noctilio outbreaks and their relation with environmental factors at a landscape scale; and (2) characterize the spatial pattern of attacked trees at the stand scale. We surveyed the spatial distribution of S. noctilio outbreaks in three pine plantation landscapes, and we assessed potential associations with topographic variables, habitat characteristics, and distance to other outbreaks. We also looked at the spatial distribution of attacked trees in 20 stands with different levels of infestation, and assessed the relationship of attacks with stand composition and management. We found that the spatial pattern of pine stands with S. noctilio outbreaks at the landscape scale is influenced mainly by the host species present, slope aspect, and distance to other outbreaks. At a stand scale, there is strong aggregation of attacked trees in stands with intermediate infestation levels, and the degree of attacks is influenced by host species and plantation management. We conclude that the pattern of S. noctilio damage at different spatial scales is influenced by a combination of both inherent population dynamics and the underlying patterns of environmental factors. Our results have important implications for the understanding and management of invasive insect outbreaks in forest systems.     

Effect of trap type, trap position, time of year, and beetle density on captures of the redbay ambrosia beetle (Coleoptera: Curculionidae: Scolytinae)

The exotic redbay ambrosia beetle, Xyleborus glabratus Eichhoff (Coleoptera: Curculionidae: Scolytinae), and its fungal symbiont Raffaellea lauricola Harrington, Fraedrich, and Aghayeva are responsible for widespread redbay, Persea borbonia (L.) Spreng., mortality in the southern United States. Effective traps and lures are needed to monitor spread of the beetle and for early detection at ports-of-entry, so we conducted a series of experiments to find the best trap design, color, lure, and trap position for detection of X. glabratus. The best trap and lure combination was then tested at seven sites varying in beetle abundance and at one site throughout the year to see how season and beetle population affected performance. Manuka oil proved to be the most effective lure tested, particularly when considering cost and availability. Traps baited with manuka oil lures releasing 5 mg/d caught as many beetles as those baited with lures releasing 200 mg/d. Distributing manuka oil lures from the top to the bottom of eight-unit funnel traps resulted in similar numbers of X. glabratus as a single lure in the middle. Trap color had little effect on captures in sticky traps or cross-vane traps. Funnel traps caught twice as many beetles as cross-vane traps and three times as many as sticky traps but mean catch per trap was not significantly different. When comparing height, traps 1.5 m above the ground captured 85% of the beetles collected but a few were caught at each height up to 15 m. Funnel trap captures exhibited a strong linear relationship (r2 = 0.79) with X. glabratus attack density and they performed well throughout the year. Catching beetles at low densities is important to port of entry monitoring programs where early detection of infestations is essential. Our trials show that multiple funnel traps baited with a single manuka oil lure were effective for capturing X. glabratus even when no infested trees were visible in the area.     

云南切梢小蠹对云南松树的蛀干危害及致死机理

蛀干危害是云南切梢小蠹致死云南松树的关键环节。通过控制云南切梢小蠹蛀干密度,对云南切梢小蠹在自然条件下蛀干行为与危害进行了首次探讨。结果表明,云南切梢小蠹蛀干密度与云南松存活率呈负相关,蛀干密度直接决定云南松死亡或存活。研究发现,蛀干密度115坑/m2是云南松树的最低致死密度阈值,云南松树在蛀干密度低于26.4坑/m2情况下存活,在26.4-115坑/m2有部分存活,超过115坑/m2以后将被害致死。云南切梢小蠹对树干攻击形成有卵和无卵两类坑道。形成无卵坑道的蛀干攻击可导致树势衰弱,形成有卵坑道的蛀干危害严重破坏了韧皮组织,是导致云南松死亡的直接原因。     

Spatial spread and infestation risk assessment in the Asian longhorned beetle,Anoplophora glabripennis

The Asian longhorned beetle (ALB), Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae), is recognised as potentially one of the most damaging invasive insects in Europe and North America. International trade has increased the risk of accidental introduction of ALB. An eradication programme was initiated in Northeast Italy in June 2009, when an ALB infestation was discovered. The infestation was monitored by annual surveys of all host‐tree species growing in the eradication area. Infested trees were cut down and chipped. This study analyses the spatiotemporal distribution of infested trees for a 5‐year period from 2008 to 2012 using a generalised linear model approach. The results show that spread and infestation risk were significantly affected by (1) distance of suitable hosts from the nearest infested trees, (2) number of infested trees in the surroundings, and (3) annual variation. The significant differences in beetle dispersal between years reflect to some extent the onset of the eradication programme. The model allowed the estimation of arbitrary probability‐based management boundaries surrounding ALB‐infested trees. For example, the model estimated a 0.1% probability of attack on a suitable host tree 1 910 m from an existing attack.     

云南松三种同域共存切梢小蠹梢转干期的空间分布格局

分布在我国西南地区的横坑切梢小蠹,云南切梢小蠹和短毛切梢小蠹同域危害寄主云南松,给林业生产带来巨大损失。为探讨同域切梢小蠹种群在共存下对其空间分布格局的影响,采用传统聚集指标法和地统计学方法研究了三者在梢转干期不同受害云南松纯林树冠中的空间分布型。结果表明重度受害样地中云南切梢小蠹种群密度显著高于横坑切梢小蠹,在轻度受害样地则相反;传统聚集指标法结果显示同域共存的3种切梢小蠹种群在不同受害程度云南松中均为聚集分布,横坑切梢小蠹和云南切梢小蠹聚集是由环境因素和昆虫本身的聚集习性引起;地统计学结果表明除重度受害样地中短毛切梢小蠹呈随机分布外,其余切梢小蠹在不同种群密度下均呈聚集分布;除重度受害样地横坑切梢小蠹外,其他小蠹的空间依赖范围为4.01—7.45 m。横坑切梢小蠹和云南切梢小蠹在不同受害林分中拟合的半变异函数模型在球形模型和高斯模型之间转换。同域共存关系不影响不同种群密度下的切梢小蠹种群空间分布类型,但影响其半变异函数模型和理论参数。     

Past attacks influence host selection by the solitary bark beetle Dendroctonus micans

1. A spatio‐temporal study of host selection and local spread of a solitary bark beetle attacking live spruce Dendroctonus micans (Kugelann) was carried out using a combination of standard statistical methods, geostatistical analyses, and modelling. The study was based on data from three plots (150–300 trees, 0.3–1 ha) from 1978 to 1993. All trees were mapped and successful and abortive bark‐beetle attacks on each tree were counted annually. Because the attacked trees usually survived, temporal attack patterns as well as spatial patterns could be analysed. 2. The distribution of successful insect attacks on the trees was slightly aggregative, indicating some degree of choice rather than totally random establishment. 3. The level of yearly individual attacks per tree was very stable, suggesting that D. micans usually leave the host in which they develop. 4. The attacked trees were distributed randomly in the plots; at the study's spatial scale, the insects dispersed freely throughout the plot (no spatial dependence). 5. On the other hand, time dependence was strong; some trees were attacked repeatedly while others were left untouched. 6. Among a choice of scenarios (random attack, fixed variability in individual host susceptibility, induced host susceptibility following random attack), the best fit was obtained with the model involving induced individual host susceptibility. This type of relation to the host tree contrasts strongly with patterns generally described in host–plant relationships (including gregarious, tree‐killing bark beetles), where local herbivore damage results in induced resistance. 7. These results suggest that the first attacks in a new stand are made at random, that all or most of the beetles emerging from a tree disperse and resample the stand, and that they settle preferentially on trees that were colonised successfully by previous generations.     

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.     

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.

     

Vertical stratification in the spatial distribution of the beech scale insect (Ultracoelostoma assimile) in Nothofagus tree canopies in New Zealand

Abstract. 1. The degree of infestation by New Zealand sooty beech scale insects (Ultracoelostoma assimile, Homoptera: Margarodidae) varies dramatically among adjacent southern beech trees (Nothofagus spp., Fagaceae), but has previously been assumed to be uniformly or randomly distributed within individual host trees. In this study, a full‐census survey was conducted from ground level to canopy level on 14 naturally occurring, canopy‐dominant red beech (Nothofagus fusca) trees (size range 38.7–107.6 cm diameter at breast height) to determine the degree of within‐tree heterogeneity in herbivore density. 2. The within‐tree distribution of the sooty beech scale was vertically stratified and highly heterogeneous, with the greatest densities occurring on bark surfaces in the canopy rather than on the trunk, and on the lower rather than upper sides of the branches. The spatial distribution was strongly negatively correlated with trunk and branch diameter, and increasing bark thickness (as a function of diameter) provides a plausible explanation for differences in the establishment and population density of sooty beech scale insects with trunk and branch size. Furthermore, there was a significant change in the spatial distribution of scale insect populations on trunks and branches of trees of increasing diameter at breast height. This indicates a strong temporal component to the spatial dynamics of the sooty beech scale insect driven by changing host phenology. Future studies on phytophagous insects infesting large host trees need to consider more explicitly changes in population dynamics through space and time. 3. Because of the high degree of within‐tree heterogeneity in population density, the total population size of scale insects on an individual tree could not be predicted from any measure of population density low on the trunk. However, the dry weight biomass of sooty mould fungi growing on the ground beneath infested trees was a remarkably accurate predictor of the total population size of scale insects. The use of sooty mould fungi as a relative measure of population size could be incorporated into studies of other honeydew‐producing hemipterans, since the growth of sooty mould is a distinctive feature synonymous with high concentrations of honeydew production worldwide.     

A local risk map using field observations of the Asian longhorned beetle to optimize monitoring activities

The Asian longhorned beetle Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae) is one of the most dangerous xylophagous pests affecting broadleaf trees in the world. Eradication programmes are undertaken in non‐native regions, requiring extensive resources and involving high costs. An adapted strategy must be set up to optimize the ratio cost/probability of success. We developed a method to generate a risk index of A. glabripennis presence at a local scale, in the surrounding area of an infestation, using field observations (counts of adult insects, exit holes and infested trees). The method, mathematically based on the bivariate symmetric Laplace distribution, has thus reasonable input requirements. The output risk map is easy to interpret and can be directly used by decision‐makers. We used our approach in three infestations in Switzerland. The risk map represented well the insect pressure (beetle population density). We highlighted the fact that survey boundaries, commonly chosen using constant distances from the infestation, should be selected regarding the spatial distribution of the insect pressure, to prioritize monitoring activities. The risk map provides a helpful instrument for advanced survey planning after a first overview, for example to decide which area and which host trees should be inspected for infestations.     

THE EVOLUTION OF AGRICULTURE IN BEETLES (CURCULIONIDAE: SCOLYTINAE AND PLATYPODINAE)

Abstract Beetles in the weevil subfamilies Scolytinae and Platypodinae are unusual in that they burrow as adults inside trees for feeding and oviposition. Some of these beetles are known as ambrosia beetles for their obligate mutualisms with asexual fungi—known as ambrosia fungi—that are derived from plant pathogens in the ascomycete group known as the ophiostomatoid fungi. Other beetles in these subfamilies are known as bark beetles and are associated with free‐living, pathogenic ophiostomatoid fungi that facilitate beetle attack of phloem of trees with resin defenses. Using DNA sequences from six genes, including both copies of the nuclear gene encoding enolase, we performed a molecular phylogenetic study of bark and ambrosia beetles across these two subfamilies to establish the rate and direction of changes in life histories and their consequences for diversification. The ambrosia beetle habits have evolved repeatedly and are unreversed. The subfamily Platypodinae is derived from within the Scolytinae, near the tribe Scolytini. Comparison of the molecular branch lengths of ambrosia beetles and ambrosia fungi reveals a strong correlation, which a fungal molecular clock suggests spans 60 to 21 million years. Bark beetles have shifted from ancestral association with conifers to angiosperms and back again several times. Each shift to angiosperms is associated with elevated diversity, whereas the reverse shifts to conifers are associated with lowered diversity. The unusual habit of adult burrowing likely facilitated the diversification of these beetle‐fungus associations, enabling them to use the biomass‐rich resource that trees represent and set the stage for at least one origin of eusociality.     

Modeling the Influence of Resource Availability on Population Densities of Oxyops vitiosa (Coleoptera: Curculionidae), a Biological Control Agent of the Invasive Tree Melaleuca quinquenervia

The Australian snout beetle, Oxyops vitiosa, was introduced to south Florida, USA, as a biological control agent of the invasive tree Melaleuca quinquenervia during the spring of 1997. As part of ongoing post-release evaluations of this weevil, we sought to quantify the population density that can be maintained by O. vitiosa larvae when fully exploiting the available melaleuca foliage. Seasonal population densities were modeled as a function of tree size distribution and density, plant phenology, quantity of acceptable foliage (suitable for larval development) and larval consumption values. Larval densities varied dramatically due to fluctuating resource availability and ranged from 830 946 larvae/ha in July–August to >4.5 million larvae/ha in January–February. The expected population density for a generalized (mixed sized) tree distribution was estimated to be 13.9 million larvae/ha. Larval densities increased to 18.8 million larvae/ha per year when large trees (>20 cm diameter at breast height) represented the dominant size class, whereas habitats dominated by medium (10–19 cm) and small (1.3–9 cm) trees were predicted to have larval densities of 11.3 and 6.3 million/year, respectively. Validation data obtained from smaller, more accessible plants suggested that the model overestimated realized yearly larval densities by 9% or 15 416 larvae/ha per year.     

Pre-treatment with salicylic acid induces phenolic responses of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis>) bark to bark beetle (<Emphasis Type="Italic">Ips typographus</Emphasis>) attack

Keymessage

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.

Abstract

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.