Host finding in the pine weevil Hylobius abietis: effects of conifer volatiles and added limonene

The attraction of Hylobius abietis (L.) (Coleoptera: Curculionidae) to -pinene was recently shown to be inhibited by limonene. This study shows that added limonene also affects the response of H. abietis to odours from natural host material. The experiments included both pieces of Scots pine suspended in pitfall traps and planted pine seedlings that had either been manually wounded or left intact. By adding a limonene dispenser, the catch of pine weevils in pine-baited traps was reduced by about 80–90% in spring and autumn and by about 40–50% in summer. Limonene substantially reduced the rate at which manually wounded seedlings were attacked by weevils in spring and autumn. For intact seedlings, a limonene-induced reduction in attack rate was found only in autumn. In the absence of limonene, a small wound made on the stem of a seedling increased its probability of being attacked by about fivefold. This figure was estimated by comparing data from all experimental periods with a model assuming a constant ratio of attack rates for the two kinds of seedlings.     

Orientation of the pine weevil Hylobius abietis to underground sources of host volatiles

Adults of Hylobius abietis (L.) (Coleoptera: Curculionidae) were found to locate conifer roots suitable for oviposition by utilizing host volatiles diffusing through the soil. Underground sources of host volatiles were presented to weevils in a laboratory bioassay. A cold-trapping condensate of Scots pine, Pinus sylvestris L., and fractions of it were tested. Various fractions containing host terpenes attracted weevils in the bioassay, but the complete pine condensate caused the highest response. Ethanol was also found to be attractive. Weevils caged underground in the absence of host material did not attract weevils on the surface.

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Zusammenfassung Rüsselkäfer (Hylobius abietis) können auf der Erdoberfläche die Lage im Boden verborgenen Brutmaterials feststellen und Wurzeln senkrecht grabend auffinden. Mit einer neu entwickelten Methode für Laborversuche, die sich auf die spezifische Reaktion des Eingrabens zum Brutmaterial gründet, wurden das Orientierungsverhalten der Käfer und die zum Brutmaterial führenden Geruchsstoffe des Wirtes studiert.Beide Geschlechter von H. abietis reagierten gleichartig mit Eingraben auf die Geruchsstoffe des Wirtes. In Wahlversuchen zwischen gleichwertigen Anlockungspunkten kam häufig Aggregation der Käfer an einem Punkt vor. Die stärkere Ansammlung an einzelnen Stellen stand im Zusammenhang mit dem Vorhandensein einer Erdröhre zur Quelle der Geruchsstoffe. In den folgenden Versuchen wurde deshalb den Tieren jeweils nur eine Geruchsstoffquelle angeboten. Weder Weibchen noch Männchen im Boden übten eine Anziehung auf Käfer an der Oberfläche aus.Kiefernstücke und Kondensate flüchtiger Kieferinhaltsstoffe sowie Kondensat nach Passage durch die Kolonne des Gaschromatographen waren stark attraktiv. Alle durch präparative Gaschromatographie hergestellten Fraktionen des Kondensats waren ebenfalls, aber schwächer attraktiv. Zehnfache Verdünnung des Kondensats und der Fraktionen verminderte ihre Attraktivität mit etwa 40–70%. Auch Äthanol, Methanol und Pentan wurden geprüft; nur Äthanol hatte eine mässig anlockende Wirkung.

     

Feeding by the pine weevil Hylobius abietis in relation to sun exposure and distance to forest edges

Abstract 1 The intensity of feeding by adult pine weevils Hylobius abietis (L.) on the stem bark of Norway spruce Picea abies (L.) Karst. seedlings planted in rows with a north–south orientation across a clear‐cutting, was measured throughout a growth season. The feeding was then correlated to light interception, soil temperature and distance to the nearest forest edge. 2 Feeding was at least twice as intense on seedlings in the central part of the clear‐cutting compared to those at the edges. The decline began approximatety 15 m from the edge and was of similar proportions on both the sun‐exposed and shaded sides. 3 Measures of global radiation and soil temperature correlated well with consumption on the shaded side. However, on the sun‐exposed side, there were no apparent correlations with global radiation or soil temperature that could explain the decline in consumed bark area. 4 We conclude that the decline in feeding towards the forest edges was mainly due to factors other than the microclimate variables we monitored. We suggest that the presence of roots of living trees along the forest edge may reduce damage to seedlings, since they provide an alternative source of food for the weevils. This alternative‐food hypothesis may also explain why seedlings in shelterwoods usually suffer less damage from pine weevils than seedlings in clear‐cuttings.     

Feeding on roots in the humus layer by adult pine weevil, Hylobius abietis

1 The consumption by adult pine weevil, Hylobius abietis, of the bark of roots present in the humus layer was assessed in a field study conducted in southern Sweden during two years (1998 and 2002). The study sites were divided into two areas: (i) a shelterwood where 80–100 mature Scots pine trees per hectare remained after cutting and (ii) a clearcut where no trees were left. 2 On average, 3741 m² per hectare of root bark was present in the humus layer, of which 135 m² was not coniferous but comprised species such as bilberry and broadleaved trees. 3 The mean area debarked by pine weevils was 2.9 m² per hectare; 2.6 m² of conifer roots and 0.3 m² of bilberry roots. Roots of broadleaved trees were almost never consumed. No clear preferences for roots of a specific level of vitality were observed. 4 No consistent difference between the shelterwood and clearcut was found, either in the amount of root bark area available or in the extent of root feeding by pine weevil. 5 A weak negative correlation between debarked areas on roots and seedlings was found, indicating that root feeding may have reduced damage to seedlings. 6 It is concluded that conifer roots in the humus layer constitute a major food source for the pine weevil and can be utilized for a considerable period in both clearcuts and shelterwoods.     

Oviposition patterns of the pine weevil Hylobius abietis

Spatial and temporal distributions of eggs laid by the pine weevil Hylobius abietis (L.) (Coleoptera: Curculionidae) were studied by taking root and soil samples around pine stumps on a clear-cutting in central Sweden. In addition, first-instar larvae migrating in the soil were sampled using traps baited with host-odour. Eggs were found in the soil rather than in the bark of stump roots, which previously has been regarded as the usual oviposition site. Based on an oviposition experiment and additional field observations we conclude that eggs are laid in the bark of roots only when the surrounding material is likely to dry out. We suggest two explanations for why weevils oviposit mainly in the soil, although they are known to show stereotypic behaviour when inserting eggs in stump roots: (1) egg predation by other arthropods or by conspecifics is avoided, and (2) newly hatched pine weevil larvae are better than ovipositing females at locating suitable sites for larval feeding.     

Soil type and microtopography influencing feeding above and below ground by the pine weevil Hylobius abietis

Abstract 1 The influence of soil type and microtopography on above and below ground feeding by adult pine weevils Hylobius abietis (L.) (Coleoptera: Curculionidae) was evaluated in a field experiment with enclosed weevil populations of known size. 2 Four soil treatments, each with a food source at the centre, were presented within each enclosure: (i) a flat surface with fine‐grained, cultivated humus; (ii) a flat surface with sand; (iii) a conical mound of sand; and (iv) a conical pit in sand. The food source consisted of a stem section of Scots pine Pinus sylvestris L. extending both above and below ground. 3 The majority of feeding on the half buried stem sections occurred below ground; only 2.7% of the total bark area consumed was situated above ground. The variation over time in bark area consumed was not significantly associated with any of the tested weather factors. 4 The amount of feeding was 10‐fold higher on food sources placed in fine‐grained humus than those in areas of flat sand. 5 Less pine bark was consumed on mounds of sand than flat sand surfaces, and there was more feeding in sandy pits than on flat sand. These effects on feeding are explained by the observation that the weevils had difficulties climbing the sandy slopes (27° gradient). 6 We conclude that pine weevil damage to conifer seedlings can be considerably reduced by planting on mounds of pure mineral soil and that planting deeply in the soil increases the risk of damage.     

Host-plant acceptance on mineral soil and humus by the pine weevil Hylobius abietis (L.)

1 The pine weevil Hylobius abietis (L.) (Coleoptera, Curculionidae) is an economically important pest of conifer forest regeneration in Europe and Asia. 2 Soil scarification, which usually exposes mineral soil, is widely used to protect seedlings from weevil attack. However, the mechanism behind this protective effect is not yet fully understood. 3 Field experiments were conducted to determine the pine weevil's responses to visual and odour stimuli from seedlings when moving on mineral soil and on undisturbed humus surface. 4 One experiment measured the number of pine weevils approaching seedlings, with and without added host odour, on mineral soil and undisturbed humus. Seedlings with added host odour attracted more weevils on both soil types. Unexpectedly, somewhat more weevils approached seedlings surrounded by mineral soil. 5 In a similar experiment, feeding attacks on seedlings planted directly in the soil were recorded. Only half as many seedlings were attacked on mineral soil as on undisturbed humus. 6 In the first experiment, the weevils were trapped 2.5 cm from the bases of the seedlings' stems, whereas they could reach the seedlings in the experiment where seedlings were planted directly in the soil. We conclude that the pine weevils' decision on whether or not to feed on a seedling is strongly influenced by the surrounding soil type and that this decision is taken in the close vicinity of the seedling. The presence of pure mineral soil around the seedling strongly reduces the likelihood that an approaching pine weevil will feed on it.     

Effects of green tree retention, prescribed burning and soil treatment on pine weevil (Hylobius abietis and Hylobius pinastri) damage to planted Scots pine seedlings

1 Feeding damage and mortality caused to planted Scots pine seedlings by the pine weevils Hylobius abietis and Hylobius pinastri were studied on burned and unburned sites with 0, 10 and 50 m³ per hectare levels of green tree retention from the second to the fourth summer after logging and burning of the sites. 2 The rate of severe feeding damage to pine seedlings caused by pine weevils was higher on burned clearcut sites than on unburned ones, whereas burning did not increase the feeding damage rate on sites with groups of retention trees. The damage rate in the fourth summer was approximately the same on burned and unburned sites. 3 Pine weevil feeding was the major cause of mortality of freshly planted pine seedlings on unburned sites. On burned sites, mortality was higher than the rate of severe feeding damage, particularly in the second summer after burning, possibly owing to fungal attack and abiotic factors. 4 At a retention tree level of 50 m³, feeding damage to the seedlings was lower than on clearcuts and at a 10 m³ retention tree level. Furthermore, on sites with 50 m³ of retention trees, scarification of the soil was found to decrease feeding damage more effectively than on clearcuts and 10 m³ sites. If the seedlings were situated in the centre of scarified patches, scarification alone was as effective as insecticide treatment on unscarified soil for decreasing feeding damage and mortality. 5 The results suggest that when burning is applied as silvicultural treatment after clear‐cuts, retention of trees is recommended to reduce the damages caused by pine weevils on pine seedlings.     

A flexible sand coating (Conniflex) for the protection of conifer seedlings against damage by the pine weevil Hylobius abietis

1 A new method for the physical protection of conifer seedlings against feeding damage by Hylobius abietis (L.), is described and evaluated in field trials in Swedish forest plantations.
2 The lower 60% of the stem of the seedling is protected by the Conniflex coating, consisting of fine sand (grain size = 0.2 mm) embedded in an acrylate dispersion that remains flexible after drying.
3 Seedlings are treated in the nursery by a large-scale application procedure involving four steps: (i) spraying the seedlings with water; (ii) application of fixative to the lower sections of the stems, (iii) application of fine sand to the fixative; and (iv) drying of the fixative.
4 A field experiment over three seasons demonstrated a significant increase in survival for coated seedlings compared with untreated seedlings. The survival rate increased from 29% to 97% for Scots pine and from 26% to 86% for Norway spruce. Coating the lower 30% of the stem (instead of 60%) provided inferior protection, resulting in only 64% survival in spruce.
5 Field trials in 11 commercial plantation areas indicated that the Conniflex sand coating was as effective in protecting seedlings as treatment with the insecticide imidacloprid.
6 The new method of coating conifer seedlings with fine sand provides an effective and environmentally sound alternative to insecticide treatment.     

Sub-lethal responses of the large pine weevil, Hylobius abietis, to the pyrethroid insecticide lambda-cyhalothrin

Abstract.  The response of the large pine weevil ( Hylobius abietis ) to the pesticide lambda-cyhalothrin, a pyrethroid, is investigated. Both behavioural (feeding preferences) and nutritional and physiological (lipid content) responses are recorded. Hylobius abietis shows both a significant avoidance of pesticide-treated food sources and a decrease in lipid content after exposure, but a full recovery after feeding on untreated food. It is proposed that the mechanism for the pesticide avoidance and altered lipid levels is due to an anti-feedant effect of the pesticide. Implications for pest management programmes are described.     

Growth and defence in young pine and spruce and the expression of resistance to a stem-feeding weevil

Defence in young trees has been much less studied than defence in older ones. In conifers, resin within ducts in bark is an important quantitative defence, but its expression in young trees may be influenced by developmental or physical constraints on the absolute size of the resin ducts as well as by differential allocation of resources to growth and resin synthesis. To examine these relationships, we used nitrogen fertilisation of 1- and 2-year-old pine and spruce to produce trees of different sizes and measured the effect on the number and size of resin ducts and the amount of resin they contained. All of these variables tended to increase with stem diameter, indicating a positive relationship between resin-based defence and growth of 1- and 2-year-old trees. In pine, however, the mass of resin flowing from severed ducts was much lower relative to duct area in 1- than in 2-year-old trees, suggesting that the older trees allocated a higher proportion of the carbon budget to resin synthesis. Resin-based defence in 1-year-old pines appears to be both positively related to growth and resource limited. In spruce, resin production was generally lower, and age-related differences were not observed, suggesting that resin-based defence is less important in this species. Bio-assays of 2-year-old trees with the pine weevil, Hylobius abietis, emphasised the importance of resin as a defence against this bark feeding insect. Nitrogen fertilisation had a limited influence on resistance expression. One-year-old trees remained susceptible because of their small size, low resin production and limited response to fertilisation. The strong growth response of 2-year-old trees to fertilisation increased resin-based defence, but most spruce trees remained susceptible, while most pines were resistant at all levels of fertilisation.     

Olfactory and visual stimuli used in orientation to conifer seedlings by the pine weevil, Hylobius abietis

Abstract. The influence of noncontact plant cues is investigated on the likelihood that individual conifer seedlings will be found by walking adults of the pine weevil, Hylobius abietis, in the field. Traps with solely odour or solely visual stimuli catch significantly more weevils than stimulus-free traps, and traps with the combination of odour and visual stimuli catch more weevils than traps with odour or visual stimuli alone. There is essentially an additive effect between odour and visual stimuli. The reactions to odour and visual stimuli are similar for three phases of the pine weevil's life cycle associated with three ages of clear-cuttings (i.e. sites where all trees have been harvested). Visual stimuli appear to be at least as important as odour for the pine weevil in finding an undamaged conifer seedling.     

Reproductive maturation in the large pine weevil Hylobius abietis: the relative importance of larval and adult diet

1 Larval and adult Hylobius abietis (L.) can feed and, in the case of the adults, cause commercially significant damage to many species of conifer. The present study aimed to evaluate the impact of larval and adult feeding on different host plants on subsequent reproductive fitness. 2 Hylobius abietis larvae were reared on logs of four host species known to vary in suitability for development; Pinus nigra ssp. laricio, Picea sitchensis, Larix kaempferi and Pseudotsuga menziesii. Adult females were collected on emergence and allocated to complete maturation feeding on one of four seedling conifer species, P. nigra spp. laricio, P. sitchensis, P. menziesii and Larix ×marschlinsii. It was assumed that variation in both larval and adult host plant quality had the potential to influence subsequent reproductive behaviour. 3 The time taken for females to attain reproductive maturity was assessed, and reproductive output was measured over an average period of 17 days. 4 The best predictor of the time to attain reproductive maturity was found to be the initial adult body weight. This was strongly related to the larval host species, with the largest adults emerging from P. nigra ssp. laricio. The time taken to begin oviposition was in the range 10–49 days (mean 23 days). The subsequent mean rate of oviposition for individual females was in the range 0.1–3.7 eggs per day. Egg volumes varied between and within individual females, in the range 0.241–1.079 mm³. 5 The species of seedling provided for maturation feeding had no significant impact on reproductive behaviour over the course of the experiment. It is possible, however, that the quality of the adult diet might exert a greater influence on behaviour over a longer period because H. abietis is a relatively long‐lived species.     

Identification of plant volatiles activating single receptor neurons in the pine weevil (Hylobius abietis)

Naturally produced plant volatiles, eliciting responses of single olfactory receptor neurons in the pine weevil, have been identified by gas chromatography linked with mass spectrometry. The receptor neurons (n = 72) were classified in 30 types, according to the compound which elicited the strongest response in each neuron, 20 of which compounds were identified. Most potent for 14 types of neurons (n = 50) were monoterpenes, including bicyclic (e.g. α-pinene, camphor and myrtenal) for 8 types (n = 32), monocyclic (limonene, carvone, α-terpinene) for 3 types (n = 12) and acyclic (e.g. β-myrcene and linalool) for 3 types (n = 6). Other compounds eliciting strongest responses of a neuron were five sesquiterpenes, including α-copaene and a farnesene-isomer, and an anethole type which has no biosynthetic relationship with terpenes. Within one type, receptor neurons with quite selective responses to the most potent compound as well as neurons with additional responses to several, structurally similar compounds were found, indicating that the neurons may have the same functional types of membrane receptors, but different sensitivities. Response spectra of neurons within the bicyclic-, mono-cyclic and acyclic types showed more overlapping than across the neuron types. Minimal overlapping response spectra was found between monoterpene and sesquiterpene neurons. The results suggest that this structure-activity relationship is significant for encoding plant odour information in the pipe weevil. Accepted: 6 January 1997     

Pine weevil damage to Norway spruce seedlings: effects of nutrient‐loading,soil inversion and physical protection during seedling establishment

• 1 The pine weevil Hylobius abietis (L.) feeds on the bark of young conifer seedlings and is one of the most economically important forest pests in Europe.
• 2 In a field experiment, we examined the combined effects of the treatments: nutrient‐loading of seedlings, planting in scarified plots and protection of seedlings against pine weevil damage for either half a season or a full season.
• 3 Nutrient loading had no significant effect on the amount of pine weevil feeding.
• 4 Seedling mortality was significantly reduced when seedlings were protected from pine weevil feeding during establishment. This occurred even though the debarked area of protected seedlings 5 weeks after the shields had been removed was similar to that of the unprotected seedlings. This indicates that initial protection rendered the seedlings more tolerant to later wounding by pine weevil.
• 5 Planting in soil inversion significantly reduced feeding compared with planting in humus.
• 6 We conclude that nutrient‐loading of seedlings in the autumn before planting would not increase pine weevil feeding after planting. Mortality could be reduced by treatments that postpone the start of pine weevil feeding on seedlings. Such treatments, combined with planting in soil inversion, would result in increased seedling growth, vitality and tolerance to pine weevil attack.

     

Larval mortality of pine sawflies [Hymenoptera: Diprionidae] in relation to pollution level: A field experiment

Larval mortality ofNeodiprion sertifer (Geoffroy),Diprion pini (L.) andGilpinia pallida (Klug) were studied in field experiments around a factory complex in southwestern Finland. Larval colonies were transferred on the shoots of Scots pines growing at different distances from the emission source. Larval mortality was highest near the factories. InN. sertifer, larval mortality caused by the nuclear polyhedrosis virus was higher and cocoon mortality caused by parasitoids was lower near the pollutant source. The most abundant parasitoid species wereSynomelix scutulata (Hartig) andLamachus eques (Hartig). 16–67% of theN. sertifer, 0–5% of theD. pini and the 73–100% ofG. pallida cocoons contained parasitoids oviposited during the larval period of the sawflies.      

Influence of insecticide, wax and biofungicide treatments, applied to Pinus sylvestris and Picea abies, on the olfactory orientation of the large pine weevil, Hylobius abietis L. (Coleoptera: Curculionidae)

1 A four-arm olfactometer was used to test the effect of permethrin ‘Gori 920^®’ and the wax treatment ‘CereNat^® E30’ on the response of mature adult Hylobius abietis to olfactory stimuli from seedlings of Pinus sylvestris and Picea abies. 2 Also investigated in the olfactometer was the effect of Rotstop^®, a biofungicide for the control of Fomes root and butt rot, Heterobasidion annosum, on the responses of mature adult female H. abietis to stump material of both P. sylvestris and P. abies. 3 Hylobius abietis was significantly more attracted to untreated P. sylvestris seedlings compared to those treated with CereNat^® E30 or permethrin Gori 920^®. No differences in response to P. abies seedlings were found. 4 With Rotstop^®, no changes in the attractiveness of P. sylvestris stump material were found, but that of P. abies was significantly increased compared to untreated material.     

Monitoring of gas exchange cycles and ventilatory movements in the pine weevil Hylobius abietis: respiratory failures evoked by a botanical insecticide

The large pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), is the most important insect pest of young coniferous plants. The implementation of new control methods requires not only a profound knowledge of the ecology and behaviour of the pest, but particularly of its physiology. Standard metabolic rate (SMR) and discontinuous gas exchange cycles (DGCs) were recorded in parallel with abdominal ventilation movements in adults of H. abietis using a differential electrolytic respirometer‐actograph. Quiescent weevils displayed DGCs of the constriction, flutter, and ventilation phases of the CFV type, while bursts of carbon dioxide were always accompanied by abdominal pumping movements, i.e., muscular ventilation in the closed subelytral cavity (SEC). In some beetles the C phase was absent and thus (C)FV cycles were recorded. In addition, at the beginning and often at the end of a burst, the SEC was rhythmically opened and closed by movements of the last abdominal segments. Continuous pumping movements and an absence of DGCs were signs of stress imposed by handling or by a new environment, even if the beetle was not moving. All individuals showed clear DGCs after recovering from handling and apparatus stress lasting 2–3 h. The results show that in the monitoring of DGCs, it is essential to determine whether they are of the constriction, flutter, and open phases (CFO), or the CFV subtype of the constriction, flutter, and burst (CFB) cycles. Use of our simple closed‐system respirometer enables non‐invasive simultaneous recording of SMR, oxygen uptake, DGCs, and active ventilation in H. abietis and other beetles. The topical application of adult H. abietis with sublethal doses of a botanical insecticide, NeemAzal T/S, caused essential respiratory failures: cyclic gas exchange was lost and irregular pumping movements appeared. In the treated beetles normal DGCs did not resume.     

Dispersal of the nuclear polyhedrosis virus of Neodiprion sertifer from soil to pine foliage with dust

Inclusion bodies (PIB) of nuclear polyhedrosis viruses (NPV) remain infectious for long periods in forest soils, but their transfer from soil to foliage is not well understood. Two instances of dispersal of the NPV of Neodiprion sertifer (Fourcroy) (Hymenoptera: Diprionidae) by dust are reported. The distributioo of diseased larval colonies in a young Scots pine stand indicated that dust caused by logging during winter could have started an epizootic. At another site there was a higher frequency of infected colonies near a forest road than further into the stand. The dust transmission hypothesis was tested by bioassays of road surface material and of samples of airborne dust collected on membrane filters. Both soil and air samples contained viable PIBs. Thus, virus dispersal with dust provided a likely explanation for the observed distributions of diseased larval colonies.

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Zusammenfassung Mortalität von N.sertifer Larven durch das wirtsspezifische Kernpolyedervirus ist zwischen den Massenvermehrungen der Sägewespe selten. Eine ständige Infektionskette bleibt nicht erhalten. Das einzige bekannte Reservoir von Virusinoculum zwischen den Massenvermehrungen ist der Waldboden. Die Übertragung von Virus vom Boden auf Kiefernnadeln wird als seltenes Ereignis betrachtet. Sie dürfte häufiger sein, wenn die bodenbedeckende Vegetation gestört ist. Es wird von zwei Studien berichtet, in denen offenbar NPV in windverfrachtetem Staub die Krankheit bei Larven verursachte.In einem jungen Bestand schottischer Kiefer, in dem keine neueren Meldungen von Virusauftreten vorlagen, wurden Proben genommen. Die Proben stammten aus verschiedenen Distanzen von einem ausgewachsenen, im vorherigen Winter gefällten Bestand. Die Befallsintensität sank hier scharf ab mit zunehmender Distanz vom Rand.Ferner wurde der Krankheitsbefall in Larvenkolonien an kleinen Bäumen nahe einer schmalen Forst-strasse und an Bäumen im Bestandesinnern untersucht: In Strassennähe war der Befall höher. Luftmuster wurden nahe der Strasse durch Membranfiltern gesaugt und in einem Biotest auf Virus untersucht. Wenn die Strassenoberfläche ungestört war, wurde kein Virus festgestellt. Wurde Staub aufgewirbelt, konnten infektiöse PIBs gefunden werden. Material von der Strassenoberfläche hatte eine hohe Virusaktivität.Die Übertragung des Virus vom Boden auf Kiefernnadeln dürfte demnach wichtig sein, um Krankheitsausbrüche auszulösen. Die beschriebenen Störungen waren vom Menschen verursacht. Ähnliches könnte sich unter andern Umständen ereignen, z.B. bei Windfall oder bei der Aktivität grösserer Tiere.