Plant resistance, plant traits, and host plant choice of the leaf-folding sawfly on the arroyo willow

Abstract. 1. The hypotheses that genetic variation in host plant resistance of the arroyo willow affected leaf folder ( Phyllocolpa sp.) (Hymenoptera: Tenthredinidae) density and that genetic variation in shoot length and leaf length was correlated with resistance were tested.
2. Willows grown in pots and exposed to ovipojsition by the leaf folding sawfly in cages had significantly different densities among clones, indicating variation in resistance caused by genetic differences among conspecific host plants.
3. There was a general correspondence between leaf folder density on potted cuttings and on the plants in the field that were the sources of cuttings.
4. In behavioural choice experiments, susceptible clones (with highest leaf fold densities) had the highest oviposition activity of female leaf folders compared to clones that were resistant to the leaf folder.
5. Clones differed significantly in shoot length and leaf length among clones grown in pots, among clones in the field, and between shoots with galls and shoots without galls on clones in the field.
6. Leaf folder density was significantly positively correlated with mean shoot length on field clones in 1985 and 1986, but was not correlated with leaf length, although leaf length and shoot length were correlated.
7. Leaf length variation among willow clones accounted for a significant portion of the variation in resistance of potted willows, but shoot length was unimportant.     

Rapid gall midge adaptation to a resistant willow genotype

Abstract 1 We conducted two experiments to investigate why a basket willow Salix viminalis L. genotype, known to be highly resistant to the leaf-roller gall midge Dasineura marginemtorquens (Bremi), should support very high gall densities in a field plantation at Tälle, south Sweden.
2 The first experiment was a field test of the hypothesis of fine-scale host adaptation in the gall midge/willow system. Support for the hypothesis would be established if midges originating from resistant willows and those originating from nearby susceptible willows differed in their abilities to initiate galls and complete development on resistant plants.
3 The objective of the second experiment was to explore whether there was a genetic basis to the trait for virulence in the midge population and to investigate any potential trade-offs this trait may entail.
4 Our results indicate that there was a fine-scaled microgeographic genetic structure to the midge population at Tälle. Midges originating from resistant plants had a heritable trait that enabled them to establish galls on resistant plants.
5 Midges able to initiate galls on the resistant genotype had longer developmental time on the susceptible genotype. This suggests that there is a physiological cost associated with being adapted to the resistant willow genotype.
6 We suggest that driving forces behind the observed host adaptation are selection imposed on the midge population by very strong willow resistance and restricted gene flow in the midge populations due to the special life history features of D. marginemtorquens .     

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

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

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

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

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

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

Is selection of host plants by Plagiodera versicolora based on plant‐related performance?

Plant-related performance may be one of the most important factors in the selection of host plants by insect herbivores. We investigated the importance of plant-related performance in host selection by the willow leaf beetle, Plagiodera versicolora (Laicharting) (Coleoptera: Chrysomelidae), on four willow species: Salix chaenomeloides Kimura, Salix eriocarpa Fr. et Sav., Salix   integra Thunb., and Salix serissaefolia Kimura (Salicaceae). Bagging experiments in the field revealed that the performance of P. versicolora adults and larvae differed significantly among willow species under enemy-free conditions and at constant densities. Egg clutch and larval abundance were positively related to adult abundance. Plagiodera versicolora adults did not discriminate strongly among willow species for feeding and oviposition. Larval performance did not differ among willow species in the presence of natural enemies, suggesting that interspecific differences in host quality were overridden by mortality from natural enemies. Adult and egg clutch abundance of P. versicolora changed seasonally despite the temporal stability of adult and larval performance under enemy-free field conditions. Thus, plant-related performance of P. versicolora adults and larvae may contribute little to population growth and temporal dynamics of host use in P. versicolora . Potential factors that reduce discrimination of P. versicolora among host willow species are discussed.     

Variation in herbivore density among host plants and its consequences for community structure

Summary The densities of four species of gall-forming sawflies were found to vary significantly among willow host plant clones. Two of the speices varied among host plants at four sites in each of three years. The other two species varied in density among host plants at most of the sites in two of the three years. Total sawfly density also varied significantly among clones. Individual species densities on willow clones were significantly positively correlated between years when all sites were combined and frequently when sites were considered separately. Most pairwise species combinations were independent in density between years, but some negative correlations existed between the stem galler and the leaf galler. Gall-former densities also were largely independent among clones within years with all sites combined and with sites considered separately. The significant correlations were nearly all positive. At all four sites the combination of significant variation in sawfly densities among willow clones in the field and independence of species densities among clones resulted in significantly different communities (relative abundance of species) among willow clones in three years. Although sawfly abundances differed substantially among the four sites, this remained true. It is argued that the pattern of community structure among clones is the result of variation in host plant quality of clones. We propose an hypothesis to account for patterns of herbivore species associations based on intrapopulation host plant variation.     

Progressive restriction of host plant exploitation along a climatic gradient: the willow psyllid Cacopsylla groenlandica in Greenland

1. Host plant selection by the endemic willow psyllid Cacopsylla groenlandica was studied at eight sites in three locations along a N–S climatic gradient in west Greenland.
2. C. groenlandica oviposited and developed on four willow species: Salix glauca, S. arctophila, S. uva-ursi and S. herbacea. Development took place on female catkins of all species and on leaf shoots of S. glauca .
3. At sites in southern Greenland, C. groenlandica fully utilized all four willow species. However, there was progressive reduction of host plant species and tissues exploited as latitude increased, such that towards the northern limit of its range the psyllid became highly specialized, feeding only on female catkins of S. glauca , despite the presence of alternative hosts. Where plant growth was tightly constrained within a restricted growing season, the demands of synchronizing with more than one species may have become too great, resulting in specialization on a single host.
4. The probable mechanisms producing specialization were between-species differences in host plant phenology, linked to variation in the growing season, set against the partially non-overlapping geographical distribution patterns of the host willows.
5. A small degree of phenological asynchrony between host plants can be advantageous for the insect. Under favourable conditions the reproductive season can be extended by using a series of host plants with staggered phenologies. Some 'early' hosts also create predator-free space. However, as asynchrony increases, the advantages disappear as the insect becomes unable to exploit the full range of host options.
6. The importance of host plant synchrony in the life histories of arctic insects is discussed.     

Shading enhances the quality of willow leaves to leaf beetles – but does it matter?

Plant shading is commonly recognised as a factor, which increases susceptibility of plants to attack by herbivorous insects. In this study we experimentally investigated the effect of host plant shading on two willow-feeding leaf beetles, Galerucella lineola feeding upon Salix phylicifolia and Phratora vitellinae feeding upon Salix myrsinifolia . Both beetle species were more abundant on potted willows growing in open habitats than on the same clones placed under the shade of trees. However, in the laboratory the food preference by adults and larval performance showed that the shaded willows are actually better food for both beetle species. On the contrary, when larvae were reared in the field under natural abiotic conditions, we found no difference in larval performance, or if any, even better performance in open habitats. Apparently, higher and more variable daily temperatures in open habitats accelerated the growth of the larvae. When adults were let to emigrate from or immigrate to potted willows, which had been grown in the same conditions but placed either in the open or shady habitats, adults preferred exposed willows. Invertebrate predators were more abundant in open habitats, but we found no differences in leaf beetle mortality by natural enemies between the habitats. Although the larval performance appeared to be approximately equal in the two habitats during the unusually warm study period, we suggest that under suboptimal temperatures the better abiotic conditions of open sites can easily override the better food provided by shaded habitats. The selection of abiotic habitat thus plays a significant role in the adaptive habitat and host plant selection of these beetles within the gradient of shadiness.     

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

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

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

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

Bottom-up cascade in a tri-trophic system: different impacts of host-plant regeneration on performance of a willow leaf beetle and its natural enemy

Abstract.  1. Plant quality can directly and indirectly affect the third trophic level. However, little attention has been paid to how changes in plant quality affect the performance of predators through trophic levels, and which herbivores or predators are affected more strongly by host-plant quality. The present study examined the effects of artificial cutting of willows on the performance of a willow leaf beetle ( Plagiodera versicolora Laicharting) and its predatory ladybird beetle ( Aiolocaria hexaspilota Hope).
2. Laboratory experiments showed that performance (survival rate, developmental time, and adult mass) of the willow leaf beetle was higher when fed with leaves of cut willows than when fed with leaves of uncut willows. Performance (developmental time and adult mass) of the predatory ladybird was also improved when it was fed on the leaf beetle larvae that had been fed on leaves of cut willows, compared with those that had been fed on leaves of uncut willows. This indicates that a bottom-up cascade occurs in the tri-trophic system.
3. In a comparison of improved performance parameters between the leaf beetle and the ladybird, regenerated willows shortened the developmental time of the willow leaf beetle more than that of the ladybird. This indicates that the impacts of willow cutting on insect performance differ between the second and third trophic levels.     

Effects of plant hybridization on herbivore-parasitoid interactions

We studied the effects of host plant hybridization on the survival and mortality of the leaf-mining moth Phyllonorycter salicifoliella on hybrid and parental willow plants in the field and in a common garden experiment. P. salicifoliella survival differed significantly among three willow taxa in the field in 1994 but not in the field in 1995 or in the common garden. Parasitism by eulophid wasps differed significantly among taxa in 1994 and appeared to account for the variation in their survival. In the field in 1995, host feeding predation varied significant among taxa. The theory of tritrophic interactions predicts that plant genotype can affect natural enemy impact, and this study supports this prediction. Significant variation in survival and eulophid parasitism was also found among genotypes within taxa in the field in both years and in the common garden experiment. The common garden results show that genetic differences in plants affect the herbivore-parasitoid interaction. Variation among years in the patterns of survival and causes of mortality among field plants suggest that genotype by environment interactions may be important. Received: 1 March 1996 / Accepted: 4 November 1996     

Trade-off in oviposition strategy: choosing poor quality host plants reduces mortality from natural enemies for a salt marsh planthopper

Abstract.  1. Both host plant nutrition and mortality from natural enemies have been predicted to significantly impact host plant selection and oviposition behaviour of phytophagous insects. It is unclear, however, if oviposition decisions maximise fitness.
2. This study examined whether the salt marsh planthopper Pissonotus quadripustulatus prefers higher quality host plants for oviposition, and if oviposition decisions are made so as to minimise mortality at the egg stage.
3. A controlled laboratory experiment and 4 years of field data were used to assess the rates of planthopper oviposition on higher quality 'green' and lower quality 'woody' stems of the host plant Borrichia frutescens . The numbers and percentages of healthy eggs and eggs that were killed by parasitoids or the host plant were recorded.
4. In all years, including the laboratory experiment, Pissonotus planthoppers laid more eggs on lower quality woody stems than on higher quality green stems. While host plant related egg mortality was higher in woody stems, the percentage of eggs parasitised was much greater in green stems. This resulted in a lower total mortality of eggs on woody stems.
5. The results of this study demonstrate that, although Pissonotus prefers lower quality host plants for oviposition, this actually increases fitness. These data seem to support the enemy free space hypothesis, and suggest that for phytophagous insects that experience the majority of mortality in the egg stage, oviposition choices may be made such that mortality is minimised.     

Oviposition site preference and larval mortality in a leaf-mining moth

Abstract. 1. The univoltine leaf-mining moth, Lithocolletis quercus Ams., is endemic to Israel, where it spends its 10.5 month larval period feeding only in the leaves of Quercus calliprinos Webb.
2. We compared patterns of egg deposition and sources of larval mortality to test whether oviposition patterns and site preferences confer an enhanced likelihood of larval survival.
3. Dominant sources of larval mortality were premature leaf abscission and death from unknown causes, whereas predation, parasitism and intraspecific interference accounted for relatively little larval mortality.
4. Eggs, and thus mines, were aggregated among leaves of host trees even though premature leaf abscission was positively correlated with density of mines per leaf. Interference competition among larvae was the only other density-dependent mortality factor.
5. Oviposition patterns within leaves mitigated the probability of death from larval interference, and probably also from early leaf abscission.
6. Despite these density-dependent mortality factors, overall probability of larval survival to pupation was independent of initial density of mines on a leaf.
7. The long larval period allows synchrony between oviposition flights and times of predictable resource availability.     

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.     

Genetic variation in resistance and tolerance to insect herbivory in Salix cordata

1. This study investigated whether sand-dune willow Salix cordata , exhibits genetic variation in resistance and tolerance to herbivory.
2. A field experiment using cuttings from nine willow clones demonstrated genetic variation in resistance to the specialist herbivore Altica subplicata , as measured by beetle densities. Willow clones differed significantly in both total biomass and leaf trichome densities, and herbivore densities were marginally correlated with both of these parameters.
3. Tolerance to herbivory was measured in a greenhouse experiment by comparing growth response of plants experiencing 50% artificial defoliation and plants experiencing no defoliation. Clones showed significant differences in tolerance to herbivory for some growth measures (changes in height and number of leaves), but not for other growth measures (stem diameter growth and final biomass).
4. Despite the significant genetic variation in both resistance and tolerance, no trade-off was found between resistance and tolerance to herbivory.     

The significance of bottom-up effects for host plant specialization in Chrysomela leaf beetles

The leaf beetle species Chrysomela lapponica, which belongs to the so‐called C. interrupta group, forms distinct allopatric populations either on willows (Salicaceae) or birches (Betulaceae). It was recently suggested that, on several occasions, host plant shifts from Salicaceae to Betulaceae occurred independently within the C. interrupta group. Our study aims to elucidate bottom‐up effects of the host plants that might have shaped the evolution of host plant specialization in the populations of C. lapponica, and thus, to shed some light on the driving forces of host shifts within the C. interrupta group, too. We compared the oviposition behaviour and performance of two C. lapponica populations, one of which has adapted to birches and the other to willows. The studies were conducted under laboratory conditions, eliminating the impact of natural enemies. Experiments involving the transfer of individuals of the birch‐specialized population to willows and vice versa with individuals of the willow‐specialized population to birches aimed to examine the plasticity in host plant use. Females of each population almost exclusively chose their natural host plant for oviposition, when offering birch and willow in dual choice experiments. When specimens of the two C. lapponica populations were reared on their natural host plants, the birch specialists suffered higher mortality, needed a longer period of development and produced less larval defensive secretion than the willow specialists. When the birch specialists were fed with willow, these performance parameters decreased even more. Other parameters, such as body weight and fecundity, did not differ between birch and willow specialists when they were fed with their natural host plant. While individuals of the birch‐specialized population could be reared on willow, all neonate larvae from the willow‐specialized population died after being transferred to birch. The significance of these bottom‐up effects for the evolution of host plant specialization in C. lapponica is discussed.     

Selection of large host plants for oviposition by a monophagous leaf beetle: nutritional quality or enemy‐free space?

Abstract.  1. Oviposition site selection is crucial for the reproductive success of herbivorous insects. According to the preference–performance hypothesis, females should oviposit on host plants that enhance the performance of their offspring. More specifically, the plant vigour hypothesis predicts that females should prefer large and vigorously growing host plants for oviposition and that larvae should perform best on these plants.
2. The present study examined whether females of the monophagous leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) prefer to oviposit on large host plant individuals of the meadow clary and whether large host plants are of higher nutritional quality than small host plants. Subsequently, it was tested whether the female preference correlates with offspring performance and survival.
3. In the field, females preferred large host plant individuals for oviposition and host plant quality, i.e. leaf nitrogen content, was significantly higher in leaves of large than of small host plants.
4. In the laboratory, larval development time was shorter on leaves of large host plant individuals than on small host plant individuals, but this could not be shown in the field.
5. However, a predator-exclusion experiment in the field resulted in a higher survival of larvae on large host plants than on small host plants when all predators had free access to the plants. On caged host plants there was no difference in survival of larvae between plant size categories.
6. It is concluded that females of C. canaliculata select oviposition sites that enhance both performance and survival of their offspring, which meets the predictions of the plant vigour hypothesis.     

Host phylogeny and nutrient content drive galler diversity and abundance on willows

1. Different groups of specialised herbivores often exhibit highly variable responses to host plant traits and phylogeny. Gall‐forming insects and mites on willows are highly adapted to their hosts and represent one of the richest communities of gallers associated with a single genus of host plants. 2. The present study evaluated the effects of host plant secondary metabolites (salicylates, flavonoids, condensed tannins), physical traits (trichome density), nutrient content (N:C) and phylogeny on the abundance and richness of gall‐forming arthropods associated with eight willow species and Populus tremula. 3. Galler abundance was affected by N:C rather than by willow defensive traits or phylogeny, suggesting that gallers respond differently to host plant traits than to less specialised guilds, such as leaf‐chewing insects. None of the studied defensive traits had a significant effect on gall abundance. Gall morphospecies richness was correlated with the host phylogeny, mainly with the nodes representing the inner division of the willow subgenus Vetrix. This suggests that the radiation of some willow taxa could have been important for the speciation of gallers associated with willows. 4. In conclusion, it is shown that whereas willow traits, such as nutrient content, appeared to affect abundances of gallers, it is probably willow radiation that drives galler speciation.