The distribution, abundance and host plant relationships of Salix- feeding psyllids (Homoptera: Psylloidea) in arctic Alaska

Abstract. 1. Five species of psyllid occurred on seven species of Salix at Meade River, Alaska. Studies were made on the two common species Psylla pclmeni Löw and P.phlebophyllae Hodkinson. The former feeds on the phanerophy tes Salix pulchra, S.lanata, S.alaxensis and S.glauca , the latter on the chamaephytes S.phlebophylla and S.reticulata.
2. Both P.palmeni and P.phlebophyllae had a 1-year life cycle and nymphal development took place on the female Salix catkin. The life cycle was generally closely synchronized with the period of catkin development. However, only a few eggs were laid on S.glauca
3. Seasonal perturbation of the host plant by flooding, ice movement and blown sand prevented psyllids breeding in certain areas colonized by the host plant.
4. In P.palmeni densities and 'feeding pressure', measured as biomass of psyllids per gram of catkin, on the different host plants followed the sequence S.pulchra>S.lanata> S.alaxensis > S.glauca. In P.phlebophyllae densities and feeding intensities were similar on S.phlebophylla and S.reticulata and grazing intensity was comparable with P.palmeni on S.pulchra.
5. A highly significant negative correlation was found between psyllid density and catkin dry weight in S.pulchra, S.phlebophylla and S.reticulata , suggesting that psyllid feeding is affecting catkin growth.
6. Predation of psyllid nymphs by syrphid larvae was heavy but there was no evidence of parasitism.
7. The life history strategies of the five psyllid species are discussed within the context of the constraints imposed by the arctic environment.     

Variation in resource exploitation along an altitudinal gradient: the willow psyllids (Cacopsylla spp.) on Salix lapponum

Three congeneric species of Salix -fedding psyllid Cacospsylla spp. Occur along an altitudinal gradient (988 1222 m. a.s.l.) in southern Norway. The 27-km transect covered the local altitudinal range of the three species. Each species showed a different characteristic altitudinal distribution: C. propinqua occurred over the entire altitudinal range of the transect C. palmeni was restricted to higher altitude and C brunneipennis occurred manly at lower altitudes. This pattern of local altitudinal distribution mirrored the broader geographical distributions of these psyllids along latitudinal gradients. All three species developed on willow catkins, but also made use of leaves. Species differed in their use of leaves with >10% of sampled individuals of C. brunncipennis or C. Palmeni developing on leaves compared with up to 80% in C. propinqua. The proportion of C. propinqua nymphs developing on catkins increased with altitude and was also related to the temporal and spatial availability of catkins but was not a result of competitive displacement from catkins. Nitrogen concentration of host plants did not affect the proportions of C propinqua on leaves or catkins and was not related to adult size or psyllid density on catkins. Patterns or resource use are discussed in relation to differences in the distribution and abundance of the three species along the transect.     

Experimental assemblage of novel plant–herbivore interactions: ecological host shifts after 40 million years of isolation

Geographic isolation is the first step in insect herbivore diet specialization. Such specialization is postulated to increase insect fitness, but may simultaneously reduce insect ability to colonize novel hosts. During the Paleocene‐Eocene, plants from the order Zingiberales became isolated either in the Paleotropics or in the Neotropics. During the Cretaceous, rolled‐leaf beetles diversified in the Neotropics concurrently with Neotropical Zingiberales. Using a community of Costa Rican rolled‐leaf beetles and their Zingiberales host plants as study system, we explored if previous geographic isolation precludes insects to expand their diets to exotic hosts. We recorded interactions between rolled‐leaf beetles and native Zingiberales by combining DNA barcodes and field records for 7450 beetles feeding on 3202 host plants. To determine phylogenetic patterns of diet expansions, we established 20 experimental plots in the field, in which we planted plots five exotic Zingiberales, recording beetles feeding on these exotic hosts. In the laboratory, using both native and exotic host plants, we reared a subset of insect species that had expanded their diets to the exotic plants. The original plant–herbivore community comprised 24 beetle species feeding on 35 native hosts, representing 103 plant–herbivore interactions. After exotic host plant introduction, 20 percent of the beetle species expanded their diets to exotic Zingiberales. Insects only established on exotic hosts that belong to the same plant family as their native hosts. Laboratory experiments show that beetles are able to complete development on these novel hosts. In conclusion, rolled‐leaf beetles are preadapted to expand their diets to novel host plants even after millions of years of geographic isolation.     

Host plant growth characteristics as determinants of abundance and phenology in jumping plant-lice on downy willow

1. Salix lapponum host plants at an upper altitudinal site differed significantly in size, structural density, phenology, growth performance, and spatial isolation from those growing at a lower site. 2. Plant differences were paralleled by significant differences in psyllid population density and phenology parameters, with psyllid population density, percentage of catkins occupied, and phenological development relatively lower or retarded at the upper site. Population densities at the upper site, nevertheless, remained high. 3. Plant measurements were good predictors of insect density, often explaining up to 73% of the variance in abundance among plants at a given site. 4. Sets of four plant characters identified by best subsets regression were better predictors of psyllid density and development than single factors, although differences were often not great and the combinations of characters selected by multiple regression sometimes differed from the best single predictors. 5. Best single predictors of psyllid density on catkins were measurements of plant size, particularly height, length, and basal stem diameter. Shoot density and catkin phenology were occasionally important but plant isolation and prior growth performance were less important. 6. By contrast with density, age structure of the psyllid population was predicted best from plant phenological measurements, notably catkin phenology.     

Effects of temperature on phenological synchrony and altitudinal distribution of jumping plant lice (Hemiptera: Psylloidea) on dwarf willow (Salix lapponum) in Norway

Summary.

• 1 The geographical distributions of three species of jumping plant lice (psyllids) along an altitudinal transect (988–1300 m a.s.l.) in southern Norway were restricted within the range of their host plant Salix lapponum. One species, Cacopsylla propinqua, occurred at all sampling locations between 988 and 1222 m, whereas C.palmeni was confined to higher altitudes (1153–1222 m) and C.brunneipennis was more abundant at lower altitudes (988–1101 m).
• 2 C.brunneipennis and C.palmeni developed only on female catkins. Development times of catkins and psyllids were similar (approximately 50 days) and successful psyllid development depended on close phenological synchrony with catkins.
• 3 Thermal requirements for development of female catkins were greater at low altitude (988 m) compared with higher altitude (1222 m), showing local adaptation of S.lapponum to altitude. In general, thermal requirements of psyllids were less than those of catkins at the same location. C.brunneipennis had higher thermal requirements than C.palmeni.
• 4 Field experiments, using polythene enclosures to elevate temperatures at two sites at different altitudes (by 0.6–1.4 deg. C), showed that insects had an enhanced relative rate of development under elevated temperatures compared with their host plants.
• 5 Indices of phenological synchrony were calculated from thermal requirements of psyllids and catkins. Under elevated temperatures, phenological synchrony decreased at both sites. This resulted in the subsequent development of smaller adult insects at low altitude, although at higher altitude, insects developing under elevated temperatures were larger and had a higher survival rate compared with controls.
• 6 Effects of temperature on phenological synchrony may explain the limits to the geographical range of psyllids. The consequences of climate change on psyllid populations will depend on the effects of decreased phenological synchrony on insect development and this may differ within the insect's geographical range.

     

Pollen limitation of reproductive effort in willows

Summary Pollen limitation of seed set differs from resource limitation in its implications for the evolution of floral traits. Willow flowers attract insects, but also abundantly produce wind-dispersed pollen. I demonstrated pollen limitation in single branches bearing 2–4 inflorescences (catkins) in a field experiment with five species by artificially increasing or decreasing the pollen load. Because the responses by single branches might be explained by diversion of resources to better-pollinated branches within a plant, a second experiment with one species tested both pollen limitation of whole plants and the autonomy of catkins. Seed set of single willow catkins is unaffected by experimental alterations of seed set in other catkins on the same plant. Hand-pollination of single catkins and of whole plants increased seed set to the same degree, suggesting there is little or no competition for resources between catkins only 5–10 cm apart. Thus, seed set in willows appears to be pollen limited, favoring insect pollination and the evolution of entomophilous traits. The data support previous views that willows have a dual pollination system utilizing wind and insects.     

What limits the altitudinal distribution of Craspedolepta species (Sternorrhyncha: Psylloidea) on fireweed?

Abstract.  1.  Craspedolepta nebulosa and C. subpunctata were studied on their shared host plant Epilobium angustifolium along an altitudinal transect in the Hardangervidda area of southern Norway.
2. The following was investigated: whether (a) the altitudinal distribution and abundance of each psyllid species was determined by summer heat budgets, acting directly on rates of insect development and/or through their host plant and (b) whether parasitism, predation, and interspecific competition played any role in restricting the distribution of the two species .
3.  Craspedolepta nebulosa extended to higher elevations than C. subpunctata but within their ranges neither species showed a clear altitudinal trend in abundance per plant. However, the former species grew to a smaller size with increasing elevation.
4. There was little evidence for interspecific competitive effects and observed levels of parasitism and predation were negligible across the transect.
5. Development and distribution of both species mirrored significant differences in the phenology, size, leaf number, and reproductive output of E. angustifolium with increasing altitude.
6. Distributions were related to available heat budgets acting on psyllid/plant development rates, with C. nebulosa developing more efficiently at lower heat availability.
7. Population transplant experiments supported the overall conclusions, with neither psyllid species able to establish viable populations above their current upper altitudinal range limit, despite the presence of their host plant.     

Head shape variation in cerambycid saproxylic beetles as a function of host plant selection

Saproxylic insects depend on deadwood for larval development, and a certain degree of specialization may be involved in their choice of host plants and/or wood in a particular stage of degradation. The plant species chosen for oviposition in turn act as an environmental pressure on the head morphology of larvae and it is expected that head shape plasticity varies directly with the number of woody plant species used for larval development in each insect species. We analyzed head shape variation in saproxylic beetles with respect to host plant species, maximum time of larval emergence and season of the year when insects colonized branches. Generalist species in the use of host plants showed significant variation in head shape and size. Time of emergence and season did not appear to affect head shape, although season was a determinant factor of abundance and possibly head size variation.     

Local adaptation and ecological genetics of host-plant specialization in a leaf beetle

The tendency of insect species to evolve specialization to one or a few plant species is probably a major reason for the remarkable diversity of herbivorous insects. The suggested explanations for this general trend toward specialization include a range of evolutionary mechanisms, whose relative importance is debated. Here we address two potentially important mechanisms: (i) how variation in the geographic distribution of host use may lead to the evolution of local adaptation and specialization; (ii) how selection for specialization may lead to the evolution of trade‐offs in performance between different hosts. We performed a quantitative genetic experiment of larval performance in three different populations of the alpine leaf beetle Oreina elongata reared on two of its main host plants. Due to differences in host availability, each population represents a distinctly different selective regime in terms of host use including selection for specialization on one or the other host as well as selection for utilizing both hosts during the larval stage. The results suggest that selection for specialization has lead to some degree of local adaptations in host use: both single‐host population had higher larval growth rate on their respective native host plant genus, while there was no difference between plant treatments in the two‐host population. However, differences between host plant treatments within populations were generally small and the degree of local adaptation in performance traits seems to be relatively limited. Genetic correlations in performance traits between the hosts ranged from zero in the two‐host population to significantly positive in the single‐host populations. This suggests that selection for specialization in single host populations typically also increased performance on the alternative host that is not naturally encountered. Moreover, the lack of a positive genetic correlation in the two host‐population give support for the hypothesis that performance trade‐offs between two host plants may typically evolve when a population have adapted to both these plants. We conclude that although there is selection for specialization in larval performance traits it seems as if the genetic architecture of these traits have limited the divergence between populations in relative performance on the two hosts.     

Preference and performance are correlated in the spittlebug Aphrophora pectoralis on four species of willow

Abstract 1. There was a positive correlation between oviposition and feeding preferences and offspring performance in the spittlebug Aphrophora pectoralis Matsumura (Homoptera: Cercopoidea, Aphrophoridae) on four species of willow Salix sp. (Salicaceae) growing near Sapporo, Japan. Spittlebugs preferred rapidly growing shoots where performance was highest.
2. When the effects of shoot length were removed, egg densities on willow species were associated with offspring performance on three of four species. Egg densities and survival rates were low on Salix integra and Salix miyabeana . Survival rates were high on Salix sachalinensis , which had high egg densities, and Salix hultenii , which had low egg densities.
3. Aphrophora pectoralis formed mating aggregations almost exclusively on S. sachalinensis but then dispersed to other willow species to oviposit.
4. Nymphs dispersed from the oviposition site to feed on nearby shoots within the same plant but they did not disperse to other willow plants. Nymphs had the same preference for rapidly growing shoots as ovipositing females, so they were able to refine the maternal choice by moving to larger shoots near the shoot on which they had eclosed.
5. The spittlebugs were highly aggregated at all life stages so that even at high densities only a small proportion of the most vigorously growing shoots was utilised.     

Plant-insect interactions: double-dating associated insect and plant lineages reveals asynchronous radiations

An increasing number of plant-insect studies using phylogenetic analysis suggest that cospeciation events are rare in plant-insect systems. Instead, nonrandom patterns of phylogenetic congruence are produced by phylogenetically conserved host switching (to related plants) or tracking of particular resources or traits (e.g., chemical). The dominance of host switching in many phytophagous insect groups may make the detection of genuine cospeciation events difficult. One important test of putative cospeciation events is to verify whether reciprocal speciation is temporally plausible. We explored techniques for double-dating of both plant and insect phylogenies. We use dated molecular phylogenies of a psyllid (Hemiptera)-Genisteae (Fabaceae) system, a predominantly monophagous insect-plant association widespread on the Atlantic Macaronesian islands. Phylogenetic reconciliation analysis suggests high levels of parallel cladogenesis between legumes and psyllids. However, dating using molecular clocks calibrated on known geological ages of the Macaronesian islands revealed that the legume and psyllid radiations were not contemporaneous but sequential. Whereas the main plant radiation occurred some 8 million years ago, the insect radiation occurred about 3 million years ago. We estimated that >60% of the psyllid speciation has resulted from host switching between related hosts. The only evidence for true cospeciation is in the much more recent and localized radiation of genistoid legumes in the Canary Islands, where the psyllid and legume radiations have been partially contemporaneous. The identification of specific cospeciation events over this time period, however, is hindered by the phylogenetic uncertainty in both legume and psyllid phylogenies due to the apparent rapidity of the species radiations.     

The influence of the host plant on the population dynamics of Acizzia russellae (Homoptera: Psyllidae)

Abstract. 1. The role of the host plant, Acacia karroo , in the population dynamics of the indigenous non-pest psyllid, Acizzia russellae , was investigated.
2. Population levels of A.russellae in early summer were almost 10 times greater on the regenerative foliage of pruned trees (cut back in spring) than on normal trees. During late summer and winter, populations declined far more slowly on pruned trees, so that very much higher numbers of psyllids were maintained on these trees through to the next spring. This finding, and others, suggest that natural enemies and climate are not major determinants in the population dynamics of this insect under normal conditions in the field.
3. Laboratory measurements of several chemical and physical characteristics of the foliage of pruned and normal trees did not reveal any differences which would account for the observed effects.
4. Measurements of organic nitrogen concentrations in the leaves of normal A. karroo trees in the field showed significant correlations between psyllid numbers and leaf nitrogen, but only during summer in November and January.
5. Psyllid performance in the laboratory (duration of life cycle, egg production, and dry weight of emergent females) was significantly better for psyllids reared on plants containing high amino nitrogen.
6. It is suggested that the availability of quantities of suitably high quality nutrients in the leaves of pruned A. karroo trees explains the "epidemic" population levels achieved by the psyllid on pruned plants. It seems that characteristics of the host plant impose the major limitation on psyllid population growth, which may explain the permanent low endemic population levels of this insect in the field.     

Implications of a temperature increase for host plant range: predictions for a butterfly

Although changes in phenology and species associations are relatively well‐documented responses to global warming, the potential interactions between these phenomena are less well understood. In this study, we investigate the interactions between temperature, phenology (in terms of seasonal timing of larval growth) and host plant use in the polyphagous butterfly Polygonia c‐album. We found that the hierarchy of larval performance on three natural host plants was not modified by a temperature increase as such. However, larval performance on each host plant and temperature treatment was affected by rearing season. Even though larvae performed better at the higher temperature regardless of the time of the rearing, relative differences between host plants changed with the season. For larvae reared late in the season, performance was always better on the herbaceous plant than on the woody plants. In this species, it is likely that a prolonged warming will lead to a shift from univoltinism to bivoltinism. The demonstrated interaction between host plant suitability and season means that such a shift is likely to lead to a shift in selective regime, favoring specialization on the herbaceous host. Based on our result, we suggest that host range evolution in response to temperature increase would in this species be highly contingent on whether the population undergoes a predicted shift from one to two generations. We discuss the effect of global warming on species associations and the outcome of asynchrony in rates of phenological change.     

Testing the trend towards specialization in herbivore-host plant associations using a molecular phylogeny of Tomoplagia (Diptera: Tephritidae)

Herbivorous insects are abundant and diverse and insect-host plant associations tend to be specialized and evolutionarily conserved. Some authors suggested that generalist insect lineages tend to become specialists, with host specialization leading to an evolutionary dead-end for the parasite species. In this paper, we have examined this tendency using a phylogenetic tree of Tomoplagia (Diptera: Tephritidae), a parasite of asteracean plants. We have tested the trend towards specialization in different hierarchical degrees of host specialization. The topology of the tree, the inference of ancestral hosts, and the lack of directional evolution indicated that specialization does not correspond to a phylogenetic dead-end. Although most Tomoplagia species are restricted to a single host genus, specialization does not seem to limit further host range evolution. This work emphasizes the advantages of the use of different levels of specialization and the inclusion of occasional hosts to establish a more detailed scenario for the evolution of this kind of ecological association.     

Evolution of specialization: a phylogenetic study of host range in the red milkweed beetle (Tetraopes tetraophthalmus)

Specialization is common in most lineages of insect herbivores, one of the most diverse groups of organisms on earth. To address how and why specialization is maintained over evolutionary time, we hypothesized that plant defense and other ecological attributes of potential host plants would predict the performance of a specialist root-feeding herbivore (the red milkweed beetle, Tetraopes tetraophthalmus). Using a comparative phylogenetic and functional trait approach, we assessed the determinants of insect host range across 18 species of Asclepias. Larval survivorship decreased with increasing phylogenetic distance from the true host, Asclepias syriaca, suggesting that adaptation to plant traits drives specialization. Among several root traits measured, only cardenolides (toxic defense chemicals) correlated with larval survival, and cardenolides also explained the phylogenetic distance effect in phylogenetically controlled multiple regression analyses. Additionally, milkweed species having a known association with other Tetraopes beetles were better hosts than species lacking Tetraopes herbivores, and milkweeds with specific leaf area values (a trait related to leaf function and habitat affiliation) similar to those of A. syriaca were better hosts than species having divergent values. We thus conclude that phylogenetic distance is an integrated measure of phenotypic and ecological attributes of Asclepias species, especially defensive cardenolides, which can be used to explain specialization and constraints on host shifts over evolutionary time.     

The Effect of Host-Plant Phylogenetic Isolation on Species Richness,Composition and Specialization of Insect Herbivores: A Comparison between Native and Exotic Hosts

Understanding the drivers of plant-insect interactions is still a key issue in terrestrial ecology. Here, we used 30 well-defined plant-herbivore assemblages to assess the effects of host plant phylogenetic isolation and origin (native vs. exotic) on the species richness, composition and specialization of the insect herbivore fauna on co-occurring plant species. We also tested for differences in such effects between assemblages composed exclusively of exophagous and endophagous herbivores. We found a consistent negative effect of the phylogenetic isolation of host plants on the richness, similarity and specialization of their insect herbivore faunas. Notably, except for Jaccard dissimilarity, the effect of phylogenetic isolation on the insect herbivore faunas did not vary between native and exotic plants. Our findings show that the phylogenetic isolation of host plants is a key factor that influences the richness, composition and specialization of their local herbivore faunas, regardless of the host plant origin.     

Herbivory by Strongylocoris leucocephalus (Hemiptera: Miridae) on a novel host plant Adenophora triphylla var. japonica in Japan

When monophagous and oligophagous insect herbivores colonize new areas, they sometimes use novel hosts. The availability of the new hosts can facilitate further expansion into regions beyond the geographic range of the original hosts, resulting in specialization on different host plant taxa in different parts of the herbivore's geographic range. Strongylocoris leucocephalus Linnaeus (Hemiptera: Miridae), which is a plant bug attacking Campanula spp., is widely distributed in the Eurasian continent, North Africa, and northern Japan. In the process of geographic range expansion, S. leucocephalus may incorporate novel host plants into their host range. I investigated the host plant species of S. leucocephalus in central Japan. Field observations revealed that the plant bug fed on the novel host plant Adenophora triphylla (Thunb.) A. DC. var. japonica (Regel) H. Hara (Campanulaceae), although feeding by S. leucocephalus was not associated with plant mortality. However, S. leucocephalus never fed on Campanula punctata Lam. var. hondoensis (Kitam.) Ohwi, the only Campanula species at the study site. The plant bug S. leucocephalus and the novel host plant A. triphylla var. japonica are not entirely distributed sympatrically, suggesting that the plant bug has expanded its geographic range by incorporating new hosts.     

Host-associated differences in fitness within and between populations of a seed beetle (Bruchidae): effects of plant variability

Summary Experiments were conducted with the sexually reproducing seed beetle Stator limbatus and its hosts in north-central Arizona to determine if it was substructured into units, each specialized for higher fitness on a specific host species. Unlike many studies, we incorporated scale, i.e., conducting experiments between and within beetle populations on seeds from within and between plant species. Of particular interest was whether intraspecific plant variability prevented beetle specialization within beetle populations. Results suggest that S. limbatus is specialized to certain hosts. On the palo verde Cercidium floridum, beetles originally reared from this host had significantly higher emergence compared to beetles transferred from other hosts. We did not test directly for a genetic basis for this. Alternative hypotheses of variation in symbiotic microorganisms in larval guts and maternal effects were assessed. Essentially no bacteria, yeast or protozoa were found, and maternal effects as expressed by varying egg weights were not detected; however, other microorganisms might have been present and maternal effects through inducible enzymes was possible. Caution, then, is needed in any genetic interpretations of our results. The differences on C. floridum were detected from tests between and within beetle populations. Evidence for specialization was not detected on the other hosts, Cercidium microphyllum and Acacia greggii. On the other hosts, beetles performed well regardless of their source. Significant differences were detected among individual plants of C. floridum as to the suitability of their seeds for deveoopment of S. limbatus. No such differences were detected among the other host plants. These patterns of conspecific plant variability are opposite of what is expected if plant variability prevents specialization of beetles to particular species of hosts. Thus, the data suggest seed variability among plants does not prevent specialization to host species in this system. We discuss how the patterns of host use in this study relate to the hypothesis of sympatric host race formation.     

Plant genetic differences influence herbivore community structure: evidence from a hybrid willow system

To determine the influence of plant genetic variation on community structure of insect herbivores, we examined the abundances of 14 herbivore species among six genetic classes of willow: Salix eriocephala, S. sericea, their F₁ and F₂ interspecific hybrids, and backcross hybrids to each parental species. We placed 1-year-old plants, grown from seeds generated from controlled crosses, in a common garden. During the growing season, we censused gall-inducing flies and sawflies, leaf-mining insects, and leaf-folding Lepidoptera to determine the community structure of herbivorous insects on the six genetic classes. Our results provided convincing evidence that the community structure of insect herbivores in this hybrid willow system was shaped by genetic differences among the parental species and the hybrid genetic classes. Using MANOVA, we detected significant differences among genetic classes for both absolute and relative abundance of herbivores. Using canonical discriminant analysis, we found that centroid locations describing community structure of the insect herbivores differed for each genetic class. Moreover, the centroids for the four hybrid classes were located well outside of the range between the centroids for the parental species, suggesting that more than additive genetic effects of the two parental species influenced community formation on hybrid classes. Line-cross analysis suggested that plant genetic factors responsible for structuring the herbivore community involved epistatic effects, as well as additive and dominance effects. We discuss the ramifications of these results in regard to the structure of insect herbivore communities on plants and the implications of our findings for the evolution of interspecific interactions.     

Interactions between willows and insect herbivores under enhanced ultraviolet-B radiation

We studied the effects of elevated ultraviolet-B radiation on interactions between insect herbivores and their host plants by exposing two species of phytochemically different willows, Salix myrsinifolia and S. phylicifolia, to a modulated increase in ultraviolet radiation in an outdoor experiment and monitoring the colonisation of insect herbivores on these willows. We examined the effect of increased ultraviolet-B (UV-B) radiation on (1) the quality of willow leaves, (2) the distribution and abundance of insect herbivores feeding on these willows, (3) the resulting amount of damage, and (4) the performance of insect larvae feeding on the exposed plant tissue. Six clones of each of the two willow species were grown in eight blocks for 12 weeks in the UV-B irradiation field. The clones were exposed to a constant 50% increase in UV-B radiation (simulating 20-25% ozone depletion), to a small increase in UV-A radiation or to ambient solar irradiation. We allowed colonisation on the willows by naturally occurring insects, but also introduced adults of a leaf beetle, Phratora vitellinae, a specialist herbivore on S. myrsinifolia. Increased UV-B radiation did not affect any of the measured indices of plant quality. However, numbers of P. vitellinae on S. myrsinifolia were higher in plants with UV-B treatment compared with UV-A and shade controls. In laboratory tests, growth of the second-instar larva of P. vitellinae was not affected by UV-B treatment of S. myrsinifolia, but was retarded on UV-B treated leaves of S. phylicifolia. In addition, naturally occurring insect herbivores were more abundant on willows exposed to elevated UV-B radiation compared to those grown under control treatments. In spite of the increased abundance of insect herbivores, willows treated with elevated UV-B did not suffer more herbivore damage than willows exposed to ambient solar radiation (shade control). The observed effects of UV-B on herbivore abundance, feeding and growth varied significantly due to spatial variation in environment quality, as indicated by the UV-treatment x block interaction. The results suggest that (1) environmental variation modifies the effects of UV-B radiation on plant-insect interactions and (2) specialist herbivores might be more sensitive to chemical changes in their secondary host plants (S. phylicifolia) than to changes in their primary hosts (S. myrsinifolia).