Factors affecting willow leaf beetles (Phratora vulgatissima) when selecting overwintering sites

Abstract

• 1 Leaf beetles are major pests in willow plantations used as short‐rotation coppice for biomass production. The beetles overwinter mainly outside the plantations. An understanding of the factors affecting adult leaf beetles seeking overwintering sites may provide information that could be valuable when developing methods to control the beetles.
• 2 We conducted a field experiment in a willow plantation with a high abundance of the leaf beetle, Phratora vulgatissima. We positioned overwintering constructions (OWCs) made out of bundles of reed (Phragmites australis) outside the plantation.
• 3 Leaf beetles preferred (i) to fly towards silhouettes; (ii) OWCs positioned 1.5 m above the ground compared with ones 0.5 m above the ground; (iii) OWCs orientated vertically compared with OWCs orientated horizontally; and (iv) reeds of a diameter of approximately 5 mm.
• 4 Leaf beetle size was not correlated with reed diameter.
• 5 It is concluded that the leaf beetle P. vulgatissima selects an overwintering site based on factors at both larger and smaller scales. The possibilities for using OWCs made of reed bundles as tools to monitor and control are discussed.

     

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.     

Susceptibility of willow clones (Salix spp.) to herbivory by Phyllodecta vulgatissima (L.) and Galerucella lineola (Fab.) (Coleoptera, Chrysomelidae)

Levels of damage by mixed natural infestations of the leaf-feeding chrysomelid beetles, Phyllodecta vulgatissima (L.) (the blue willow beetle) and Galerucella lineola (Fab.) (the brown willow beetle), were determined in replicated field plots of 24 Salix clones at Long Ashton (Bristol, UK) during 1993–94. Over the same period, the host plant preferences of both chrysomelids were investigated in a standard multiple-choice laboratory procedure, where beetles were enclosed in Petri dishes with leaf discs cut from young pot-grown trees propagated from shoot cuttings taken from 20 of the 24 willow clones represented in the field study. The laboratory experiments indicated that P. vulgatissima and G. lineola had similar host plant preferences in the range of willows examined (r >0.85). In both field and laboratory, the least preferred Salix clones and hybrids were those of 5. eriocephala, followed by S. purpurea, S. burjatica, S. dasyclados and S. triandra. Clones of S. eriocephala and S. purpurea were frequently rejected altogether in laboratory tests. Most preferred were clones of S. viminalis and several hybrids of S. viminalis, S. aurita, S. caprea and S. cinerea. These results substantiate the reports that P. vulgatissima and G. lineola are deterred from feeding on willows which have relatively high concentrations of phenolic (salicylate) glucosides in the leaves. The least preferred willows, particularly S. eriocephala, S. purpurea and S. burjatica, could be of great potential value in plant breeding for resistance to these willow beetle pests.     

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.     

Phenolic glucosides as feeding cues for willow-feeding leaf beetles

The effects of individual phenolic glucosides and total glucoside fractions on the feeding behaviour of three willowfeeding leaf beetles (Coleoptera: Chrysomelidae) were tested in the laboratory. Feeding preferences of the tested leaf beetles were strongly influenced by certain phenolic glucosides which are typical secondary compounds of willows (Salicaceae:Salix). Two of the tested leaf beetles,Galerucella lineola andLochmaea capreae showed rather similar responses to glucoside treatments. Both of them were strongly stimulated by total glucoside fractions fromSalix triandra and by its major glucoside salidroside. The third species,Phatora vitellinae, was attracted most by the fractions fromS. myrsinifolia andS. pentandra, and by two related salicylate glucosides, tremulacin and salicortin. Food selection pattern of the tested beetles in the laboratory concords fairly well with their distributions in the field and with the occurrence of phenolic glucosides in their host willows. Phenolic glucoside extracts stimulated more feeding than individual pure glucosides. This indicates that different compounds have synergistic effects in the feeding behaviour of leaf beetles. Our results clearly show that willow leaf beetles select their food based on phenolic glucosides of their host plants.     

A comparative study of interspecies mating of Phratora vulgatissima and P. vitellinae using behavioural tests and molecular markers

The leaf beetle genus Phratora L. (Coleoptera: Chrysomelidae) has been used to study the ecology of host plant chemicals in herbivore preference, and the evolution of host use in chemical defence. Phratora vulgatissima and P. vitellinae are sympatric species distributed widely across Europe. Their trophic niches are largely separate due to strong differences in their host feeding preference, but they have occasionally been recorded together, feeding on Salix burjatica‘Germany’ and, only in early spring, on Populus trichocarpa (Torr & A. Gray) ‘Trichobel’. Using behavioural tests and recently developed species‐specific microsatellite markers, the intra‐ and interspecific mating of both beetle species were investigated. The microsatellite markers provided evidence that interspecific mating occurred under field conditions. Interspecific mating also took place under laboratory conditions, but less frequently than mating within species. Females of both species laid fewer eggs, and fewer eggs per clutch, when isolated with an interspecific male than with a conspecific male. Female P. vulgatissima were polyandrous, as microsatellite markers showed that their larvae were the progeny of both P. vulgatissima males that had been isolated with a single female. While only 0.55% of eggs laid in interspecific pair combinations hatched, microsatellite markers provided evidence of hybridisation between beetle species; however, these larvae died within a week when reared in a Petri dish containing ‘Germany’ and P. trichocarpa leaves. It can therefore be inferred that reproductive isolation is complete. The results are discussed in relation to species integrity and the implications for diverse mixtures of short‐rotation coppice willow plantations.     

Interactions between the willow beetle Phratora vulgatissima and different genotypes of Salix viminalis

Abstract 1 Planting of species mixtures is a strategy for the non‐chemical management of willow beetles in short‐rotation coppice willows. However, the relatively susceptible Salix viminalis genotypes and their hybrids dominate current high‐yielding willows. Interactions between Phratora vulgatissima and different genotypes of S. viminalis were examined under laboratory conditions to determine if these genotypes exhibit genetic differences in susceptibility to willow beetle damage. 2 Seven S. viminalis genotypes and four hybrids were tested for the feeding preference of adult P. vulgatissima, larval performance and within‐season plant response to manual defoliation (50% and 75%). 3 The feeding preference of adult beetles, the growth rate of larvae, and the weight at 33 days of larvae and pupae differed significantly between genotypes. Genotypes also differed significantly in their height and weight responses to mechanical defoliation. Two genotypes were taller, with longer internodes, after defoliation than were undamaged plants. Two hybrids and their S. viminalis parent showed no significant reduction in final dry weight between 0% and 50% defoliation treatments. 4 Susceptibility of genotypes to adult feeding was not correlated with their tolerance to defoliation in terms of weight or height responses; however, larval growth rate on genotypes was negatively correlated both with final height and number of leaves after 75% defoliation and with the susceptibility of genotype to adult feeding. 5 Salix viminalis showed genetic differences for all parameters tested. This suggests that the planting of a mixture of these genotypes would contain genetic differences with respect to host susceptibility to P. vulgatissima. Some parameters showed similarities between a hybrid and its parent, whereas others showed differences between siblings. This offers potential for effective breeding of desirable traits.     

Evidence for host race formation in the leaf beetle Galerucella lineola

We examined preference and performance of four Finnish Galerucella lineola F. populations on alder and willow. In standardized two‐choice laboratory feeding trials with alder and willow, only two naturally alder‐associated G. lineola populations accepted alder. Two conspecific willow‐associated populations preferred willow. These preferences seem to be unstable, however, because they can be modified by the beetles’ experience. Thus, there probably is not a complete host preference‐based isolation of alder‐ and willow‐associated G. lineola beetles in nature. In performance experiments, larvae of all four populations survived better on willow than on alder. This may indicate that willows are the ancestral hosts for G. lineola. Nevertheless, larvae of the two alder‐associated G. lineola populations survived better on alder than larvae of the two willow‐associated populations. On the other hand, larvae of the two willow‐associated populations survived better on willow than larvae of the two alder‐associated populations. This performance trade‐off suggests that G. lineola encounters different selective pressures on alders and willows. On both of them, selection probably disfavours those G. lineola genotypes that are the most successful and abundant on alternative hosts. This may reduce the effects of gene flow that is likely to occur as a consequence of incomplete host preference‐based isolation of alder‐ and willow‐associated G. lineola populations. Data from pupal weights support the idea that alder‐ and willow‐associated G. lineola populations may be genetically differentiated. Pupae of the two alder‐associated populations were heavier than those of the willow‐associated populations irrespective of whether larvae had fed on alder or on willow. Overall, our results indicate host race formation in G. lineola. This process may be enforced by the variable abundance of alders and willows in local communities.     

Spatio-temporal dynamics of willow beetle (Phratora vulgatissima) in short-rotation coppice willows grown as monocultures or a genetically diverse mixture

1 Phratora vulgatissima (Chrysomelidae) is the major pest of short-rotation coppice willows in the U.K., capable of causing severe defoliation in monoculture plantations. As this beetle shows feeding preferences between willow clones, knowledge of the spatio-temporal dynamics of P. vulgatissima is needed in order to assess the effects of mixed clonal plantings on the management of this pest. 2 Distribution patterns of adult P. vulgatissima were monitored over a season at a spatial scale of every tree or every other tree in single clone plantings (monocultures) of three willow clones and in a regular row mixture of these clones. 3 Distribution indices (deviation from Complete Spatial Randomness) demonstrated that P. vulgatissima adults were spatially aggregated on favourable clones in the monocultures and the mixture. The degree of aggregation (measured by the k-parameter of the negative binomial) differed between willow clones. Beetles were highly aggregated at the start of the season, but less so later. 4 Dispersal studies of P. vulgatissima adults over time showed a pattern of beetle infestation along rows of the preferred clones in a regularly structured mixture. In the monocultures, movement was less directional. 5 Plantation design offers some potential for more effective non-chemical pest management for P. vulgatissima. Monitoring of the pest must take account of our findings that adults occur in mobile aggregations.     

Leaf beetle grazing does not induce willow trichome defence in the coppicing willow Salix viminalis

Abstract 1 Willows are frequently attacked and defoliated by adult leaf beetles (Phratora vulgatissima L.) early in the season and the plants are then attacked again when new larvae emerge. The native willow Salix cinerea has previously been shown to respond to adult grazing by producing new leaves with an increased trichome density. Subsequent larval feeding was reduced on new leaves. This type of induced plant response may reduce insect damage and could potentially be utilized for plant protection in agricultural systems. 2 Here, we investigated if the willow species most commonly used for biomass production in short rotation coppice, Salix viminalis, also responds to adult beetle grazing by increasing trichome density. Larval performance and feeding behaviour on plants previously exposed to adult beetles was compared with that on undefoliated control plants in a greenhouse. 3 We found an overall decrease in trichome density within all the plants (i.e. trichome density was lower on new leaves compared to that for older basal leaves on S. viminalis). However, leaves of beetle defoliated plants had a higher trichome density compared to control plants. Larval growth and feeding was not affected by this difference between treatments. Larvae appeared to remove trichomes when feeding on S. viminalis, a behaviour that might explain the lack of difference between treatments.     

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.     

Phenotypic plasticity in host plant preference of the willow leaf beetle Phratora vulgatissima: the impact of experience made by adults

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THE EVOLUTION OF HOST-PLANT USE AND SEQUESTRATION IN THE LEAF BEETLE GENUS PHRATORA (COLEOPTERA: CHRYSOMELIDAE)

Leaf beetles in the genus Phratora differ in host plant use and in the chemical composition of their larval defensive secretion. Most species specialize on either poplars or willows (family Salicaceae), but two species feed on birch (family Betulaceae). Phratora vitellinae utilizes salicylates from the host plant to produce its larval secretion, which contains salicylaldehyde, while other Phratora species produce an autogenous secretion. To reconstruct the evolutionary history of host plant use and the larval secretion chemistry in this genus, we sequenced 1383 base pairs of the mt cytochrome oxidase I gene for six European and one North American Phratora species and three outgroup taxa. Bootstrap values of the complete nucleotide sequence were 99-100% for six of eight nodes in the maximum parsimony tree. They were 71% and 77% for the two other nodes. The maximum parsimony tree and the maximum likelihood tree based on nucleotide sequence showed the same relationships as a maximum parsimony tree based on the amino acid sequence. Beetle phylogeny overlapped broadly with host plant taxonomy and chemistry, and it revealed historical constraints influencing host plant use. However, there was one host shift from the willow family (Salicaceae) to the birch family (Betulaceae). The use of host plant phenol glycosides for the larval defensive secretion evolved along the lineage that led to P. vitellinae. Phratora vitellinae feeds on the taxonomically widest range of host plants, which are characterized by moderate to high levels of salicylates. The results support the hypothesis that the use of salicylates for the larval secretion evolved twice independently in chrysomeline leaf beetles.     

Geographic variation in phenotypic traits in Phratora spp. and the effects of conditioning on feeding preference

The leaf beetle genus Phratora (L.) (Coleoptera: Chrysomelidae) includes important pests of cultivated willows and poplars. The feeding preference of P. vulgatissima and P. vitellinae collected from different geographical locations was studied under laboratory conditions. There were geographic differences in the amount consumed of different willow host genotypes by each beetle species. Correlation analysis showed that, regardless of these individual differences between willow genotypes and locations, the ranking of preferred host genotypes was significantly related between locations. There were differences in the preference of host genotypes by P. vulgatissima adults and larvae after being confined to a specific willow genotype for a month, suggesting the possibility of a conditioning effect on food preference. This study also permitted the assessment of possible geographic differentiation in morphological traits. Both beetle species generally showed an increase in body size, width, and percentage fat of dry body weight with a more northerly location. These results are discussed in relation to using varietal mixtures as a pest management strategy.     

Plant-mediated effects of different Salix species on the performance of the braconid parasitoid Perilitus brevicollis

The blue willow beetle, Phratora vulgatissima, is considered to be the most damaging herbivorous pest in Salix short-rotation coppices throughout Europe. The braconid parasitoid Perilitus brevicollis is an important natural enemy of Phratora. As several different Salix species are used in coppices, I investigated the bottom-up (tritrophic) effects of Salix on the parasitoid. Three host plants were studied: the introduced fast-growing S. viminalis, which is highly susceptible to the beetle; S. dasyclados, which is introduced and moderately-resistant to the beetle; and the native slow-growing Salix cinerea, which is not currently used in coppices. The identity of the host-plant species had significant effects on parasitoid larval development time; parasitoids developed rapidly on the susceptible S. viminalis and slowly on the moderately resistant S. dasyclados. Increased development time resulted in reduced adult longevity. Host-plant species identity also affected larval survival; 57%, 64%, and 49% of the parasitoids successfully completed larval development in beetles fed S. viminalis, S. cinerea, and S. dasyclados, respectively. Parasitoid development was also correlated with the body size of their beetle host, but this effect was independent of the identity of the host-plant species. The results of this study suggest that the parasitoid has higher survival and growth rates when it parasitizes beetles feeding on the common coppice species S. viminalis, but the performance of the parasite is reduced when the beetle feeds on the moderately-resistant S. dasyclados. Conversely, the omnivorous biocontrol agents sometimes used in these systems appear to perform better on S. dasyclados compared to S. viminalis. The results of this study suggest that Perilitus parasitoids and omnivorous beetle predators may provide complementary protection to Salix and therefore be useful in coppice management.     

The influence of phenolic compounds on the suitability of three Salix species as hosts for the willow beetle Phratora vulgatissima

The phenolic glycosides salicin and salicortin were found to influence larval growth and development rates and adult feeding preference of Phratora vulgatissima in laboratory feeding studies. Salicortin was more toxic to larvae than salicin, and none of the third instar larvae fed on Salix viminalis leaves amended with 1.52% (fresh mass) salicortin pupated. Condensed tannins (proanthocyanadins) did not affect larval performance. It was concluded that Salix burjatica resistance to willow beetle is due to the high levels of salicortin which occur in leaves of this species.     

Intra- and interspecific effects of larval secretions in some chrysomelids (Coleoptera)

The larval secretions of the Chrysomelinae (Coleoptera, Chrysomelidae) are generally considered to be chemical defense substances against predators and parasites. The experiments presented in this paper provide evidence that the range of activity of these larval secretions also extends to interactions between different conspecific developmental stages and between competing phytophagous species. Four chrysomeline species were tested: Gastrophysa viridula De Geer, Phaedon cochleariae (F.), Plagiodera versicolora (Laich.), and Phratora vitellinae (L.). In the latter species, we did not test the intraspecific effect of the natural larval secretion, but the activity of its purchasable main component, salicylaldehyde (synthetic larval secretion). In G. viridula, Ph. cochleariae, and Ph. vitellinae the (synthetic) larval secretions act as oviposition deterrents against conspecific females. An investigation of the persistence of the oviposition-deterring activity of the larval secretion in G. viridula revealed that the deterrent effect is lost after 20 to 26 hours. In each of the four tested species the (synthetic) larval secretion deters feeding of conspecific adults. Since P. versicolora and Ph. vitellinae may occur on the same host plant (e.g. the willow Salix fragilis L.), the interspecific effect of the larval secretion between competing phytophagous species was examined in these two willow leaf beetles. The larval secretions of both species and salicylaldehyde, respectively, act as interspecific repellent and feeding deterrent against adults of the other species.

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Zusammenfassung Die Sekrete der Larven der Chrysomelinae (Fam. Chrysomelidae) werden im allgemeinen als chemische Abwehrstoffe gegen Prädatoren und Parasiten betrachtet. Die hier dargestellten Untersuchungen zeigen, daß die Wirkung der Larvensekrete nicht nur auf die Feindabwehr begrenzt ist. Das Wirkungsspektrum umfaßt darüber hinaus auch Interaktionen zwischen verschiedenen Entwicklungsstadien einer Art bzw. zwischen konkurrierenden phytophagen Arten. Folgende Chrysomelinenarten wurden untersucht: Gastrophysa viridula De Geer, Phaedon cochleariae (F.), Phratora vitellinae (L.) und Plagiodera versicolora (Laich.). Da das Larvensekret von Ph. vitellinae Salicylaldehyd als Hauptkomponente enthält, wurde hier-wenn nicht anders erwähnt-die Wirkung von synthetischem Salicylaldehyd getestet. Bei G. viridula, Ph. cochleariae und Ph. vitellinae wirkt das Larvensekret bzw. die authentische synthetische Substanz als hochwirksames Eiablage-Deterrens auf konspezifische Weibchen. Die eiablagehemmende Wirkung des Larvensekretes von P. versicolora wurde nicht getestet. Eine Untersuchung der Wirkungsdauer der eiblagehemmenden Aktivität des Larvensekretes von G. viridula ergab, daß die Hemmwirkung nach 20 bis 26 Stunden nicht mehr nachweisbar ist. Bei allen oben genannten vier Arten hemmt das. Larvensekret bzw. dessen authentische synthetische Substanz den Fraß von Adulten derselben Art. Diese fraßhemmende Aktivität ist bei G. viridula, Ph. cochleariae und Ph. vitellinae mindestens 18 Stunden wirksam. Das Larvensekret von P. versicolora zeigte bei 18-stündiger Versuchsdauer keine fraßhemmende Wirkung auf Adulte derselben Art; erst bei Reduktion der Versuchsdauer auf drei Stunden konnte hier eine signifikant fraßhemmende Wirkung nachgewiesen werden. P. versicolora und Ph. vitellinae können als Weidenblattkäfer an derselben Wirtspflanze auftreten. Deshalb wurde auch die interspezifische Wirkung der Larvensekrete dieser beiden Chrysomelinenarten geprüft. Das Larvensekret von Ph. vitellinae sowie auch dessen authentische synthetische Substanz (Salicylaldehyd) wirkt auf adulte P. versicolora als Repellent und Fraßhemmstoff. Ebenso wirkt umgekehrt das Larvensekret von P. versicolora abschreckend und fraßhemmend auf adulte Ph. vitellinae.

     

Willow genotype, but not drought treatment, affects foliar phenolic concentrations and leaf-beetle resistance

In a greenhouse experiment we examined the effect of willow genotype and irrigation regime (moderate drought and well‐watered) on plant growth parameters, foliar nitrogen, and phenolic concentrations, as well as on the preference and performance of the blue leaf beetle, Phratora vulgatissima (L.) (Coleoptera: Chrysomelidae). The 10 vegetatively propagated willow genotypes in the experiments were F2 full‐sibling hybrids, originated from a cross between Salix viminalis (L.) (Salicaceae) (high in condensed tannins) and Salix dasyclados (L.) (Salicaceae) (rich in phenolic glycosides). Insect bioassays were conducted on detached leaves in Petri dishes as well as with free‐living insects on intact potted plants. The 10‐week long irrigation treatments caused statistically significant phenotypic differences in the potted willow saplings. Total biomass was somewhat higher in the well‐watered treatment. The root to total biomass ratio was higher in the drought‐treatment plants. There was significant genotypic variation in foliar nitrogen concentrations, and they were higher in the drought‐treatment plants. There was also a strong genotypic variation in each of the phenolic substances analyzed. Condensed tannins, which accounted for the greatest proportion of total phenolic mass, were higher in the well‐watered treatment. There was, however, no difference in levels of the other phenolics (salicylates, cinnamic acid, flavonoids, and chlorogenic acid) between irrigation treatments. The sum of these phenolics was higher in the well‐watered treatment. There was a strong variation in P. vulgatissima larval development on different willow genotypes, and larval performance was negatively correlated with levels of salicylates and cinnamic acid. There was, however, no effect of irrigation treatment on larval performance. Phratora vulgatissima preferred to feed on well‐watered plants, and we found a preference for oviposition there, but neither feeding nor oviposition site preference was affected by willow genotype. Adult feeding and oviposition preferences were not correlated with larval performance.     

Influence of phenolglucosides and trichome density on the distribution of insects herbivores on willows

The effects of both trichome density and phenolglucoside content of leaves of 76 willow hybrids (Salix alba x fragilis) were measured to estimate their influence on the distribution of Phratora vitellinae (L.), Plagiodera versicolora Baly (Coleoptera: Chrysomelidae) and Pontania proxima (Lepeletier 1823) (Hymenoptera: Tenthredinidae) in a nursery at Gramont, Belgium.The willows showed differences in their phenolglucoside content and pilosity of leaves and are classified on these basis into four groups by a clustering method. Correlations and multiple regressions showed that these chemical and physical characteristics are good predictors of the abundance of insects. First, the abundance of larvae of Ph. vitellinae, adults of Pl. versicolora and galls of P. proxima is correlated positively with a high phenolglucoside content and a low pilosity of the leaves. Secondly, the distribution of adults of Ph. vitellinae and of larvae of Pl. versicolora is influenced by neither the chemical nor the physical leaves characteristics studied.     

Effects of host variety on blue willow beetle Phratora vulgatissima performance

Phratora vulgatissima adults exhibit feeding preferences among willow varieties, yet little is known of the effects of willow variety on larval and adult performance. The effects of host variety on adult fitness and fecundity as well as on larval mortality and development were studied under laboratory conditions for 35 willow varieties. The host variety significantly affected the number of days that adults survived and the rates of weight change. On average, males lost weight and females gained weight. The total number of eggs, eggs day¹, eggs clutch^?1 and the length of the oviposition period were also significantly affected by willow variety. Progeny from eggs laid by adults fed on the different willow varieties showed significant differences in days to pupation and pupal weight when subsequently reared on Salix × dasyclados or on the same variety as fed to the adults. However, there was no correlation between these parameters on S. × dasyclados and the variety fed to adults. The willow variety fed to larvae significantly affected larval mortality (four varieties caused 100% mortality), the shape of larval growth curves (as measured by predicted final weight and time to half the final weight), the number of days to pupation and pupal weight. There were significant positive correlations between previously determined adult P. vulgatissima feeding preferences of the 35 willow varieties and the following: number of eggs laid, length of the oviposition period, larval mortality and development and change in adult weight. There was a considerable degree of variation in these correlations and some varieties did not follow the general trend indicated by the size or sign of particular correlations, for example, having a high feeding preference ranking yet few eggs laid, low larval weight and longer time to pupation. The differences found between varieties for adult and larval performance in conjunction with previously established feeding preferences offer great potential for utilising plant resistance to P. vulgatissima as a means of strategic control. Despite the general correlation of feeding preference and performance parameters, the results found here suggest that it appears to be possible for plant breeders to circumvent this trend.