Ants protect conifer seedlings from feeding damage by the pine weevil Hylobius abietis

• 1 Ants that protect food resources on plants may prey on (or deter) herbivores and thereby reduce damage. Red wood ants (of the Formica rufa group) are dominant ants in boreal forests of Eurasia and affect the local abundance of several herbivorous species.
• 2 The pine weevil Hylobius abietis (L.) is a herbivore that causes severe damage by feeding on the bark of coniferous seedlings within areas of forest regeneration.
• 3 We investigated whether ants can protect conifer seedlings from pine weevil feeding. In a manipulative experiment, ants were attracted to sugar baits attached to spruce seedlings and the damage caused by pine weevils was compared with control seedlings without ant‐baits.
• 4 The feeding‐scar area was approximately one‐third lower on the seedlings with ant‐baits compared with the controls. Besides red wood ants, Myrmica ants were also attracted in high numbers to the ant baits and the relative effects of these species are discussed.
• 5 The results obtained in the present study support the trophic cascade hypothesis (i.e. damage to herbivores is suppressed in the presence of predators). The decreased pine weevil feeding on the baited seedlings was probably a result of nonconsumptive interactions [i.e. the presence of (or harassment by) ants distracting pine weevils from feeding].
• 6 Understanding the role of ants may have important implications for future strategies aiming to control pine weevil damage. For example, maintaining suitable conditions for ants after harvesting stands may be an environmentally friendly but currently unexploited method of for decreasing weevil 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.     

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

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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.     

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.     

Feeding and oviposition rates in the pine weevil Hylobius abietis (Coleoptera: Curculionidae)

Feeding and oviposition in the pine weevil Hylobius abietis (Linnaeus) were monitored under laboratory conditions in two long-term experiments lasting over an extended breeding season. Data were also collected from weevils under semi-natural conditions outdoors. In addition, the effects of crowding and starvation were studied in separate experiments. During the main peak oviposition period, female H. abietis consumed 50% more bark tissue than males. When oviposition ceased, the feeding rate of the females declined to the same level as in the males. The rates and spatial distribution patterns of oviposition and feeding were clearly affected by climatic conditions and the degree of crowding. Females were estimated to lay on average 0.8 eggs per day during the season under outdoor conditions. The realized fecundity of a female weevil during the first season was estimated to be approximately 70 eggs. The estimated average rate of feeding was 23 mm2 of Scots pine bark per weevil per day. This implies that planted seedlings can only constitute a minor part of the food resources needed to sustain H. abietispopulations of the size that usually appear on fresh clear-cuttings in northern Europe.     

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.     

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.     

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.

     

Movement behaviour of the pine weevil Hylobius abietis in relation to soil type: an arena experiment

Pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), movement behaviour on different substrates was studied in an indoor arena using a video camera and digital image processing technique. We analysed individual variation in movement characteristics, i.e. turning angles, movement directions, and distance moved per unit time on the bare and level arena surface which consisted of mineral soil (sand) and/or humus sections in various spatial configurations. Pine weevils on humus did not turn back when they came to the border with the sand. However, most individuals moved faster on sand than on humus. Thus, the results suggest that interactions between substrate differences and individual movement behaviour may to some extent explain why pine weevils have been observed to feed less frequently on coniferous seedlings planted on mineral soil than on those planted on humus.     

Testing associational resistance against pine weevils mediated by Lasius ants attending conifer seedlings

Natural enemies attracted to plants may provide those plants with protection against herbivores but may also protect neighbouring plants, that is through associational resistance. Ant attendance may be an important mechanism for the occurrence of such effects because ants can reduce the damage caused by a wide variety of herbivorous insects. Ants have been shown, in a previous field experiment, to decrease the damage caused by the pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), a pest species that causes high seedling mortality in forest regeneration areas. In this study, we specifically tested whether seedlings planted close to ant‐attended seedlings experience associational resistance. We did this under laboratory conditions using the ant species Lasius niger (L.) (Hymenoptera: Formicidae). The feeding damage by pine weevils was significantly reduced on seedlings attended by ants. The neighbouring seedlings, however, did not experience associational resistance. Nevertheless, some associational effects were observed as the number of weevils recorded on both ant‐attended and neighbouring seedlings was significantly lower compared with ant‐excluded seedlings.     

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.     

Influence of host log age and refuge from natural enemies on colonization and survival of Phoracantha semipunctata

The pine cone weevil, Pissodes validirostris Gyll. (Coleoptera: Curculionidae), attacks seed cones of most Eurasian pine species, except these of Swiss stone pine (Pinus cembra L.). Behavioural responses of adult weevils to cone volatile emissions of Swiss stone pine and to those of a common host, mountain pine (Pinus uncinata Ram.), were compared in an olfactometer. Weevils were significantly attracted by the volatile blend emitted by mountain pine, but Swiss stone pine volatiles elicited an inverse response, with most weevils moving in the opposite direction to the odour source. However, the majority of second instar weevil larvae that were extracted from mountain pine cones and transferred into Swiss stone pine cones were capable of developing to the adult stage. This suggests that Swiss stone pine cones do not contain strong feeding deterrents that could prevent larval development. The possible factors involved in the absence of colonization of Swiss stone pine cones by cone weevils are discussed.     

Cues For Shelter Use in a Phytophagous Insect

Many insects spend a large proportion of their life inactive, often hiding in shelters. The presence of shelters may, therefore, influence where insects feed. This study examines stimuli affecting the use of shelters by adults of the pine weevil, Hylobius abietis (L.) (Coleoptera, Curculionidae). This species is an economically important forest pest in Europe since the adults feed on the stem bark of newly planted conifer seedlings. When there are hiding or burrowing places present in close proximity to a seedling, pine weevils may hide there and repeatedly return to feed on the same seedling. Experiments were conducted in a laboratory arena with above-ground or below-ground shelters and in the presence or absence of wind. Pine weevils were highly attracted to shelters both above and below ground. Weevils in shelters were often observed assuming a characteristic “resting” posture. Experiments with opaque and transparent shelters showed that visual stimuli are used for orientation towards shelters and also increase the probability of an individual remaining in a shelter. The presence of wind increased the weevils’ propensity to use shelters both above and below ground. The present study indicates that shelters have a major influence on the behavior of the pine weevil and possible implications of the results are discussed.     

Pathogenicity of Bursaphelenchus xylophilus on Pines in Minnesota and Wisconsin

The pinewood nematode, Bursaphelenchus xylophilus, was inoculated into established native jack and red pines (Pinus banksiana and P. resinosa) and exotic Austrian pine (P. nigra) in Minnesota and Wisconsin forests during summer 1981. The nematode isolates did not kill established nonstressed pine trees growing in the forest. However, the same nematode isolates killed pine seedlings under greenhouse conditions. Girdling the main stem of some trees to induce stress resulted in the death of the majority of inoculated and noninoculated branches of Austrian and jack pines, but no branch death was observed on red pine. Greater numbers of nematodes were extracted from branches of inoculated, girdled trees than from nongirdled trees. The mean number of nematodes extracted from branches of inoculated, nongirdled trees was 0.3 - 14 nematodes per gram of wood.     

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.     

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.     

Attraction of pepper weevil to volatiles from damaged pepper plants

Pioneer herbivorous insects may find their host plants through a combination of visual and constitutive host‐plant volatile cues, but once a site has been colonized, feeding damage changes the quantity and quality of plant volatiles released, potentially altering the behavior of conspecifics who detect them. Previous work on the pepper weevil, Anthonomus eugenii Cano (Coleoptera: Curculionidae), demonstrated that this insect can detect and orient to constitutive host plant volatiles released from pepper [Capsicum annuum L. (Solanaceae)]. Here we investigated the response of the weevil to whole plants and headspace collections of plants damaged by conspecifics. Mated weevils preferred damaged flowering as well as damaged fruiting plants over undamaged plants in a Y‐tube olfactometer. They also preferred volatiles from flowering and fruiting plants with actively feeding weevils over plants with old feeding damage. Both sexes preferred volatiles from fruiting plants with actively feeding weevils over flowering plants with actively feeding weevils. Females preferred plants with 48 h of prior feeding damage over plants subjected to weevil feeding for only 1 h, whereas males showed no preference. When attraction to male‐ and female‐inflicted feeding damage was compared in the Y‐tube, males and females showed no significant preference. Wind tunnel plant assays and four‐choice olfactometer assays using headspace volatiles confirmed the attraction of weevils to active feeding damage on fruiting plants. In a final four‐choice olfactometer assay using headspace collections, we tested the attraction of mated males and virgin and mated females to male and female feeding damage. In these headspace volatile assays, mated females again showed no preference for male feeding; however, virgin females and males preferred the headspace volatiles of plants fed on by males, which contained the male aggregation pheromone in addition to plant volatiles. The potential for using plant volatile lures to improve pepper weevil monitoring and management is discussed.     

Feeding by short-snouted weevils on seedlings of seven tree species planted beneath Norway spruce at differing stand densities

1 Short-snouted weevils, including Strophosoma spp. and related species, may damage tree seedlings. We investigated the damage caused by these weevils feeding on seedlings of seven tree species planted in clear-cuts and under shelterwoods with three densities (control, dense, and sparse).
2 There were no pronounced differences in damage caused by short-snouted weevils in the various shelterwood densities. Most feeding occurred in the clear-cuts during the first year, whereas more seedlings were damaged in the shelterwoods of all densities in the second year. In the third year, virtually no feeding occurred in either the clear-cuts or shelterwoods. We cannot explain this pattern of damage, but a fallow period of 2 years seems to prevent short-snouted weevil damage.
3 We determined the general feeding preferences for short-snouted weevils to be cherry, lime > beech, oak, spruce > maple, ash. However, although Strophosoma mellanogrammum [Correction added after online publication 8 December 2008: Strophosoma melanogrammum corrected to Strophosoma mellanogrammum ] was observed feeding on seedlings, no full short-snouted weevil inventory was conducted, so caution must be exercised when drawing conclusions from this study regarding weevil damage patterns and feeding preferences.
4 Short-snouted weevils can sometimes occur in large numbers and may destroy entire reforestation projects, but in our study their influence on seedling growth and survival was minor.     

Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration

The boreal forest is expected to experience the greatest warming of all forest biomes, raising concerns that some of the large quantities of soil carbon in these systems may be added to the atmosphere as CO₂. However, nitrogen deposition or fertilization has the potential to increase boreal forest production and retard the decomposition of soil organic matter, hence increasing both tree stand and soil C storage. The major contributors to soil‐surface CO₂ effluxes are autotrophic and heterotrophic respiration. To evaluate the effect of nutrient additions on the relative contributions from autotrophic and heterotrophic respiration, a large‐scale girdling experiment was performed in a long‐term nutrient optimization experiment in a 40‐year‐old stand of Norway spruce in northern Sweden. Trees on three nonfertilized plots and three fertilized plots were girdled in early summer 2002, and three nonfertilized and three fertilized plots were used as control plots. Each plot was 0.1 ha and contained around 230 trees. Soil‐surface CO₂ fluxes, soil moisture, and soil temperature were monitored in both girdled and nongirdled plots. In late July, the time of the seasonal maximum in soil‐surface CO₂ efflux, the total soil‐CO₂ efflux in nongirdled plots was 40% lower in the fertilized than in the nonfertilized plots, while the efflux in girdled fertilized and nonfertilized plots was 50% and 60% lower, respectively, than in the corresponding nongirdled controls. We attribute these reductions to losses of the autotrophic component of the total soil‐surface CO₂ efflux. The estimates of autotrophic respiration are conservative as root starch reserves were depleted more rapidly in roots of girdled than in nongirdled trees. Thus, heterotrophic activity was overestimated. Calculated on a unit area basis, both the heterotrophic and autotrophic soil respiration was significantly lower in fertilized plots, which is especially noteworthy given that aboveground production was around three times higher in fertilized than in nonfertilized plots.