Willow leaf traits affecting host use by the leaf-gall-forming sawfly

Gall-forming insects usually have very restricted host ranges, but plant traits affecting patterns of host use have rarely been examined. The sawfly Phyllocolpa sp. (Hymenoptera: Tenthredinidae) forms leaf-roll galls on three of seven Salix (Salicaceae) species that occur together on riverbanks in central Japan. We have attempted to explain this host-use pattern by invoking three plant traits: synchrony of leaf flush with the oviposition period of the sawfly, intrinsic leaf quality as a potential larval food, and leaf morphology. Two Salix species frequently used by the sawfly, Salix eriocarpa and Salix pierotii, had similar leaf traits suitable for larval survival. The third species, Salix serissaefolia, was used relatively less often and the sawfly frequently stopped laying eggs on the plant during oviposition, suggesting ovipositional selection. S. serissaefolia had the smallest leaves, and survival of sawfly larvae was lower on S. serissaefolia than on S. eriocarpa and S. pierotii, because of gall destruction, by other herbivorous insects, and leaf-size restrictions. Among the four unused species, Salix chaenomeloides had a late leaf-flush phenology, Salix gracilistyla had inferior leaf quality, and Salix gilgiana had linear leaves; these seemed to be critical factors for non-use. Salix subfragilis was also unused, but the reason for this could not be explained by the three leaf traits studied.     

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.     

A rapid hypersensitive response associated with resistance in the willow Salix viminalis against the gall midge Dasineura marginemtorquens

Great genotypic variation in resistance against the gall midge Dasineura marginemtorquens Bremi (Diptera: Cecidomyiidae) exists within its willow host Salix viminalis L. (Salicaceae). In some resistant genotypes larvae die within 40 h after attempting to initiate galls. The present study tested the hypothesis that the hypersensitive response (HR) is involved in incompatible interactions between D. marginemtorquens and S. viminalis. By means of UV/blue light and visible light microscopy, we verified a rapidly (within 12 h after egg hatch) spreading cell death of an HR-type due to larval attack in resistant willow genotypes. Twelve h after egg hatch, the cell death had spread up to six cell layers in resistant S. viminalis genotypes whereas in susceptible genotypes only up to two cell layers were affected. In the groups of dead cells on the resistant genotypes accumulation of phenolics was observed within 24 h after egg hatch. The rapidity of the cell death induction, the early local accumulation of phenolic compounds, and the strong association of the cell death with larval mortality suggest that the formation of dead cells in resistant willow genotypes should be interpreted as a true HR.     

Preference and performance of the sawfly Diprion pini on host and non-host plants of the genus Pinus

The sawfly, Diprion pini L., is a pest of Pinus in Europe and is mainly found on P. sylvestris L. and P. nigra laricio Poiret. The relative importance of female oviposition capacity and behaviour, egg development, and larval survival on a new host plant was measured on 11 pine species. Five were natural host plants and six non-host plants, five of which are not indigenous to Europe. Oviposition choice tests showed that females discriminated between the pine species. Egg and larval development also differed between pine species. However, the female choice was not linked with hatching rate and larval development. Results of biological tests clearly indicated that there were different response patterns of D. pini life stages in relation to pine species, and these patterns were the same with insects of four different origins. We discuss the importance of each potential barrier to colonisation of a new host.     

Selection ofAzolla forms resistant to the water lily aphid,Rhopalosiphum nymphaeae Life history of Rhopalosiphum nymphaeae

Genotypic and environmentally determined differences inSalix viminalis L. suitability forDasineura marginemtorquens (Bremi) (Diptera: Cecidomyiidae) growth and survival were investigated in laboratory and field experiments. Earlier studies have documented high neonate larval mortality on certainS. viminalis genotypes. Here we present results from laboratory experiments which show no difference in larval establishment behaviour on resistant and susceptible willow genotypes. These data indicate that larvae are not able to detect the plant characteristic responsible for resistance. In a field study insect performance was measured on willow shoots of dissmilar size. Adult body size was positively correlated to willow shoot length. In turn, body size was positively correlated to potential fecundity. These results are discussed in relation to observed high densities ofD. marginemtorquens in vigorously growing willows cultivated for biomass production.     

Oviposition,larval survival and leaf damage by the willow leaf blotch miner,Micrurapteryx salicifoliella,in relation to leaf trichomes across 10 Salix species

1. Foliar trichomes clearly reduce chewing damage and efficiency of movement by some insect herbivores, but the effect of trichomes on insect oviposition is less well characterised. Trichomes are likely to have particularly strong, negative effects on species that require secure attachment of the egg to the leaf epidermis for successful transition to the feeding stage – a group that includes many leaf mining insects. 2. One such species, Micrurapteryx salicifoliella, must initially enter leaf cells directly from an egg adhered to the cuticle, but later instars can move between leaves and initiate new mines from the leaf exterior. 3. Natural patterns of occurrence by M. salicifoliella were quantified on 10 sympatric Salix species varying in trichome expression to test whether trichomes were associated with reduced oviposition, larval survival and leaf damage. 4. Mean egg density and leaf mining damage were negatively related to mean trichome density across Salix species. Survival of M. salicifoliella from egg to pupa was positively related to trichome density, suggesting that initiation of new mines by late‐instar larvae was not adversely affected by trichomes. There was no evidence that trichomes benefited leaf miner larvae indirectly by decreasing density‐dependent mortality; rather, the positive relationship between trichome density and larval survival may reflect less effective chemical defence by Salix species expressing high trichome density. 5. The results suggest that foliar trichomes serve as an effective defence against M. salicifoliella by deterring oviposition, but do not reduce the survivorship of those individuals that successfully transition from egg to larva.     

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.     

High oviposition rate ofDasineura marginemtorquens onSalix viminalis genotypes unsuitable for offspring survival

A short history of contact betweenDasineura marginemtorquens and resistant genotypes ofSalix viminalis has previously been suggested as an explanation for a poor correspondence between female oviposition preference and larval performance on certain host types. The discovery of resistant wild willows instigated a quest for an alternative hypothesis. This study presents data from laboratory and field experiments that were designed to detect variability in host preferences of individual members of the midge population. Field tests showed that the oviposition site choices of females followed the same random-natured pattern as choices of females in laboratory situations. Ovipositing females did not distinguish between willow genotypes with very high or very low larval mortality. No differences in post-alighting behavior of midges on the two willow types were discovered. Observations revealed that females usually remained and oviposited on the first plant with which they came into contact, regardless of genotype.     

Molecular phylogeny ofSalicaceae and closely relatedFlacourtiaceae: Evidence from 5.8 S, ITS 1 and ITS 2 of the rDNA

A ribosomal DNA region, including the entire 5.8S RNA gene and the internal transcribed spacers ITS 1 and ITS 2, was used for studying the phylogeny ofSalicaceae and the relationship betweenSalicaceae andFlacourtiaceae. The length of the ITS regions withinSalicaceae andFlacourtiaceae was similar to that found in other angiosperms. The GC content of both ITS regions was high, varying 62.7-72.2%. The most parsimonious tree clusters the wind-pollinatedChosenia bracteosa among theSalix species, suggesting that it should be included in the genusSalix. The grouping withinSalix leaves subg.Salix as paraphyletic, for which reason the subgeneric division is questionable.Populus was monophyletic and formed a sister group toSalix. The interspecific variation of the ITS sequences was very small inSalicaceae, which is in contradiction to the age of the group according to the evidence from fossil data.Idesia polycarpa fromFlacourtiaceae shows great sequence similarity withSalicaceae, but the analysis of 5.8S rDNA supports monophyly of the four species ofFlacourtiaceae sampled for this study.     

Fecundity and Longevity of Green Vegetable Bug,Nezara viridula, Following Parasitism byTrichopoda giacomellii

The effects of parasitism by the ArgentinianTrichopoda giacomellii(Blanchard) on reproduction and longevity of its host,Nezara viridula(L.) are reported. Parasitoid larvae suppress egg maturation, reducing by 70% the fecundity of mature female hosts during the period of larval development. Egg viability was not affected, but mating frequency was reduced by approximately 50%. When parasitized as newly eclosed adults, 84% of females fail to reproduce. In male hosts, fertility and mating frequency were not affected during the period of larval parasitoid development. In male and reproductively immature female hosts, death was coincident with, or occurred shortly after parasitoid emergence (2–4 days); in mature females, death occurred on average 2 weeks after larval parasitoid emergence. Host mortality occurred as a consequence of tissue damage incurred as the parasitoid larvae emerged from the host. Some individuals survived parasitism though no further reproductive activity (mating or oviposition) occurred. The effectiveness ofT. giacomelliias a biological control agent is discussed in relation to its impact on reproduction and survival of its host and contrasted with the action of otherTrichopodaspecies.     

Open field host selection and behavior by tamarisk beetles (Diorhabda spp.) (Coleoptera: Chrysomelidae) in biological control of exotic saltcedars (Tamarix spp.) and risks to non-target athel (T. aphylla) and native Frankenia spp.

Biological control of invasive saltcedars (Tamarix spp.) in the western U.S. by exotic tamarisk leaf beetles, Diorhabda spp., first released in 2001 after 15 years of development, has been successful. In Texas, beetles from Crete, Greece were first released in 2004 and are providing control. However, adults alight, feed and oviposit on athel (Tamarix aphylla), an evergreen tree used for shade and as a windbreak in the southwestern U.S. and México, and occasionally feed on native Frankenia spp. plants. The ability of tamarisk beetles to establish on these potential field hosts was investigated in the field. In no-choice tests in bagged branches, beetle species from Crete and Sfax, Tunisia produced 30–45% as many egg masses and 40–60% as many larvae on athel as on saltcedar. In uncaged choice tests in south Texas, adult, egg mass and larval densities were 10-fold higher on saltcedar than on adjacent athel trees after 2 weeks, and damage by the beetles was 2- to 10-fold greater on saltcedar. At a site near Big Spring, in west-central Texas, adults, egg masses and 1st and 2nd instar larvae were 2- to 8-fold more abundant on saltcedar than on athel planted within a mature saltcedar stand being defoliated by Crete beetles, and beetles were 200-fold or less abundant or not found at all on Frankenia. At a site near Lovelock, Nevada, damage by beetles of a species collected from Fukang, China was 12–78% higher on saltcedar than on athel planted among mature saltcedar trees undergoing defoliation. The results demonstrate that 50–90% reduced oviposition on athel and beetle dispersal patterns within resident saltcedar limit the ability of Diorhabda spp. to establish populations and have impact on athel in the field.     

Host selection,oviposition behaviour and leaf traits in a specialist willow sawfly on species of Salix (Salicaceae)

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Oviposition and feeding preferences of a flower-feeding weevil,Coelocephalapion aculeatum, in relation to conspecific damage to its host-plant

The oviposition and feeding preferences ofCoelocephalapion aculeatum Fall (Coleoptera: Apionidae), a host specific florivore ofMimosa pigra L. (Mimosaceae), were studied in relation to conspecific damage to its hostplant. Adults ofC. aculeatum cease ovipositing in inflorescences when the egg load reaches a number consistent with the larval carrying capacity of the inflorescence. The basis for this oviposition deterrence was examined by offering inflorescences damaged by adult feeding alone, larval feeding alone and a combination of adult feeding and oviposition. Adults preferred to oviposit on inflorescences which are not damaged by either adult feeding, larval feeding, or oviposition. No evidence for the existence of an oviposition deterring pheromone (ODP) was found. I suggest that the ability of a single host inflorescence to support the development of many larvae causes selection for the use of these oviposition deterring cues which can convey more quantitative information about the level of previous infestation than can ODPs. Adults fed a similar amount on damaged compared to undamaged inflorescences. These results assisted in the design of host range testing trials and allows predictions to be made about the effectiveness of this insect as a biological control agent.     

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

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Oviposition preferences and larval performance of a flower-feeding weevil,Coelocephalapion aculeatum, in relation to host development

The oviposition behaviour ofCoelocephalapion aculeatum Fall (Coleoptera: Apionidae), a host specific florivore exploiting an ephemeral resource, was studied in relation to host development. Immature inflorescences of its host, the weedMimosa pigra L. (Mimosaceae), were categorised into 5 developmental stages ranging from first appearance to one day before opening. The duration, weight, percent abortion, and nitrogen content of each stage were measured. Oviposition preferences ofC. aculeatum for all inflorescence stages were determined in multiple choice trials. Newly hatched, larvae were transferred into inflorescences of each stage to determine larval survival as a function of stage. Numbers of eggs laid into inflorescences generally correlated with larval survival except that oviposition was greater in stage 4 inflorescences than expected from the larval survival. Numbers of eggs laid into inflorescences generally correlated with their carrying capacity except that adults avoided stage 5 inflorescences in which larval survival was low. The evolution of oviposition preferences ofC. aculeatum appears to have been influenced by two opposing selective forces. The need to complete development before the inflorescence deteriorated would have selected for oviposition in young inflorescences. The greater availability of food and lower probability of abortion offered by older inflorescences would have selected for oviposition in these. The net outcome of these pressures has resulted in the observed oviposition strategy.     

Melampsora rust species on biomass willows in central and north-eastern Germany

Melampsora willow rusts are the most important fungal pathogens in short rotation coppices of biomass willows. In the past, breeding programmes for rust resistant biomass willows concentrated on the distinction of races within the forma specialis Melampsora larici-epitea f. sp. larici-epitea typica that colonized Salix viminalis and related clones. In a new breeding program that is based on a wider range of willow species it is necessary to identify further Melampsora species and formae specialis that are pathogens of willow species other than S. viminalis. Therefore, three stock collections with Salix daphnoides, Salix purpurea, and other shrub willow species (including S. viminalis) species were sampled in north-eastern Germany. A fourth stock collection in central Germany contributed rusts of tree willows (Salix fragilis and Salix alba) and the large shrub Salix caprea. Out of 156 rust samples, 149 were successfully sequenced for ITS rDNA. A phylogenetic analysis combining Neighbour-Joining, Maximum-Likelihood and Bayesian analysis revealed six species: Melampsora ribesii-purpureae, Melampsora allii-salicis-albae, Melampsora sp. aff. allii-fragilis, Melampsora larici-pentandrae, Melampsora larici-caprearum, and Melampsora larici-epitea. The first four species were found exclusively on the expected hosts. Melampsora larici-caprearum had a wider host range comprising S. caprea and S. viminalis hybrids. Melampsora larici-epitea can be further differentiated into two formae speciales. The forma specialis larici-epitea typica (59 samples) colonized Salix viminalis clones, Salix purpurea, Salix×dasyclados, and Salix×aquatica. In contrast to this relatively broad host range, f. sp. larici-daphnoides (65 samples) was found exclusively on Salix daphnoides. With the distinction and identification of the rust species/formae speciales it is now possible to test for race-specific resistances in a more targeted manner within the determined pairings of rust and willow species.     

Slow larval growth on a suboptimal willow results in high predation mortality in the leaf beetle Galerucella lineola

The slow growth/high mortality hypothesis predicts that herbivorous insects feeding on suboptimal host plants are subjected to higher predation mortality owing to the longer time spent in the vulnerable juvenile stages compared with conspecifics feeding on optimal plants. We tested this hypothesis for the willow-feeding leaf beetle Galerucella lineola raised on one suitable (Salix viminalis) and one unsuitable (S. dasyclados) willow species as well as on plants from an interspecific cross between the two species. Cohorts of larvae raised on caged plants (protected from enemies) and uncaged plants (exposed to enemies) were monitored daily throughout larval development in two consecutive years. Larvae raised on S. viminalis developed faster, grew larger and survived better than those raised on S. dasyclados. The suitability of the hybrid plants was intermediate to that of their parents. Our results strongly support the slow growth/high mortality hypothesis. In both years, total predation during the larval period was higher on S. dasyclados than on S. viminalis. Furthermore, the daily predation rate (i.e. the proportion of larvae preyed upon per day) was higher on S. dasyclados than on S. viminalis. When hybrid plants were included in the analysis total predation was positively correlated with both larval development time and daily predation rate. We suggest that high predation on beetles on low-quality plants is the combined result of their longer development time and elevated daily predation rate. The results are discussed in relation to the evolution of host plant selection and the paradox of sublethal plant defenses.     

Distribution of second generation European corn borer,Ostrinia nubilalis, egg masses in field corn and relationship to subsequent tunneling damage

The relationship between second generation European corn borer (Ostrinia nubilalis Hübner) egg mass numbers and subsequent field corn damage, as measured by stalk cavity numbers, was studied in 79 fields in northeastern North Carolina over three years. A mean of 0.028 egg masses per plant (645 egg masses/23400 plants) was found over the course of the study. Significant differences in oviposition rate were detected between fields and years. Ca. 85% of egg masses were deposited in a five leaf zone surrounding the primary ear; of these, 89% were found on the lower four leaves in this zone. Egg masses appeared to be distributed randomly within fields but at low rates of incidence, and oviposition was relatively uniform between sampling areas within individual fields. Under moderate to high oviposition pressure (mean number of egg masses per plant over the duration of the oviposition period >ca. 0.02), eggs laid during the early phases of the oviposition period account for more subsequent stalk damage than eggs laid during the later phases of the oviposition period. Variations in second generation egg mass numbers accounted for ca. 70% of variation in stalk cavity numbers.     

Potential of Salix as phytoextractor for Cd on moderately contaminated soils

Biomass production and Cd uptake in Salix (willow) is generally high. The aim of this study was to quantify the amounts of Cd in different plant and soil compartments in commercial Salix stands grown on different soil types and to use these data for estimation of the long-term effects of Cd removal by Salix at stem harvest on the soil Cd content by calculating Cd balances. Ten commercial short rotation willow coppice stands, planted with clone 78183 of Salix viminalis (L.) were investigated in order to include three different soil types with respect to pH. Stands with high stem biomass production generally had lower Cd concentrations in the plant and grew on more alkaline soils with low Cd availability when compared to stands with low stem biomass production. The highest output of Cd by Salix is therefore not necessarily achieved by stands with the highest yields or the highest Cd concentrations, as both these parameters have to be considered. The net removal of Cd from the plough layer by the Salix crop under different conditions varied between 2.6 and 16.5 g Cd ha^–1 year^–1. The long-term effects of Salix cultivation on soil Cd content, i.e. after 6–7 cutting cycles (25 years), were calculated and we reached the conclusion that there is a high potential in Salix to remove significant amounts of Cd at stem harvest. Salix can therefore presumably be used to counteract Cd pollution of agricultural soils in Sweden provided that the Cd in furnace ashes is taken care of.     

Preference and performance linkage of a leaf-mining moth on different Salicaceae species

The relationship between oviposition prefer-ence and offspring performance of a leaf-mining moth (Paraleucoptera sinuella) on four Salicaceae species was investigated in 1997 and 1998. We observed the egg distribution pattern on different plant species in the field and carried out oviposition experiments in the laboratory to determine the preference of ovipositing females. We also examined larval survival, pupal mass, and developmental time to compare larval performance on each plant species. Egg density in the field differed significantly among plant species. However, egg density was not correlated exactly with demonstrated oviposition preference. No larvae could develop on two Salix species. This finding indicated that larval survival is the most critical index of larval performance. Larval performance on each plant species was correlated well with oviposition preference that was revealed by a no-choice experiment in the laboratory. However, this correlation was not found in the field. These results indicate that the preference–performance linkage that was observed under laboratory conditions, was not always maintained in the field. Received: September 25, 2000 / Accepted: April 27, 2001