Monoterpenes emitted by the large pine weevil, Hylobius abietis (L.) feeding on Scots pine, Pinus sylvestris L.

Abstract. Attraction of the large pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae) to pine logs was enhanced by the presence of weevils already on the logs. We investigated the volatiles emitted by insects feeding on host trees to determine the mechanism of the attraction. Chemical analyses of the collected volatiles showed large amounts of monoterpenes. There were no qualitative differences in the collections, between Scots pine alone and pine plus feeding weevils, nor between males and females, mated or virgin. Strong quantitative differences in monoterpenes existed that were correlated to the surface of cut bark. Electroantennograms (EAGs) were recorded on male and female antennae stimulated by increasing doses of collected volatiles. EAGs for pine alone and pine plus weevils at the same stimulus loading in (a + P)-pinene gave superimposable curves. EAGs of male and female volatiles were also identical. These results suggest an attraction mediated by host plant allelochemicals rather than by an aggregation pheromone.     

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.     

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.     

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.     

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.     

Reproductive development and maternal effects in the pine weevil Hylobius abietis

1. In a laboratory study of maturation feeding of female pine weevil Hylobius abietis on current and 1‐year‐old stem bark of transplants of Scots and Corsican pine, Norway and Sitka spruce, Douglas fir, and hybrid larch, the length of the pre‐oviposition period was influenced by the species on which weevils fed. The shortest pre‐oviposition period was on hybrid larch (11.8 days) and the longest on Douglas fir (15.5 days). 2. The species on which weevils fed also affected fecundity but there was evidence of a species–year interaction. Over a period of 36 days, most eggs were laid by weevils feeding on current stem of Norway spruce and Corsican and Scots pine and fewest on current stem of Sitka spruce. 3. Significant maternal effects on egg size were observed both in relation to female size and conifer species. The largest eggs were laid on Corsican pine and the smallest on Douglas fir, with no evidence of a trade‐off between number of eggs laid and their size. 4. There was a positive relationship between egg and larval size and between larval size and survival on logs of four conifer species. Residual resistance mechanisms in the bark of recently cut stumps and larval competition are discussed briefly in relation to the importance of the observed maternal effects on weevil population dynamics.     

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.     

Peptide actions on oviduct contractions in the large pine weevil,Hylobius abietis

Abstract The peptides proctolin, crustacean cardioactive peptide (CCAP) and FMRFamide, which are known to modulate insect muscle contractions, were assayed for their action on oviduct contractions in Hylobius abietis. A video microscopy technique and computer‐based method of data acquisition and analysis were used to investigate the effects of theses peptides on spontaneous contractions of continuously perfused oviducts. All three peptides tested stimulate spontaneous contraction activity of the pine weevil oviduct, increasing the frequency and amplitude of phasic contractions in a dose‐dependent manner. Proctolin is more potent as a stimulator than CCAP. For proctolin a threshold response of oviduct muscles is at concentration of peptide 10^?11–10^?10 mol/L and for CCAP at concentration range 10^?10–10^?9 mol/L. FMRFamide exerts a weak stimulatory effect on the oviduct, and at higher concentrations of the peptide (above 10^?8 mol/L). The peptides exert different responses on oviduct contractions and they may play a role as functional regulators in such processes as egg movement and oviposition.     

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.     

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.     

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.     

Non-pesticide alternatives for reducing feeding damage caused by the large pine weevil (Hylobius abietis L.)

The large pine weevil (Hylobius abietis L.) is an important pest of young forest stands in Europe. Larvae develop under the bark of freshly cut pine and spruce stumps, but maturing weevils feed on the bark of coniferous seedlings. Such seedlings frequently die because of bark consumption near the root collar. We tested the effect of three treatments (the insecticide alpha cypermethrin, a wax coating and a glue coating) on the feeding damage caused by H. abietis on Douglas fir (Pseudotsuga menziesii) and Norway spruce (Picea abies) seedlings under semi-natural conditions. In two experiments (one in 2016 and another in 2017) seedlings in cages were subjected to pine weevil feeding for 16 weeks under shaded outdoor conditions. The experiment in 2016 compared insecticide and wax treatments and an untreated control on Douglas fir and Norway spruce, and the experiment in 2017 compared insecticide, wax and glue treatments and an untreated control on Norway spruce. In both experiments, all treatments significantly reduced H. abietis feeding damage at week 8 at the end of both experiments (week 16); the effect of treatments was significant only on spruce seedlings. The damages on Douglas fir seedlings was less on treated seedlings than on untreated control seedlings but differences were not significant. Coating stems with glue and especially with wax was generally effective at reducing weevil damage and in most cases provided control that was not significantly different from that provided by insecticide treatment. Our results suggest that a wax coating has the potential to replace the protection of seedlings provided by insecticides.     

Olfactory sensilla on the antennae of the pine weevil,Hylobius abietis

Summary Olfactory sensilla on the antennal club of the pine weevil, Hylobius abietis, have been studied by scanning and transmission electron microscopy. Two main types, sensilla basiconica and sensilla trichodea, have been classified. The former are found in two variants, one containing one sensory cell and the other two. Sensilla trichodea have one sensory cell. The two sensillum types are localized in different fields of the club. The numerous sensilla basiconica are confined exclusively to the two constriction bands. Sensilla trichodea, fewer in number, are mainly restricted to the distal part where they also form a third ring. In addition, they are found immediately distal to the two construction bands. The fine structure of the two types suggests an olfactory function which could be proven by electrophysiological studies. The other hairs on the club are shown to belong to mechano- and taste sensilla and uninnervated setae.I am grateful to the Norwegian Research Council for Science and Humanities NAVF and the Norwegian Forest Research Institute for financial support; to Professor D. Schneider for providing laboratory facilities at Seewiesen; to him and his colleagues, particularly Dr. R.A. Steinbrecht, for helpful discussions and to Miss Barbara Müller for electron microscopy assistance. I also wish to thank Dr. T. Braathen and his staff for help with scanning electron microscopy facilities at the University of Oslo.     

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.     

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.     

Egg allocation by Bracon hylobii Ratz., the principal parasitoid of the large pine weevil (Hylobius abietis L.), and implications for host suppression

1 The braconid parasitoid Bracon hylobii Ratz. is one of the few specialist natural enemies of the large pine weevil, Hylobius abietis L., a destructive pest of conifer transplants. An assessment of its role as an agent of biological control requires a detailed knowledge of the allocation of its reproductive effort. 2 Parasitoid females were continuously observed in laboratory culture with individually reared host larvae in bark discs. The outcome of sequential parasitoid–host encounters was recorded by subsequent examination of hosts and by rearing all parasitoids. 3 Parasitoids avoided ovipositing on host larvae < 100 mg fresh weight, even though such larvae represented sufficient biomass for complete parasitoid development. All larger larvae were vulnerable to attack, which leaves a window of vulnerability for parasitoids of about 90% of weevil larval life. 4 Parasitoids presented with a range of host sizes showed no preference above 100 mg for the size of host first attacked, but allocated more eggs and a greater total handling time to larger hosts. 5 Most eggs were deposited on the first host attacked, with progressively fewer allocated to subsequent hosts. However, oviposition experience did not affect the time spent on the next host. 6 From these results it is anticipated that when weevil larval size is reduced by less favourable feeding substrates, fewer parasitoid eggs will be allocated to each but more host larvae will ultimately be attacked. 7 Generation time, host finding, oviposition rate, clutch size, life expectancy and diapause induction are strongly affected by temperature. Life expectancy is substantially shorter for parasitoids deprived of non‐host food supplement. At 15 and 20 °C the number of hosts attacked and the number of eggs deposited decreased with female age. 8 Bracon hylobii is inevitably poorly synchronized with a variable life‐cycle host; it is egg‐limited and can enter diapause at a relatively high field temperature. None of these characteristics suggest that it could stabilize the abundance of its host below an economically acceptable threshold density. However, the reproductive potential of the parasitoid suggests that it could make a significant contribution to larval mortality and suppress adult recruitment, thus complementing other control strategies.     

Adult large pine weevils Hylobius abietis feed on silver birch Betula pendula even in the presence of conifer seedlings

1 The feeding preference of the adult pine weevil Hylobius abietis (L.) (Coleoptera: Curculionidae) for Betula pendula Roth was studied in no‐choice and paired‐choice feeding experiments. 2 In the first no‐choice test, large quantities of silver birch bark in Petri dishes were consumed; on average, the daily consumption of each weevil was 67 mm². 3 In the second no‐choice test, the weevils were offered 1‐year‐old silver birch seedlings for 6 days. Initially, the weevils fed mostly on the stem bases; later, they moved upward to feed on other parts of the stems. In addition to the main shoots, scars caused by gnawing were also found on leaf bases, blades, veins and petioles. Feeding resulted in the death of the main stems in 15% of the seedlings. 4 In the paired‐choice tests, the conifers were preferred to silver birch, even though a large amount of silver birch was also consumed in the presence of conifers. 5 In the paired‐choice tests, equal amounts of Scots pine and Norway spruce were always consumed. When hybrid aspen was offered, only small amounts were gnawed.     

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.

     

The effect of red wood ant abundance on feeding damage by the pine weevil Hylobius abietis

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A fungal metabolite masks the host plant odor for the pine weevil (Hylobius abietis)

The pine weevil is one of the most important pest insects of conifer reforestation areas in Europe. Female pine weevils cover their eggs with chewed bark and feces (frass) resulting in avoidance behavior of feeding conspecifics towards egg laying sites. It has been suggested that microorganisms present in the frass may be responsible for producing deterrent compounds for the pine weevil. The fungi Ophiostoma canum, O. pluriannulatum, and yeast Debaryomyces hansenii were isolated from aseptically collected pine-weevil frass. The isolated fungi were cultured on weevil frass broth and their volatiles were collected by SPME and identified by GC–MS. D. hansenii produced methyl salicylate (MeS) as a major compound, whereas, in addition, O. canum and O. pluriannulatum produced 6-protoilludene. In a multi-choice lab bioassay, MeS strongly reduced pine weevil's attraction to the Pinus sylvestris volatiles. Thus, a fungal metabolite was found that strongly affects the pine weevil host-odor search.