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.     

Response of the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae), to conophthorin and other angiosperm bark volatiles in the avoidance of non-hosts

1 Seventeen non‐host angiosperm bark volatiles, seven of which are antennally active to Ips pini (Say), the pine engraver (PE), were tested for their ability to disrupt the response of the PE to pheromone‐baited traps. 2 Four green leaf volatiles (GLVs) were tested [1‐hexanol (Z)‐3‐hexen‐1‐ol, hexanal, and (E)‐2‐hexenal]. None had any disruptive effect singly, as a group or in all possible blends based on functional groups, despite the fact that the two aldehydes were antennally active. These compounds may have, in some instances, actually masked the disruptive effect of other compounds. The PE thus differs in its response from other Scolytidae, including other Ips spp. 3 Eight non‐host volatiles that were antennally active to other bark beetles, but not to PEs, had no disruptive effect, validating the use of coupled gas chromatographic‐electroantennographic detection analyses to detect compounds with potential behavioural activity. 4 The bicyclic spiroacetal conophthorin, (E)‐7‐methyl‐1,6‐dioxaspiro[4.5]decane, was disruptive when tested alone. When blends of two aldehydes [salicylaldehyde and nonanal] plus an alcohol and a phenol [benzyl alcohol and guaiacol] were combined with conophthorin, an enhanced disruptive effect was revealed. No single compound, other than conophthorin, disrupted the pheromone‐positive response and no blend that did not contain conophthorin was consistently disruptive to both sexes. Conophthorin seems to be a critical component in the non‐host angiosperm message for I. pini during its host selection phase. 5 Combination of the repellent synomones, verbenone and ipsenol, with the five disruptive non‐host volatiles may provide a potent treatment to protect trees, logs or stands from attack by the PE.     

Incomplete barriers to mitochondrial gene flow between pheromone races of the North American pine engraver,Ips pini (Say) (Coleoptera,Scolytidae)

The pine engraver Ips pini (Say) is known to include three pheromone races, but gene flow between these races has not been investigated. We used maternally inherited mitochondrial DNA (mtDNA) variation to infer gene flow between 22 widely distributed North American populations of I. pini for a total of 217 individuals, based on 354 bp of the cytochrome oxidase I gene. Gene flow was estimated cladistically as migrants per generation (Nm) and as haplotype variation between populations (N_st). Three distinct mtDNA haplotype lineages, generally corresponding to eastern (I), Rocky Mountain (II) and western (III) regions of North America, were resolved with a total of 34 distinct I. pini haplotypes. The distributions of these lineages were largely congruent with the geographical ranges of the ''New York'', ''California'' and ''Idaho–Montana'' pheromone races. Only individuals with lineage I mtDNA were observed among eastern populations, whereas individuals with lineage II or III mtDNA predominated among western populations. Gene flow (Nm and N_st) was generally moderate between all populations. However, the presence of lineage I mtDNA on the eastern side of western North America and the absence of lineage II and III mtDNA in eastern North America suggest directional gene flow from east to west. These results indicate that female-controlled assortative mating among pheromone races may disrupt gene flow between conspecifics, reflecting incomplete pre-mating barriers.     

Effect of host tree seasonal phenology on substrate suitability for the pine engraver (Coleoptera: Scolytidae): implications for population dynamics and enemy free space

This study evaluated the effects of seasonal phenology on the substrate quality of susceptible hosts to the pine engraver, Ips pini (Say). We also determined the effects of the duration and method of storage on host quality for purposes of laboratory rearing. Live red pine trees were felled at various times during the season, and I. pini adults from a laboratory colony were established on the logs. Subsamples of logs were stored for various intervals, and then provided to beetles. Subsamples of stored logs were waxed at both ends to prevent water loss before being submitted to the same assays. Suitability of red pine phloem tissue in susceptible hosts declined for I. pini throughout the growing season. As the season progressed, the number of beetle progeny that emerged from colonized hosts dropped substantially. This decline was associated with simultaneous reductions in phloem moisture content. Reduction in host suitability may partially offset any advantage I. pini may gain from colonizing trees after the major predators have become less abundant. Bark beetle brood production decreased significantly with length of storage, regardless of the month of tree felling or the method of storing. Implications for bark beetle population dynamics and laboratory rearing systems are discussed.     

Efficacy of systemic insecticides for protection of loblolly pine against southern pine engraver beetles (Coleoptera: Curculionidae: Scolytinae) and wood borers (Coleoptera: Cerambycidae)

We evaluated the efficacy of the systemic insecticides dinotefuran, emamectin benzoate, fipronil, and imidacloprid for preventing attacks and brood production of southern pine engraver beetles (Coleoptera: Curculionidae: Scolytinae) and wood borers (Coleoptera: Cerambycidae) on standing, stressed trees and bolt sections of loblolly pine, Pinus taeda L., in eastern Texas. Emamectin benzoate significantly reduced the colonization success of engraver beetles and associated wood borers in both stressed trees and pine bolt sections. Fipronil was nearly as effective as emamectin benzoate in reducing insect colonization of bolts 3 and 5 mo after injection but only moderately effective 1 mo after injection. Fipronil also significantly reduced bark beetle-caused mortality of stressed trees. Imidacloprid and dinotefuran were ineffective in preventing bark beetle and wood borer colonization of bolts or standing, stressed trees. The injected formulation of emamectin benzoate was found to cause long vertical lesions in the sapwood-phloem interface at each injection point.     

Longevity of needle fascicles of Pinus longaeva (Bristlecone pine) and other North American pines

Summary Variation in leaf longevity of gymnosperms has received surprisingly little attention despite its likely adaptive significance. Pinus longaeva, a pine of arid, subalpine environments in the western United States, has the record among conifers for needle longevity, with a maximum dwarf shoot (needle fascicle) retention time of up to about 45 years. Most low elevation pines have dwarf shoot retention times (DSRs) of two to four years. Literature data for the 37 species of Pinus native to the United States and Canada and field data for eight taxa (21 populations) of pines growing at various elevations in California each show a strong positive correlation between elevation and DSR, respectively, r=+0.65, df=35, p<0.001 and r=+0.82; df=19, p<0.001. We propose that extended needle fascicle longevity represents an adaptation to arid and especially high elevation environments. Field data from native stands and common gardens indicate that differences between taxa in DSR relate to both genetic and environmental factors. When grown at the same sites certain species (eg, P. longaeva, P. monophylla) had much longer DSRs than others, indicating a genetic basis for differences in needle fascicle longevity. For six of seven taxa that were each studied at more than one elevation there was a statistically significant increase in DSR in going from the lowest to the highest elevation site, indicating strong environmental control of needle fascicle longevity.The physiological control of dwarf shoot senescence and abscission is poorly understood. For P. longaeva dwarf shoots of a particular age class are not shed simultaneously; rather there is a more or less gradual attrition of dwarf shoots from the long shoot. Although different types of long shoots of pines are known to differ physiologically, for P. longaeva there was no consistent difference in DSR on various types of lateral long shoots (eg, vegetative, pollen cone-bearing, seed cone-bearing), nor was there a statistically significant difference in DSR on trunks versus on their lateral long shoots. In addition, for P. contorta ssp. bolanderi and P. muricata needle fascicle longevity was not affected by the degree of edaphically induced dwarfing (ie, stunting) of the trees.     

Limonene: attractant kairomone for white pine cone beetles (Coleoptera: Scolytidae) in an eastern white pine seed orchard in western North Carolina

I report on the attraction of the white pine cone beetle, Conophthoru.s coniperda (Schwarz) (Coleoptera: Scolytidae), to traps baited with the host monoterpene limonene in western North Carolina. Both (+)- and (-)-limonene attracted male and female cone beetles to Japanese beetle traps in an eastern white pine, Pinus strobus L., seed orchard near Murphy, NC. Catches of cone beetles were directly proportional to the release rate of (-)-limonene; (+)-limonene was not tested for dose response. Attraction of cone beetles to the pheromone (+/-)-trans-pityol was increased significantly by both enantiomers of limonene. In all experiments, catches of C. coniperda were strongly male biased with no treatment effect on sex ratio. (- )-Limonene had no effect on trap catches of the predator Enoclerus nigripes (Say) to pityol, whereas (+)-limonene interrupted the attraction of E. nigripes to traps baited with pityol. Of six monoterpenes commonly found in white pine cones, only (-)-alpha-pinene elicited attraction of E. nigripes to Japanese beetle traps.     

Evaluation of insecticides for protecting Southwestern ponderosa pines from attack by engraver beetles (Coleoptera: Curculionidae: Scolytinae)

Insecticides that might protect pine trees from attack by engraver beetles (Ips spp.) have not been rigorously tested in the southwestern United States. We conducted two field experiments to evaluate the efficacy of several currently and potentially labeled preventative insecticides for protecting high-value ponderosa pine, Pinus ponderosa Dougl ex. Laws., from attack by engraver beetles. Preventative sprays (0.19% permethrin [Permethrin Plus C]; 0.03, 0.06, and 0.12% bifenthrin [Onyx]; and 1.0 and 2.0% carbaryl [Sevin SL] formulations) and systemic implants (0.875 g per capsule acephate [Acecap] and 0.650 g per capsule dinotefuran) were assessed on bolts (sections of logs) as a surrogate for live trees for a period of 13 mo posttreatment. The pine engraver, Ips pini (Say), was the most common bark beetle found attacking control and treated bolts, but sixspined ips, Ips calligraphus (Germar), and Ips lecontei Swain also were present. After approximately 13 mo posttreatment in one experiment, the spray treatments with 2.0% carbaryl, 0.19% permethrin, and 0.06 or 0.12% bifenthrin prevented Ips attack on the bolts at a protection level of > or = 70%. The acephate and dinotefuran systemic insecticides, and the 0.03% bifenthrin spray, provided inadequate (< or = 36%) protection in this experiment. For the other experiment, sprayed applications of 1.0% carbaryl, 0.19% permethrin, and 0.06% bifenthrin prevented beetle attack at protection levels of > or = 90, > or = 80, and > or = 70%, respectively, when bolts were exposed to Ips beetle attack for approximately 9-15 wk posttreatment. The sprays with 0.19% permethrin and 0.06% bifenthrin also provided > or = 90% protection when bolts were exposed for approximately 15-54 wk posttreatment. We concluded that under the conditions tested, 1.0 and 2.0% carbaryl, 0.19% permethrin, and 0.06 and 0.12% binfenthrin were acceptable preventative treatments for protecting ponderosa pine from successful engraver beetle attack for one entire flight season in the U.S. Southwest.     

Feeding behaviour of the black pine beetle,Hylastes ater (Coleoptera: Scolytidae)

1 The feeding behaviour of Hylastes ater was investigated at 15 °C and 20 °C on a range of tree species. The role of an aggregation pheromone and the part played by olfactory cues in food selection was also investigated. 2 A distinct hierarchical preference at both 15 °C and 20 °C such that Pinus radiata > Pinus sylvestris > > Picea abies > > Rubus fruticosus=Betula pendula was shown. 3 Feeding on P. sylvestris led to the highest weight gain, closely followed by P. radiata and P. abies. Slightly elevated feeding was observed at 20 °C compared with 15 °C. 4 Reduced feeding in the presence of B. pendula suggests the possibility of an antifeedant effect and predamaged wood was found to be less nutritious, possibly due to greater plant defences. 5 The role of an aggregation pheromone was not supported. Olfactory cues enabled the detection of woody hosts, although differentiation between host was not observed. 6 These results are likely to be important in helping to design an effective integrated control approach against this pest.     

Frass clearing by male pine engraver beetles (Ips pini; Scolytidae): paternal care or paternity assurance

Male parental care and paternity assurance are often associatedwith long-duration pair bonds. The mating system of the pineengraver beetle, Ips pini, includes an association between themale and female that persists for most of the prolonged oppositionperiod. The male beetles remove frass that arnmmlatn as thefemales lay their eggs in die phloem tissue of the host tree.Experiments and field observations were done to test possiblebenefits to males that stay in the galleries removing frasswhile die females are ovipositing. Two hypotheses were thatclearing frass (1) provides some form of care that results inmore offspring being produced and (2) is part of a paternityassurance mechanism. Male removal experiments in the field producedno evidence that male presence significantly influenced anyof five measures of offspring production. Laboratory experimentsin which virgin females were bred reciprocally to sterile andfertile males showed that, while there is no strong patternof last-male pr, last-male paternity does increase over time.Field observations revealed that female pine engravers oftencarry sperm from previous maringi when they solicit entry toa male's breeding gallery. The pattern of paternity and thefemale's sperm storage capacity suggest that males must maintainprolonged mating access to females in order to ensure high paternity.Hence, frass clearing is necessary to maximize paternity     

Responses of single receptor cells in the pine engraver beetle,Ips pini (SAY) (Coleoptera: Scolytidae) to its aggregation pheromone,ipsdienol, and the aggregation inhibitor,ipsenol

Summary Responses of single olfactory cells on the antennal club ofIps pini have been recorded electrophysiologically. The majority of cells were strongly activated by either/or of the two behavior-modifying chemicals, the aggregation pheromone ipsdienol and the aggregation inhibitor ipsenol. Simultaneous stimulation with these terpence alcohols showed that ipsenol had no inhibitory effect on the receptor responses to ipsdienol. It appears that the behavioral inhibition by ipsenol on the attraction of ipsdienol results from central integration of the information from separate receptor cells rather than blockage at the peripheral receptors.This work was carried out at the College of Environmental Sciences and Forestry, SUNY, Syracuse. We are grateful to Professor R.M. Silverstein and his staff for provision of purified compounds and laboratory facilities. We also acknowledge with thanks the provision of equipment and laboratory facilities by Professor D. Tapper, College of Veterinary Medicine, Cornell University.     

Tomicus piniperda (Coleoptera: Scolytidae) shoot-feeding characteristics and overwintering behavior in Scotch pine Christmas trees

Overwintering behavior of Tomicus piniperda (L.) was studied in a Scotch pine (Pinus sylvestris L.) Christmas tree plantation in Indiana (1992-1994) and a plantation in Michigan (1994). In general, adults feed inside shoots during summer, then move to overwintering sites at the base of trees in autumn. In early autumn, adults were most often found in shoot-feeding tunnels that were still surrounded by green needles, whereas few were in tunnels surrounded by yellow or brown needles. For all years and sites combined, the range in the percentage of recently tunneled shoots that contained live T. piniperda adults decreased from 89 to 96% in mid-October, to 15- 66% in early November, to 2-10% in mid-November, and to 0-2% by late November to early December. In each year, the first subfreezing temperatures in autumn occurred in October, before most adults left the shoots. Of 1,285 T. piniperda-tunneled shoots, one to seven tunnels (mean = 1.6) and zero to three adults were found per infested shoot. Of these 1,285 attacked shoots, 55% of the shoots had one tunnel, 33% had two, 9% had three, 3% had four, and <1% had five to seven tunnels each. When two or more tunnels occurred in a single shoot, adults were most commonly found in the innermost (most basal) tunnel. For the 2,070 tunnels found in the 1,285 shoots, average shoot thickness at the tunnel entrance was 6.0 mm, average distance from the tunnel entrance to the shoot tip was 6.3 cm, and average tunnel length was 2.3 cm. Four Scotch pine Christmas trees were dissected in January 1993. Eighty percent of the tunneled shoots were in the upper quarter of the tree crown and 98% were in the upper half. For the four trees inspected in January, one live adult was found in a shoot and 85 adults were found in the outer bark along the lower trunk from 1 cm below the soil line to 19 cm above the soil line. No overwintering adults were found outside the trunk in the duff or soil near the base of each test tree. Implications of these results are discussed in terms of surveying, timing the cutting of Christmas trees, and cutting height for Christmas trees.     

Interactions of Hylastes species (Coleoptera: Scolytidae) with Leptographium species associated with loblolly pine decline

Hylastes spp. (Coleoptera: Scolytidae) were evaluated as potential vectors of Leptographium spp. fungi. Bark beetles were trapped from stands ofloblolly pine, Pinus taeda L., exhibiting a range of decline symptoms in central Alabama. Under controlled conditions, field-collected adult Hylastes salebrosus Eichoff (Coleoptera: Scolytidae) and Hylastes tenuis Eichoff (Coleoptera: Scolytidae), which had been surface-sterilized and inoculated with Leptographium terebrantis Barras & Perry and Leptographium serpens (Goid.) Wingfield, transmitted the fungi into 100% of wounded and unwounded loblolly root sections with which they were confined. None of the sterilized and uninoculated beetles transmitted any Leptographium spp. to roots. Significantly more H. salebrosus and H. tenuis brood emerged from roots infected with Leptographium species than from sterile roots, indicating an enhancement of Hylastes reproduction.     

Evaluation of funnel traps for characterizing the bark beetle (Coleoptera: Scolytidae) communities in ponderosa pine forests of north-central Arizona

Lindgren funnel traps baited with aggregation pheromones are widely used to monitor and manage populations of economically important bark beetles (Coleoptera: Scolytidae). This study was designed to advance our understanding of how funnel trap catches assess bark beetle communities and relative abundance of individual species. In the second year (2005) of a 3-yr study of the bark beetle community structure in north-central Arizona pine (Pinus spp.) forests, we collected data on stand structure, site conditions, and local bark beetle-induced tree mortality at each trap site. We also collected samples of bark from infested (brood) trees near trap sites to identify and determine the population density of bark beetles that were attacking ponderosa pine, Pinus ponderosa Douglas ex Lawson, in the area surrounding the traps. Multiple regression models indicated that the number of Dendroctonus and Ips beetles captured in 2005 was inversely related to elevation of the trap site, and positively associated with the amount of ponderosa pine in the stand surrounding the site. Traps located closer to brood trees also captured more beetles. The relationship between trap catches and host tree mortality was weak and inconsistent in forest stands surrounding the funnel traps, suggesting that trap catches do not provide a good estimate of local beetle-induced tree mortality. However, pheromone-baited funnel trap data and data from gallery identification in bark samples produced statistically similar relative abundance profiles for the five species of bark beetles that we examined, indicating that funnel trap data provided a good assessment of species presence and relative abundance.     

Flight of the Chinese white pine beetle (Coleoptera: Scolytidae) in relation to sex, body weight and energy reserve

The flight distance, flight time and individual flight activities of males and females of Dendroctonus armandi were recorded during 96-h flight trials using a flight mill system. The body weight, glucose, glycogen and lipid content of four treatments (naturally emerged, starved, phloem-fed and water-fed) were compared among pre-flight, post-flight and unflown controls. There was no significant difference between males and females in total flight distance and flight time in a given 24-h period. The flight distance and flight time of females showed a significant linear decline as the tethered flying continued, but the sustained flight ability of females was better than that of males. The females had higher glycogen and lipid content than the males; however, there was no significant difference between both sexes in glucose content. Water-feeding and phloem-feeding had significant effects on longevity, survival days and flight potential of D. armandi, which resulted in longer feeding days, poorer flight potential and lower energy substrate content. Our results demonstrate that flight distances in general do not differ between water-fed and starved individuals, whereas phloem-fed females and males fly better than water-fed and starved individuals.     

Prairie grasses as hosts of the northern corn rootworm (Coleoptera: Chrysomelidae)

We evaluated 27 prairie grass species thought to be among those dominant 200 yr ago in the northern midwest as larval hosts of the northern corn rootworm, Diabrotica barberi Smith and Lawrence. Maize (Zea mays L.), spring wheat (Triticum aestivum L.), and grain sorghum (Sorghum bicolor L.) were included as controls for a total of 30 species. Twenty pots of each species were planted in a randomized complete block design. Each pot was infested 5 wk later with 20 neonate northern corn rootworm larvae. Two pots within each species and block were assigned an extraction date of 7 or 14 d after infestation. The remaining two pots from each block were used to monitor adult emergence. The percentage of larvae recovered, change in larval head capsule width, and change in average dry weights varied significantly among the grass species. The highest percentage of larvae was recovered from slender wheatgrass, Elymus trachycaulus (Link), and this was significantly greater than the percentage recovered from all other species including maize for the 14-d sample date. Several additional species were also relatively good hosts, in that the percentage of larvae recovered from these species was not significantly different from maize. The average dry weight of larvae recovered was significantly greater for larvae recovered from maize than for larvae recovered from all other species except slender wheatgrass, when the two samples dates were combined. Overall, adults were produced from only 6 of the 28 species evaluated, and no analysis was performed because of the low numbers. The results of this study are discussed in relation to the potential of alternate hosts of northern corn rootworm to serve as a bridge to survival on transgenic maize.     

Descriptive biogeography of Tomicus (Coleoptera: Scolytidae) species in Spain

Aim, location Tomicus (Coleoptera, Scolytidae) species are some of the principal pests of Eurasian forest and are represented by three coexisting species in Spain, Tomicus piniperda (Linnaeus, 1758), Tomicus destruens (Wollaston, 1865) and Tomicus minor (Harting, 1834). The distribution of two taxa are unknown as they have until recently been considered separate species. Therefore, we model the potential distribution centres and establish the potential distribution limits of Tomicus species in Iberia. We also assess the effectiveness of different models by comparing predicted results with observed data. These results will have application in forest pest management. Methods Molecular and morphological techniques were used to identify species from 254 specimens of 81 plots. For each plot, a Geographical Information System was used to extract a set of 14 environmental (one topographic, six climatic) and biotic variables (seven host tree distributions). General Additive Models and Ecological Niche Factor Analysis models are applied for modelling and predicting the potential distribution of the three especies of Tomicus. Results The results of both modelling methodologies are in agreement. Tomicus destruens is the predominant species in Spain, living in low and hot areas. Tomicus piniperda occurs in lower frequency and prefers wet and cold areas of north‐central Spain. We detected sympatric populations of T. destruens and T. piniperda in Northern coast of Spain, infesting mainly P. pinaster. Tomicus minor is the rarest species, and it occupies a fragmented distribution located in high and wet areas. The remarkable biotic variable is the distribution of P. sylvestris, incorporated into the models of T. destruens and T. piniperda. Main conclusions These results indicate that in wet areas of north‐central Spain where T. piniperda occurs (and possibly the high altitudes of the southern mountains), T. destruens has a climatic distribution limit. In the northern border of this area, both species overlap their distributions and some co‐occurrences were detected. Tomicus minor potentially occurs in high and wet fragmented areas.     

Biochemical approach to the control ofXyleborus fornicatus (Coleoptera: Scolytidae)

Summary The addition of small quantity of potassium acetate to the fertilizer applied to tea bushes was an effective method of reducing infestation by the beetle pest,Xyleborus fornicatus. The mechanism of action of potassium acetate is discussed as being due to its conversion to saponins and/or sterol analogues which interfere with pupation of the beetle larvae.     

A model for the timing of swarming of Tomicus piniperda (Coleoptera: Scolytidae)

Tomicus piniperda is the first bark beetle to swarm in the spring in Fennoscandia. The intensity of swarming is directly related to air temperature above a certain threshold and reaches 100% asymptotically. While temperature sums above the threshold accumulate, swarming rate slows down. A non-linear model was built on these considerations. The model was calibrated to window trap material from northern Finland, and it was validated against published material mainly from Norway. The threshold temperatures in these materials were 10.8°C and 11.6°C, respectively. The predictions of the model were satisfactory, and the model will be tried as a part of an entire population dynamics model. The possibilities to predict not only timing, but also the frequency of attack are discussed.