Resistance level in Scots pine clones and artificial introductions of Tomicus piniperda (Col., Scolytidae) and Leptographium wingfieldii (Deuteromycetes)

Scots pine trees belonging to clones considered to be resistant or susceptible after mass inoculations of Leptographium wingfieldii , a fungus associated with Tomericus piniperda , were submitted to a low density of fungal inoculations or beetle insertions. The length of the phloem-induced reaction zone, the fungal extension and fungal density inside the reaction zone, and the length of maternal galleries of T. piniperda were measured. The length of the phloem reaction zone never differed between resistant and susceptible trees. The density of the fungus mycelium in histological sections of the phloem, and the length of the maternal galleries in the reaction zone were higher in the susceptible trees than in the resistant ones. These factors could be good indicators of resistance of Scots pine to mass attacks.     

Oviposition behavior and reproductive success of the cerambycid Acanthocinus aedilis in the presence and absence of the bark beetle Tomicus piniperda

The influence of Tomicus piniperda (L.)(Col.:Scolytidae) attacks on Acanthocinus aedilis (L.)(Col.:Cerambycidae) (1) oviposition behavior, (2) breeding material preference and (3) reproductive success was investigated in caged pine bolts with and without egg galleries of T. piniperda. In addition, A. aedilis oviposition behavior was studied in the field. In the presence of T. piniperda attacks most A. aedilis eggs were laid through the entrance holes of the bark beetle egg galleries. In the absence of bark beetle attacks most A. aedilis eggs were laid at places where the outer bark had been damaged and the phloem was exposed. In cages, A. aedilis preferred to oviposit in bolts with bark beetle attacks. Thus, A. aedilis oviposited in all 17 of bark beetle attacked bolts but in only four of nine bolts without T. piniperda attacks. Neither the number of A. aedilis offspring nor the body size of emerging adults differed significantly between bolts attacked by bark beetles and unattacked bolts. Nor was there any significant relationship between the density of bark beetle egg galleries and the production of A. aedilis offspring.     

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.     

Suitability of some southern and western pines as hosts for the pine shoot beetle, Tomicus piniperda (Coleoptera: Scolytidae)

The pine shoot beetle, Tomicus piniperda (L.), is an exotic pest that has become established in North America. Discovered in Ohio in 1992, it has since been found in at least 13 states and parts of Canada. The beetle can cause significant growth loss in pines, and it represents a potential threat to trees in areas where it has not yet become established. To evaluate this threat to native pines, field and laboratory tests were conducted on several common and important southern and western species to determine whether they are acceptable hosts for T. piniperda. Comparisons with Pinus sylvestris L., Scots pine, a preferred natural host for the beetle, were made where possible. Measurements of beetle attack success on southern pine billets showed that Pinus taeda L., Pinus echinata Miller, Pinus elliottii var. elliottii Engelmann, Pinus palustris Miller, and Pinus virginiana Miller (loblolly, shortleaf, slash, longleaf, and Virginia pine, respectively) and two western pines, Pinus ponderosa Lawson and Pinus contorta Douglas (ponderosa and lodgepole pine, respectively), were acceptable for breeding material, but brood production was highly variable. Among the southern pines, P. taeda and P. echinata were susceptible to shoot feeding by T. piniperda, whereas P. elliottii was highly resistant and P. palustris seemed to be virtually immune. Shoot feeding tests on the western pines were conducted only in the laboratory, but there was moderate-to-good survival of adults feeding on both species. It seems that if T. piniperda is introduced into the south and west it will likely establish and may cause some damage to native pines. P. taeda may be affected more than other southern pines because it is the most abundant species, it is readily attacked for brood production, which can result in moderately large broods, and the beetle survives well during maturation feeding on P. taeda shoots.     

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。     

Attraction of the bark beetle Tomicus piniperda to Scots pine trees in relation to tree vigor and attack density

The attraction of Tomicus piniperda (L.) (Coleoptera: Scolytidae) to Scots pine trees of low and higher vigor with and without previous attacks was monitored with sticky traps. Somewhat higher numbers of beetles were caught on low vigor trees than on trees of higher vigor, indicating differences in olfactory stimuly between the two classes of trees.Many more beetles were caught on trees attacked by T. piniperda and on trees with simulated bark beetle galleries (manually drilled holes) than on control trees with no or only a few attacks. The strong attraction of beetles to attacked trees is attributed to the beetles responding to host volatiles released from the galleries.

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Zusammenfassung Der Anflug von Tomicus piniperda (L.) (Coleoptera: Scolytidae) an Kiefern in schlechtem oder besserem Gesundheitszustand, mit oder ohne vorherigen Befall, wurde mit Hilfe von Leimfallen untersucht. An Bäumen in schlechtem Zustand wurden etwas mehr Käfer gefangen als an Bäumen in besserem Gesundheitszustand. Das deutet auf Unterschiede in Geruchsreizen zwischen den beiden Klassen des Baumzustands hin.An Kiefern, die von T. piniperda befallen waren oder simulierte Einbohrungen (von Hand gebohrte Löcher) hatten, wurden viel mehr Käfer gefangen als an Kontrollbäumen ohne oder mit sehr geringem Befall. Der starke Anflug von Käfern an befallene Bäume wird gedeutet als Reaktion auf Duftstoffe des Wirts, die aus den Borkenkäfergängen abgegeben werden.

     

Mountain pine beetle attack associated with low levels of 4-allylanisole in ponderosa pine

Mountain pine beetle (Dendroctonus ponderosae) is the most important insect pest in southern Rocky Mountain ponderosa pine (Pinus ponderosa) forests. Tree mortality is hastened by the various fungal pathogens that are symbiotic with the beetles. The phenylpropanoid 4-allylanisole is an antifungal and semiochemical for some pine beetle species. We analyzed 4-allylanisole and monoterpene profiles in the xylem oleoresin from a total of 107 trees at six sites from two chemotypes of ponderosa pine found in Colorado and New Mexico using gas chromatography-mass spectroscopy (GC-MS). Although monoterpene profiles were essentially the same in attacked and nonattacked trees, significantly lower levels of 4-allylanisole were found in attacked trees compared with trees that showed no evidence of attack for both chemotypes.     

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.     

Mountain Pine Beetles Use Volatile Cues to Locate Host Limber Pine and Avoid Non-Host Great Basin Bristlecone Pine

The tree-killing mountain pine beetle (Dendroctonus ponderosae Hopkins) is an important disturbance agent of western North American forests and recent outbreaks have affected tens of millions of hectares of trees. Most western North American pines (Pinus spp.) are hosts and are successfully attacked by mountain pine beetles whereas a handful of pine species are not suitable hosts and are rarely attacked. How pioneering females locate host trees is not well understood, with prevailing theory involving random landings and/or visual cues. Here we show that female mountain pine beetles orient toward volatile organic compounds (VOCs) from host limber pine (Pinus flexilis James) and away from VOCs of non-host Great Basin bristlecone pine (Pinus longaeva Bailey) in a Y-tube olfactometer. When presented with VOCs of both trees, females overwhelmingly choose limber pine over Great Basin bristlecone pine. Analysis of VOCs collected from co-occurring limber and Great Basin bristlecone pine trees revealed only a few quantitative differences. Noticeable differences included the monoterpenes 3-carene and D-limonene which were produced in greater amounts by host limber pine. We found no evidence that 3-carene is important for beetles when selecting trees, it was not attractive alone and its addition to Great Basin bristlecone pine VOCs did not alter female selection. However, addition of D-limonene to Great Basin bristlecone pine VOCs disrupted the ability of beetles to distinguish between tree species. When presented alone, D-limonene did not affect behavior, suggesting that the response is mediated by multiple compounds. A better understanding of host selection by mountain pine beetles could improve strategies for managing this important forest insect. Moreover, elucidating how Great Basin bristlecone pine escapes attack by mountain pine beetles could provide insight into mechanisms underlying the incredible longevity of this tree species.     

Physical and chemical response of juvenile Acacia tortilis trees to browsing. Experimental evidence

1. Changes in nutritional value and accessibility of leaves following browsing are important in the dynamics of plant–herbivore interactions because they influence the fitness of the plant attacked and the future utilization of it by the herbivore.
2. Hand pruning of Acacia tortilis , a spinescent tree common in savanna ecosystems of eastern Africa, resulted in higher biomass of spines and new shoots in pruned trees than in unpruned controls.
3. Pruned trees allocated a higher proportion of shoot biomass to spines than unpruned ones, whereas the proportion of leaf biomass in new shoots was slightly reduced. Because increases in spine biomass and density following pruning are coupled with an increase in shoot production, it is concluded that higher production of spines is an inducible response of Acacia tortilis to pruning.
4. No significant changes in the concentration of total phenolics, condensed tannins or leaf nitrogen were induced by pruning.
5. Irrespective of treatment, high foliar concentrations of nitrogen were correlated with an increase in twig production for a given leaf biomass and a reduction in the concentration of secondary substances in leaves. This relation may lead to a conflict between foraging efficiency and nutrition for browsers of A. tortilis.     

Alterations of the resin canal system of <Emphasis Type="Italic">Pinus pinaster</Emphasis> seedlings after fertilization of a healthy and of a <Emphasis Type="Italic">Hylobius abietis</Emphasis> attacked stand

Changes in resource availability and biotic and abiotic stress may alter the defensive mechanisms of pine trees. The effect of fertilisation on the resin canal structure of Pinus pinaster seedlings established in two trials in NW Spain, one attacked by Hylobius abietis and the other non-attacked, was studied. The leaders of 50 plants were destructively sampled and the resin canal density, the canal area and its relative conductive area in the phloem and xylem were assessed. Experimentally increased nutrient availability significantly decreased resin canal density in the phloem of the seedlings in the two analysed trials, where unfertilised seedlings presented up to 30% more resin canal density than the fertilised seedlings (mean value ± SEM = 0.32 ± 0.02 resin canals mm⁻² in the fertilised plants versus 0.45 ± 0.04 resin canals mm⁻² in the control plants). Fertilisation had no effect on the resin canal system in the xylem, but significantly increased tracheid size. Significant differences of resin canals among sites were observed mainly in the xylem; the resin canal density was 1.7-fold greater in the attacked site than in the non-attacked site. The similar structure of phloem resin canals in both sites supports that phloem resin canals are constitutive mechanisms of defence in P. pinaster, whereas xylem resin canals would be constitutive mechanisms but also inducible mechanisms of resistance following the attack of pine weevils or bark beetles. XM and LS equally contributed to this paper.     

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.     

Winter starch reserves of white oak as a predictor of attack by the twolined chestnut borer,Agrilus bilineatus (Weber) (Coleoptera: Buprestidae)

Summary The twolined chestnut borer, Agrilus bilineatus (Weber) (Coleoptera: Buprestidae), attacks oaks (Quercus spp.) and is associated with extensive mortality of trees in the eastern deciduous forests of North America. We tested the hypothesis that winter starch reserves of oak roots are an indicator of tree vigor and that only trees low in stored starch would be attacked by A. bilineatus. We measured the levels of stored starch in the roots of 200 non-infested healthy white oaks during the dormant season and determined their correlation with A. bilineatus attacks the following spring. There was a significant increase in A. bilineatus captures on sticky traps with a decrease in winter starch reserves. Trees low in stored starch that were also stressed by phloem-girdling attracted 3.7 times as many beetles as did non-girdled trees that were low in starch. However, non-girdled trees that had low winter starch reserves were also attacked. Only oaks that had had extremely low winter root starch reserves (<5mg/g dry weight of root sapwood tissue) were heavily attacked by A. bilineatus and subsequently died. One third of non-girdled low starch trees and 67% of phloem-girdled low starch trees died, whereas none of the trees with root starch >5 mg/g dry wt died. These results indicate that winter starch reserves are a good predictor of A. bilineatus attack.The investigation reported in this paper (No. 87-7-8-118) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director     

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

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Host resistance in defoliated pine: effects of single and mass inoculations using bark beetle-associated blue-stain fungi

1 In 1996, 7000 ha of pine forests were defoliated by the pine looper Bupalus piniaria in south‐western Sweden. 2 The susceptibility of trees of different defoliation classes (0, 30, 60, 90 and 100% defoliation) to beetle‐vectored blue‐stain fungi was tested in inoculation experiments. Forty and 120‐year‐old Scots pine trees were inoculated with ‘single’, i.e. a few inoculations of Leptographium wingfieldii and Ophiostoma minus, two blue‐stain fungi associated with the pine shoot beetle Tomicus piniperda. The young trees were also ‘mass’ inoculated with L. wingfieldii at a density of 400 inoculation points per m² over a 60 cm stem belt. 3 Host tree symptoms indicated that only trees with 90–100% defoliation were susceptible to the mass inoculation. 4 Single inoculations did not result in any consistent differences in fungal performance between trees of different defoliation classes, regardless of inoculated species or tree age class. 5 Leptographium wingfieldii produced larger reaction zones than O. minus, and both species produced larger lesions in old than in young trees. 6 As beetle‐induced tree mortality in the study area occurred only in totally defoliated stands, mass inoculations seem to mimic beetle‐attacks fairly well, and thus seem to be a useful tool for assessing host resistance. 7 As even severely defoliated pine trees were quite resistant, host defence reactions in Scots pine seem to be less dependent on carbon allocation than predicted by carbon‐based defence hypotheses.     

为害云南松的纵坑切梢小蠹 (Tomicus piniperda)经济阈值的研究(英文)

经过对纵坑切梢小蠹 (TomicuspiniperdaL .)种群密度和云南松 (PinusyunnanensisFranch .)生长状况的长期、仔细的调查 ,加上改进的回归分析方法 ,得出了为害云南松的纵坑切梢小蠹防治上的经济阈值 :当林木枯死率不超过 0 .5 2 %时 ,自然控制因子能控制纵坑切梢小蠹种群不向高密度方向发展 ,因而可以不必防治。当林木枯死率达到 0 .5 0 % ,对应的枯捎率为 0 .6 1 % ,以及对应的平均每株侵入孔数为 0 .6 6时 ,防治费用和挽回损失价值相当。所以 ,若既考虑到经济上的合理性 ,又考虑到控制其种群不向高密度方向发展 ,当林木枯死率超过 0 .5 % ,对应的枯梢率超过 0 .6 1 % ,以及对应的平均每株侵入孔数超过 0 .6 6时 ,就有了进行防治的必要。本文还给出了各云南松林区可以用来根据自己的具体情况计算出适合自己的经济阈值的公式。     

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     

The bark beetle Pityogenes chalcographus (L.) (Scolytidae) in living trees: reproductive success, tree mortality and interaction with Ips typographus

Abstract:  Tree-killing ability of Pityogenes chalcographus was experimentally investigated by baiting living spruce Picea abies with synthetic aggregation pheromone, thereby exposing them to beetle attack. Reproductive success was estimated by rearing beetles from stem sections taken from killed trees. The presence of Ips typographus on several killed trees allowed a comparison of reproductive success in the presence and absence of this species. Furthermore, the possibility that breeding material colonized by P. chalcographus may induce subsequent attacks by I. typographus on nearby trees was also studied. The tree killing ability of P. chalcographus was low as only 8% of the baited trees, attacked by P. chalcographus alone, were killed. Tree death was strongly related to the presence of I. typographus , which is in line with the association with aggressive wood-living fungi in this species. The reproductive success was low, with on average less than one daughter beetle produced per mother beetle. The success was significantly higher in trees also inhabited by I. typographus . Tree-parts colonized by P. chalcographus induced almost no attacks by I. typographus on nearby trees.     

Anatomical defences against bark beetles relate to degree of historical exposure between species and are allocated independently of chemical defences within trees

Conifers possess chemical and anatomical defences against tree‐killing bark beetles that feed in their phloem. Resins accumulating at attack sites can delay and entomb beetles while toxins reach lethal levels. Trees with high concentrations of metabolites active against bark beetle‐microbial complexes, and more extensive resin ducts, achieve greater survival. It is unknown if and how conifers integrate chemical and anatomical components of defence or how these capabilities vary with historical exposure. We compared linkages between phloem chemistry and tree ring anatomy of two mountain pine beetle hosts. Lodgepole pine, a mid‐elevation species, has had extensive, continual contact with this herbivore, whereas high‐elevation whitebark pines have historically had intermittent exposure that is increasing with warming climate. Lodgepole pine had more and larger resin ducts. In both species, anatomical defences were positively related to tree growth and nutrients. Within‐tree constitutive and induced concentrations of compounds bioactive against bark beetles and symbionts were largely unrelated to resin duct abundance and size. Fewer anatomical defences in the semi‐naïve compared with the continually exposed host concurs with directional differences in chemical defences. Partially uncoupling chemical and morphological antiherbivore traits may enable trees to confront beetles with more diverse defence permutations that interact to resist attack.