Carbon–nitrogen stoichiometry in the tritrophic food chain willow, leaf beetle, and predatory ladybird beetle

Although plant quality can indirectly increase the performance of the third trophic level by bottom-up cascading effects, the mechanisms of this indirect effect are still unclear. In this study the carbon–nitrogen stoichiometry in a tri-trophic system consisting of the willow, a leaf beetle, and a predatory ladybird beetle were examined to determine the mechanisms of the bottom-up cascading effect. The bottom-up cascade is initiated by increasing leaf nitrogen, because of artificial cutting of willow trees. The relative growth rate (RGR) of the leaf beetle increased when fed on cut willow leaves, because of the high leaf nitrogen in the cut willows. Ladybird beetle RGR also increased when fed on leaf beetles fed on cut willow leaves. The increased RGR of the ladybirds cannot be explained by the quality of the prey, however, because leaf beetle nitrogen was not affected by host plant quality. Thus, the carbon–nitrogen stoichiometry could not be a mechanism of the bottom-up cascade through multiple trophic levels.     

Host plant variation in plant-mediated indirect effects: moth boring-induced susceptibility of willows to a specialist leaf beetle

Abstract.  1. We examined the plant-mediated indirect effects of the stem-boring moth Endoclita excrescens (Lepidoptera: Hepialidae) on the leaf beetle Plagiodera versicolora (Coleoptera: Chrysomelidae) in three willow species, Salix gilgiana , S. eriocarpa , and S. serissaefolia.
2. When the stem-boring moth larvae damaged stems in the previous year, willows were stimulated to produce vigorously growing lateral shoots on these stems. These new lateral shoots were significantly longer and the upper leaves had significantly higher nitrogen and water content than current-year shoots on unbored stems, although the carbon content and leaf dry mass were not different between lateral and current-year shoots.
3. In the field, leaf beetle larvae and adults had significantly greater densities on lateral shoots of bored stems than on current-year shoots of unbored stems. A laboratory experiment showed that female beetles had significantly greater mass and fecundity when fed on leaves of newly-emerged lateral shoots. Thus, the stem-boring moth had a positive effect on the temporally and spatially separated leaf beetle by increasing resource availability by inducing compensatory regrowth.
4. The strength of the indirect effects on the density and performance of the leaf beetle differed among willow species, because there was interspecific variation in host quality and herbivore-induced changes in plant traits. In particular, we suggest that the differences in magnitude of the changes among willow species in shoot length and leaf nitrogen content greatly affected the strength of the plant-regrowth mediated indirect effect, coupled with host-plant preference of the leaf beetle.     

Plant phenology-mediated indirect effects: The gall midge opens the phenological window wider for a leaf beetle

We examined whether larvae of the gall midge Rabdophaga rigidae (Diptera: Cecidomyiidae) can modify the seasonal dynamics of the density of a leaf beetle, Plagiodera versicolora (Coleoptera: Chrysomelidae), by modifying the leaf flushing phenology of its host willow species, Salix serissaefolia and Salix eriocarpa (Salicaceae). To test this, we conducted field observations and a laboratory experiment. The field observations demonstrated that the leaf flushing phenology of the willows and the seasonal dynamics of the beetle density differed between shoots with stem galls and shoots without them. On galled shoots of both willow species, secondary shoot growth and secondary leaf production were promoted; consequently, leaf production showed a bimodal pattern and leaf production periods were 1 to 2 months longer than on non‐galled shoots. The adult beetle density on galled shoots was thus enhanced late in the season, and was found to change seasonally, synchronizing with the production of new leaves on the host willow species. From the results of our laboratory experiment, we attributed this synchrony between adult beetle density and willow leaf flush to beetles’ preference to eat new leaves rather than old. Indeed, beetles consumed five times more of the young leaves when they were fed both young and old leaves. These results indicate that stem galls indirectly enhance the adult beetle density by enhancing food quality and quantity late in the beetle‐feeding season. We therefore conclude that midge galls widen the phenological window for leaf beetles by extending the willows’ leaf flush periods.     

Nitrogen homeostasis in a willow leaf beetle, Plagiodera versicolora, is independent of host plant quality

While foliar nitrogen (N) content of host plants depends on environmental conditions, N content of herbivorous insects may remain relatively constant due to homeostasis. However, it is unknown to what extent insects can maintain their body elemental composition against natural variation in host plant quality. The present study examined the performance and N content of a willow leaf beetle, Plagiodera versicolora Laicharting (Coleoptera: Chrysomelidae), when fed leaves of host willow, Salix eriocarpa Franchet et Savatier (Salicaceae), with varying nutritional status. Water content, toughness, and N content of willow leaves varied seasonally, and they affected performance of the leaf beetle. The leaf beetle achieved high performance when fed young leaves. On the other hand, the N content of the leaf beetle changed little, and it was independent of that of willow leaves, indicating strong N homeostasis of the leaf beetle. We discussed the function of N homeostasis in herbivorous insects in tritrophic level interactions.     

The suitability of damaged willow leaves as food for the leaf beetle, Phgiodera versicolora

Abstract. 1. The effect of previous damage on the suitability of willow ( Mix bubylonicu and S.ulbu 'Tristis') leaves as food for the imported willow leaf beetle, Plugiodera versicoloru Laich. (Coleoptera: Chrysomelidae), was evaluated by feeding adults and larvae leaves from damaged and undamaged branchlets.
2. Females had lower fecundity when fed leaves from artificially damaged branchlets of S.ufba 'Tristis'. A similar result was obtained when beetles ate leaves from naturally damaged branchlets of S.babylonica .
3. Feeding preference tests suggest that the reductions in leaf suitability were not due to adults avoiding damaged leaves as a food source.
4. PZugiodera versicolora larvae required longer to develop and attained a lower adult weight when fed leaves from damaged branchlets.
5. Changes in leaf suitability brought about by herbivore feeding may affect the seasonal abundance of P.versicoloru on its willow hosts.     

Trunk cutting initiates bottom-up cascades in a tri-trophic system: sprouting increases biodiversity of herbivorous and predaceous arthropods on willows

We examined the effects of trunk cutting on the regrowth responses of two dominant willow species, Salix eriocarpa and S. gilgiana , and the subsequent effects on the community structure (abundance and species richness) of herbivorous and predaceous arthropods. We studied ten randomly selected pairs of cut and uncut (control) trees of each willow species. Field observations showed that when the trunks were cut, shoots sprouted from the base and developed rapidly. These shoots continued to grow until July, producing lateral shoots and leaves, whereas current-year shoots on uncut trees essentially stopped growing by April. In July, the upper leaves of cut trees were less tough and had a greater water and nitrogen content than leaves of uncut trees. Leaf consumption and abundance and species richness of both herbivorous and predaceous arthropods were significantly greater on cut trees than on uncut trees in both willow species. Overall, trunk cutting resulted in at least a two-fold increase in both relative abundance and species richness within the arthropod community on the willow species. We concluded that the severe physical damage caused by trunk cutting greatly increased the biodiversity of herbivorous and predaceous arthropods through bottom-up cascading effects.     

Do exotic generalist predators alter host plant preference of a native willow beetle?

1 Selection can favour herbivores that choose host plants benefitting their offspring either by enhancing growth rates or by increasing larval defences against native predators. For exotic predator species that feed on herbivores, their success with invading new habitats may depend upon overcoming defences used by native prey. Whether exotic predators can alter herbivore host choice has remained unexamined. Therefore, we compared the efficacy of larval defence by Chrysomela knabi (a native beetle species) that had fed on two native willow hosts: Salix sericea (a phenolic glycoside (PG)-rich species) and Salix eriocephala (a PG-poor species), when attacked by exotic generalist predators. In addition, the preference and performance of C. knabi on S. sericea and S. eriocephala was examined.
2  Chrysomela knabi preferred and performed better on S. sericea. In a common garden, adult C. knabi were nine-fold more common and oviposited five-fold more frequently on S. sericea than on S. eriocephala . In the laboratory, adult feeding preference on leaf discs and survival rates of larvae were both greater on S. sericea , and time to pupation was shorter.
3  Chrysomela knabi larvae produced significantly more salicylaldehyde when fed S. sericea leaves than when fed S. eriocephala leaves. Additionally, those larvae with greater salicylaldehyde had reduced predation by two exotic generalist predators, Harmonia axyridis larvae and juvenile Tenodera aridifolia sinensis .
4 The results obtained in the present study suggest that selection favoured the preference of C. knabi for PG-rich willow plants because larvae grew and survived better and that selection by common exotic generalist predators would reinforce this preference.     

The importance of specialist natural enemies for Chrysomela lapponica in pioneering a new host plant

Abstract.  1. The Salicaceae have been suggested as ancestral host plants of Chrysomela species (Coleoptera: Chrysomelidae). In Chrysomela lapponica , some populations are specialised on salicaceous plants, but others have switched to birch. This study aimed to elucidate the significance of natural enemies as possible selective forces for the host plant shift of C. lapponica from willow to birch.
2. Two C. lapponica populations were studied, one specialised on willow Salix borealis in Finland, and another one specialised on birch Betula pubescens in the Czech Republic. Abundances of predators and parasitoids on birches and willows were recorded at both population sites. Furthermore, field and laboratory experiments were conducted.
3. Field data do not support the hypothesis that generalist predators affected the host shift from willow to birch in C. lapponica.
4. Parasitism of C. lapponica (pre)pupae by a specialised phorid fly was significantly stronger in specimens living on willow than in birch-living ones.
5. The predatory syrphid Parasyrphus nigritarsis specialised on Chrysomelinae was only detected on willows. The syrphid preferred to orient towards substrates treated with defensive larval secretion or faeces of the willow-specialised C. lapponica specimens compared with the birch-specialised ones.
6. The data suggest that specialised parasitoids and predators might have been driving forces for C. lapponica to leave willows and to pioneer birches as sites with a lowered risk of predation and parasitism. This hypothesis is discussed with respect to results of earlier studies on the impact of bottom-up effects by the plant.     

Can herbivore‐induced plant volatiles inform predatory insect about the most suitable stage of its prey?

The leaf beetle Plagiodera versicolora (Coleoptera: Chrysomelidae) is a specialist herbivore, all of whose mobile stages feed on the leaves of salicaceous plants. Both the larval and adult stages of the ladybird Aiolocaria hexaspilota (Coleoptera: Coccinellidae) are dominant natural enemies of the larvae of the leaf beetle. To clarify the role of plant volatiles in prey‐finding behaviour of A. hexaspilota, the olfactory responses of the ladybird in a Y‐tube olfactometer are studied. The ladybird adults show no preference for willow plants Salix eriocarpa that are infested by leaf beetle adults (nonprey) over that for intact plants but move more to the willow plants infested by leaf beetle larvae (prey) than to intact plants. Moreover, ladybird larvae show no preference for willow plants infested by leaf beetle larvae or adults over intact plants. Using gas chromatography‐mass spectrometry, six volatile compounds are released in larger amounts in the headspace of willow plants infested by leaf beetle larvae than in the headspace of willow plants infested by leaf beetle adults. In addition, the total amount of volatiles emitted from willow plants that are either intact or infested by leaf beetle adults is much smaller than that from willow plants infested by leaf beetle larvae. These results indicate that volatiles from S. eriocarpa infested by P. versicolora inform A. hexaspilota adults about the presence of the most suitable stage of their prey, whereas A. hexaspilota larvae do not use such information.     

Habitat difference in abundance of willow leaf beetle Phratora vulgatissima (Coleoptera: Chrysomelidae): plant quality or natural enemies?

Herbivorous insects are influenced by both 'bottom-up' forces mediated through host plants and 'top-down' forces from natural enemies. Few studies have tried to evaluate the relative importance of the two forces in determining the abundance of insects. The leaf beetle Phratora vulgatissima Linnaeus sometimes occurs at high densities and severely damages the willow Salix cinerea in forest habitats. For willows growing in open agricultural landscapes (farmland S. cinerea), the leaf beetle generally occurs at low densities and plants receive little damage. The purpose of the present study was to evaluate the relative importance of host plant quality and natural enemies behind the observed difference in P. vulgatissima abundance. Female egg-laying and larval performance (growth and survival) were studied on caged willow branches in the field to investigate if plant quality differs between S. cinerea trees growing in forest and farmland habitats. The survival of eggs exposed to natural enemies was examined to see if predation could explain the low abundance of leaf beetles on farmland willows. The results indicated no difference in plant quality; female egg laying and larval performance did not differ between the forest and the farmland. However, heteropteran predators (true bugs) were more abundant, and the survival of eggs was lower, on plants in the farmland habitat than in the forest habitat. The data suggest that the low abundance of P. vulgatissima on farmland willows could not be explained by a poor quality of plants, but more likely by high predation from heteropterans.     

Defensive larval secretions of leaf beetles attract a specialist predator Parasyrphus nigritarsis

1. Noxious larval secretions of leaf beetles, which repel generalist predators, do not deter specialist syrphid fly predators (genus Parasyrphus ). These flies cause considerable mortality to the beetles, but little is known about their foraging behaviour.
2. Larvae of Parasyrphus nigritarsis were attracted to the volatile larval secretions produced by two prey species Phratora vitellinae and Linaeidea aenea. Parasyrphus nigritarsis feeds on both beetles in nature. Phratora vitellinae feeds on willows and utilizes host plant compounds for secretion production, while the alder-feeding L. aenea produces an autogenous secretion.
3. Fly larvae were strongly attracted to pieces of filter paper treated with larval secretion of the beetles. They attempted to feed on them for up to 7 min, and were equally attracted to the secretions of Ph. vitellinae and L. aenea . Fly larvae were also attracted to pure salicyl aldehyde, the main component of the secretion of Ph. vitellinae .
4. Fly larvae searched extensively for prey on leaves that had been damaged by beetle larvae. They also followed trails made with solutions containing faecal matter of prey larvae. They showed no differential preference for Ph. vitellinae or L. aenea , but always rejected larvae of the non-prey leaf beetle Agelastica alni .
5. Beetle secretions thus play an important, but unexpected, role in the feeding behaviour of P. nigritarsis . This predator uses the beetle secretion to locate its prey. The implications of these results for three trophic level interactions are discussed.     

Size class structure and tree dispersion patterns in old-growth cedar-hemlock forests of the northern Rocky Mountains (USA)

Summary The relationship between the food selection of four leaf beetle species (Phratora vitellinae, Plagiodera versicolora, Lochmaea capreae, Galerucella lineola) and the phenolic glycosides of willow (Salix spp.) leaves was tested in laboratory food choice experiments. Four willow species native to the study area (Eastern Finland) and four introduced, cultivated willows were tested.The willow species exhibited profound differences in their phenolic glycoside composition and total concentration. The food selection patterns of the leaf beetles followed closely the phenolic glycoside spectra of the willow species. Both the total amount and the composition of phenolic glycosides affected the feeding by the beetles. Phenolic glycosides apparently have both stimulatory and inhibitory influences on leaf beetle feeding depending on the degree of adaptation of a particular insect. Very rare glycosides or exceptional combination of several glycoside types seem to provide certain willow species with high level of resistance against most herbivorous insects. Analogously the average absolute amount of leaf beetle feeding was lower on the introduced willows than on the native species to which the local herbivores have a good opportunity to become adapted.     

A trophic cascade in a macrophyte-based food web at the land–water ecotone

Trophic cascades may purportedly be more common in aquatic than terrestrial food webs, but herbivory on freshwater vascular plants has historically been considered low. Water lilies are an exception, suffering severe grazing damage by leaf beetles. To test whether a central prediction of cascade models—that predator effects propagate downwards to plants—operates in a macrophyte-based food web, we experimentally manipulated predation pressure on a key herbivore of water lilies in the littoral zone of a lake in Michigan, USA. Field experiments comprised combinations of caging treatments to alter the number of predators (larvae of the ladybird beetle Coleomegilla maculata) that hunt the grazers of the macrophytes (larvae of the leaf beetles Galerucella nymphaeae) on the leaves of the water lily Nuphar advena. Predatory larvae of the ladybird beetles significantly reduced grazing damage to water-lily leaves by 35–43%. The predators reduced plant damage chiefly via density-mediated effects, when lower densities of grazers translated to significant declines in plant damage. Plant damage caused by the surviving herbivores was less than predicted from individual grazing rates under predator-free conditions. This suggests that trait-mediated effects may possibly also operate in this cascade. The observed strong effect of predators on a non-adjacent trophic level concurs with an essential component of the trophic cascade model, and the cascade occurred at the ecotone between aquatic and terrestrial habitats: Nuphar is an aquatic macrophyte with emergent and floating leaves, whereas both beetle species are semi-terrestrial and use the dry, emergent and floating leaves of the water lily as habitat. Also, the cascade is underpinned by freshwater macrophytes—a group for which trophic processes have often been underappreciated in the past.     

Leaf beetle larvae,Plagiodera versicolora (Coleoptera: Chrysomelidae), show decreased performance on uninfested host plants exposed to airborne factors from plants infested by conspecific larvae

Undamaged plants are known to suffer less damage from herbivores when previously exposed to airborne factors from neighboring plants that are either infested or artificially damaged. However, to date, the effects of such a defensive phenomenon on performance of herbivorous insects have not been clearly shown. Here, we studied such effects in an interaction between a willow plant, Salix eriocarpa Franchet et Savatier (Salicales: Salicaceae), and a specialist leaf beetle, Plagiodera versicolora (Laicharting) (Coleoptera: Chrysomelidae). In a wind tunnel, uninfested willow plants were placed downwind of willow plants infested by leaf beetle larvae for 4 days. As a control, we placed uninfested plants downwind of uninfested plants in the tunnel. After exposure, downwind plants were served to leaf beetle larvae. Pupal weight, larval survival rates, and the leaf area consumed by larvae all decreased significantly, and larval developmental duration increased significantly, when larvae fed on willow plants downwind of infested plants were compared with those downwind of uninfested plants. These results showed that airborne factors from infested willow plants negatively affected the performance of leaf beetle larvae. Further studies are needed to identify the active factor(s) from the infested willow plants affecting the performance of leaf beetle larvae.     

Host-plant effects on larval survival of a salicin-using leaf beetle Chrysomela aeneicollis Schaeffer (Coleoptera: Chrysomelidae)

Several species of willow leaf beetles use hostplant salicin to produce a defensive secretion that consists of salicylaldehyde. Generalist arthropod predators such as ants, ladybird beetles, and spiders are repelled by this secretion. The beetle larvae produce very little secretion when they feed on willows that lack salicylates, and salicin-using beetles prefer salicylate-rich willows over salicylate-poor ones. This preference may exist because the larvae are better defended against natural enemies on salicylate-rich willows. If this is true, the larvae should survive longer on those willows in nature. However, this prediction has not been tested. I determined the larval growth and survival of Chrysomela aeneicollis (Coleoptera: Chrysomelidae) on five willow species (Salix boothi, S. drummondiana, S. geyeriana, S. lutea, and S. orestera). These species differed in their salicylate chemistries and in leaf toughness but not in water content. The water content varied among the individual plants. Larval growth of C. aeneicollis did not differ among the five species in the laboratory, but it varied among the individual plants and it was related to the water content. In the field, C. aeneicollis larvae developed equally rapidly on the salicylate-poor S. lutea and on the salicylate-rich S. orestera. Larval survival was greater on S. orestera than on S. lutea in one year (1986), but there was no difference between them during three succeeding years. In another survivorship experiment, larval survival was low on the medium-salicylate S. geyeriana, but high on the salicylate-poor S. boothi and on S. orestera. Larval survival in the field was related to the larval growth and water content that had been previously measured in the laboratory. These results showed that the predicted relationship between the host plant chemistry and larval survival did not usually exist for C. aeneicollis. One possible reason for this was that the most important natural enemies were specialist predators that were unaffected by the host-derived defensive secretion. One specialist predator, Symmorphus cristatus (Hymenoptera: Eumenidae), probably caused much of the mortality observed in this study. I discuss the importance of other specialist predators to salicin-using leaf beetles.     

Host‐plant quality adaptively affects the diapause threshold: evidence from leaf beetles in willow plantations

1. Voltinism of herbivorous insects can vary depending on environmental conditions. The leaf beetle Phratora vulgatissima L. is univoltine in Sweden but will sometimes initiate a second generation in short‐rotation coppice (SRC) willow plantations. 2. The study investigated whether increased voltinism by P. vulgatissima in plantations can be explained by (i) rapid life‐cycle development allowing two generations, or (ii) postponed diapause induction on coppiced willows. 3. In the field, no difference was found in the phenology or development of first‐generation broods between plantations (S. viminalis) and natural willow habitats (S. cinerea). However, the induction of diapause occurred 1–2 weeks later in SRC willow plantations. 4. Laboratory experiments indicated no genetic difference in the critical day‐length for diapause induction between beetles originating from plantations and natural habitats. Development time was unaffected by host‐plant quality but critical day‐length was prolonged by almost an hour when the beetles were reared on a non‐preferred willow species (S. phylicifolia). When reared on new leaves from re‐sprouting shoots of recently coppiced willow plants, diapause incidence was significantly less than when the beetles were reared on mature leaves from uncoppiced plants. 5. The study suggests that P. vulgatissima has a plastic diapause threshold influenced by host‐plant quality. The use of host‐plant quality as a diapause‐inducing stimulus is likely to be adaptive in cases where food resources are unpredictable, such as when new host‐plant tissue is produced after a disturbance. SRC willows may allow two beetle generations due to longer growing seasons of coppiced plants that grow vigorously.     

Host plant variation in mortality of the leaf-folding sawfly on the arroyo willow

Abstract. 1. We tested the hypothesis that survival and sources of mortality of the leaf-folding sawfly ( Phyllocolpa sp.) varied significantly among host plants of the arroyo willow ( Salix lasiolepis Bent ham).
2. Survival of the leaf folder differed among field and potted willows in a common environment in two of three cases, and sources of mortality differed among plants in four of five cases.
3. Egg mortality differed among field plants but not among the potted willow plants.
4. Larval mortality and parasitism differed among field and potted willows in 2 years, and appeared to be compensatory mortality sources.
5. Leaf folder density among plants was not generally correlated with percentage of leaf folds with no egg (galls formed but no subsequent oviposition), percentage survival, or percentage mortality, indicating a general lack of density dependence.
6. The proportion of folds with no egg oviposited differed significantly among field and potted plants, and was only correlated with survival or sources of mortality in one of three years.
7. The data support the hypothesis that host plant genotype affects the interaction of the leaf folder with its natural enemies, and thus represents a three trophic level interaction.     

Chlorogenic acid as an antiherbivore defence of willows against leaf beetles

In the leaves of 13 Finnish willow species, the content of a phenolic, chlorogenic acid, was found to vary from 0 up to 18 mg g^–1 D.W. Effects of pure chlorogenic acid on insect feeding behaviour were tested using four common leaf beetle species which are in the field mainly found on willows with low-chlorogenic acid leaves. One species, Lochmaea capreae L., was invariably deterred by pure chlorogenic acid applied in naturally occurring concentrations on the willow leaves. Accordingly, in 2-choice laboratory feeding trials L. capreae was found to prefer low-chlorogenic acid leaves of four willow species over high-chlorogenic acid leaves of Salix pentandra L. and S. myrsinifolia Salisb. When presented on the leaves of S. phylicifolia L, pure chlorogenic acid inhibited also the feeding by Phratora polaris Sp.-Schn. Instead, chlorogenic acid had no significant effect on Ph. polaris when it was presented on the leaves of another willow S. cinerea L. In laboratory, Ph. polaris did not show general preference for willow species with low chlorogenic acid content in their leaves. Thus, the response of Ph. polaris to chlorogenic acid seems to depend on the plant species. Apparently variation in other traits such as leaf hairyness may easily override the potential effect of chlorogenic acid content on Ph. polaris. To two other leaf beetle species, Galerucella lineola F. and Plagiodera versicolora Laich., chlorogenic acid is an ineffective deterrent even at unnaturally high concentrations. In laboratory, G. lineola and P. versicolora did not prefer willows with low chlorogenic acid content in their leaves. Thus, among four studied leaf beetle species, only L. capreae seems to be clearly affected by this phenolic. Therefore, overall importance of chlorogenic acid as a defence against willow-feeding leaf beetles appears to be very limited.     

Influence of leaf trichome density on the efficiency of two polyphagous insect predators

Plant characteristics, such as leaf structure or hairiness, are important for the movement and attachment of insects. It has been suggested that increased trichome density on new Salix cinerea L. (Salicaceae) leaves, produced after grazing by the willow leaf beetle Phratora vulgatissima L. (Coleoptera: Chrysomelidae), function as an inducible defence against the beetle and especially its larvae. Here we studied whether and how two of the main natural enemies of P. vulgatissima, viz., Anthocoris nemorum L. (Heteroptera: Anthocoridae) and Ortothylus marginalis L. (Heteroptera: Miridae), were influenced by trichome density on S. cinerea leaves. The effect of trichome density on these two predators was studied on plants with different trichome densities, comparing natural enemy efficiency, measured as number of P. vulgatissima eggs consumed or larvae missing and/or killed. To obtain different trichome densities, cuttings of several different clones of S. cinerea were used. In the experiment using eggs as prey, an increase in trichome density was, in addition, induced through leaf beetle defoliation on half of the plants of each willow clone. Furthermore, a field study was performed to investigate whether trichome density was correlated with natural enemy abundance. The results indicate that neither the efficiency of these two natural enemies in the greenhouse, nor their abundance in the field was influenced by trichome density. A well‐known behavioural difference between the two predator species could probably account for the higher disappearance of larvae after exposure to the more active predator. These findings are relevant for the development of pest management programs, not least because the enemies are polyphagous predators. It is concluded that an induced increase in leaf hairiness in willows in response to leaf beetle grazing could be a plant resistance trait worthy of further study in this system, because no negative effects on the main natural enemies were observed.     

Effects of leaf toughness on mandibular wear of the leaf beetle, Plagiodera versicolora

Abstract. 1. The effects of leaf toughness on mandibular wear of the leaf beetle Plagiodera versicolora Laich. (Coleoptera: Chrysomelidae) were evaluated by feeding adult beetles young (tender) and old (tough) leaves of Salix babylonica and S. alba 'Tristis'.
2. Tough leaves erode the cutting surface of beetle jaws more so than tender leaves.
3. Beetles with worn jaws consumed leaves at a slower rate than ones with less mandibular wear.
4. Because rates of leaf consumption and egg production are positively correlated, increased mandibular wear may reduce beetle fecundity.
5. These results support the belief that leaf toughness may act as a potent defence affecting morphology, feeding behaviour, and ultimately spatial and temporal patterns of herbivores.