Spatial proximity between two host plant species influences oviposition and larval distribution in a leaf beetle

Everything else being equal, insect herbivores can be expected to oviposit on host plants that provide the qualitatively and quantitatively best food for larvae. However, the selection of a plant for oviposition may be influenced by such ecological factors as natural enemies, host distribution, host patch size or host patch density. We performed a field study to test whether spatial proximity between two host plant species influences the oviposition patterns and larval distribution of the alpine leaf beetle Oreina elongata. In the population studied, O. elongata oviposits and feeds on two host plants, that belong to the same family (Asteraceae): Adenostyles alliariae and Cirsiumspinosissimum. The first species contains pyrrolizidine alkaloids that are sequestered by the beetle as a chemical defence, whereas the second plant does not contain any alkaloids but has hairy and spiny leaves that might give some mechanical protection to beetle larvae.
During two consecutive summers, we quantified oviposition and larval distribution on randomly chosen C. spinosissimum that grew spatially isolated from A. alliariae, on C. spinosissimum that grew in leaf contact with A. alliariae and on A. alliariae that grew in leaf contact with C. spinosissimum (isolated A. alliariae was not considered, because it is rare in the study population). In both years, more eggs were laid on C. spinosissimum than on A. alliariae and more on those C. spinosissimum that were growing close to A. alliariae than on those growing isolated. Large numbers of larvae moved from C. spinosissimum to A. alliariae during the season. Patch size did not influence egg and larval numbers. Eggs survived better on C. spinosissimum than on A. alliariae in the field. The data suggest that C. spinosissimum may provide eggs with better protection against stormy weather. In a separate study of the same population, we found that larval performance was better on A. alliariae than on C. spinosissimum. Our present data suggest that O. elongata preferentially oviposits on plants of the species that maximizes egg survival and that grow in close proximity to plants of the species that provides better food and chemical defence.     

Geographic variation in oviposition choice of a leaf beetle: the relationship between host plant ranking, specificity, and motivation

The degree of adaptation of herbivorous insects to their local flora is an important component of the evolutionary processes that lead to host plant specialization in insects. In this study we investigated geographic variations in the oviposition preference of the leaf beetle Oreina elongata Suffrian (Coleoptera: Chrysomelidae: Chrysolini) in relation to differences in host plant specialization, in the field. We focused on the mechanisms of host choice and asked whether potential differences among populations are due to variations in host plant ranking and/or host plant specificity. We performed a combination of simultaneous choice and sequential no‐choice experiments with two of the major host plants of the beetle [Cirsium spinosissimum (L.) and Adenostyles alliariae (Gouan) (Asteraceae)]. The results suggested that spatial variation in host plant specialization has resulted in differences between populations in some aspects of the oviposition choice of O. elongata, while other aspects seem unaffected. We found no variation in host plant ranking among populations, as estimated in simultaneous choice tests. In contrast, the sequential no‐choice test indicated that host plant specificity was lower in a population that never encountered the highest ranked plant in the field. This finding agreed with our expectations, and we discuss our results in relation to the commonly used hierarchical threshold model. The results suggested that the mechanism for the differences in specificity is the variation among populations in the general motivation to oviposit, rather than quantitative differences in relative preference for the two hosts. We stress that it is essential to establish which of the two mechanisms is most important, as it will affect the probability of evolutionary change in host plant ranking.     

An alternative hibernation strategy involving sun-exposed 'hotspots', dispersal by flight, and host plant finding by olfaction in an alpine leaf beetle

Oreina cacaliae (Schrank) (Coleoptera: Chrysomelidae) has a 2‐year life cycle that it has to complete within the short warm seasons of the harsh alpine environment. Three years of field observations and experiments revealed that not all beetles overwintered in the soil next to their principal host Adenostyles alliariae (Asteraceae), as was previously assumed, but that many O. cacaliae left their host in autumn and flew to overwintering sites that were extensively sun‐exposed. In spring, these individuals became active 2 months earlier than their conspecifics that had remained in the soil close to the host plant. These early beetles flew from their hibernation sites against the direction of the prevailing wind. After a random landing in snow, they walked to the spring host Petasites paradoxus (Asteraceae) and fed on its floral stalks, the only plant parts present at that time. A few weeks later, they took flight again to locate newly emerging A. alliariae on which they would feed and deposit larvae as did individuals that had overwintered close to A. alliariae. Leaves of A. alliariae contain pyrrolizidine alkaloids (PAs), which the beetles sequester for their own defence. The dominating PA (seneciphylline) was also found to be present in the floral stalks of P. paradoxus. With additional behavioural assays in the field and laboratory, we demonstrated the importance of plant odours in the short‐range host location process. This study reveals a unique hibernation behaviour in which part of the beetle population uses exceptionally warm locations from which they emerge in spring, long before all the snow has melted. This early, but risky emergence allows them to exploit a second, highly suitable host plant, which they locate first by wind‐guided flight and then by odour‐guided walking. The well‐fed beetles then use odour again to move to their principal host plant, on which they reproduce.     

Response of a leaf beetle to two food plants,only one of which provides a sequestrable defensive chemical

Oreina elongata is a chemically defended leaf beetle. If its food plant contains pyrrolizidine alkaloids, all life stages of the beetle sequester them. However, one of the two known host-plant genera does not contain these alkaloids. In this paper we compare the adult feeding preference and larval performance of two populations, one feeding on Adenostyles alliariae (which contains alkaloids) and one on Cirsium spinosissimum (devoid of alkaloids). Adults of the population living on C. spinosissimum preferred the alkaloid-containing A. alliariae, while adults of the population feeding on A. alliariae showed no preference for either plant. On the other hand, larval growth of both populations is better on C. spinosissimum, without alkaloids. This is especially so in the population that never naturally encounters pyrrolizidine alkaloids; the population living on A. alliariae is apparently better adapted to its host's secondary compounds. The data are discussed in terms of cost of defense and trade-offs between growth and defense.     

Pyrrolizidine alkaloids of probable host-plant origin in the pronotal and elytral secretion of the leaf beetle Oreina cacaliae

Oreina cacaliae (Coleoptera, Chrysomelidae) produces in its elytral and pronotal defensive secretion seneciphylline N-oxide together with small amounts of another pyrrolizidine alkaloid tentatively identified as senecionine N-oxide. This is a strong departure from the chemical composition of the defensive secretions in related species, characterized by complex mixtures of cardenolides, synthesized by the beetles from cholesterol. It is suggested that O. cacaliae sequesters the alkaloids from its host-plant, Adenostyles leucophylla. Other specimens of O. cacaliae from far distant populations feeding on Senecio nemorensis, Petasites paradoxus or P. album also produced pyrrolizidine alkaloids, but not O. speciosissima feeding on the same food plants and producing cardenolides. In addition to pyrrolizidine alkaloids, O. cacaliae secretes ethanolamine, which is also found in all the cardenolide-producing species.     

Does Rust Infection of Willow Affect Feeding and Oviposition Behavior of Willow Leaf Beetles?

Willows are often attacked by both herbivorous insects and rust fungi. Little is known about interactions between these two willow enemies. We studied whether feeding and oviposition behavior of the willow leaf beetle Plagiodera versicolora upon the willow hybrid Salix x cuspidata is affected when the rust fungus Melampsora allii-fragilis has attacked the plant. Laboratory bioassays revealed that adult willow leaf beetles significantly avoided feeding and oviposition on rust-infected leaves when compared to healthy leaves. Further bioassays aimed to elucidate the temporal and spatial scale of effects of rust infection on feeding behavior of adults. While infected parts of leaves were avoided at all times past infection tested (8, 12, and 16 days), symptom-free parts of infected leaves were only avoided 16 days past infection. Systemic effects extended only one leaf position up and two leaf positions down from the infection site.     

Response of the elm leaf beetle to host plants induced by oviposition and feeding: the infestation rate matters

We investigated by olfactometry and feeding‐ and oviposition‐choice‐tests how the highly specialised elm leaf beetle, Xanthogaleruca luteola Müller (Coleoptera: Chrysomelidae), responds to conspecifically induced defences in the field elm Ulmus minor Miller (Ulmaceae). While egg deposition of the beetle induced elms to release volatiles attractive to the egg parasitoid Oomyzus gallerucae Fonscolombe (Hymenoptera: Eulophidae), feeding alone did not. In the present study, females of the elm leaf beetle showed preferences for the odours of twigs induced by low egg deposition and feeding over odours from uninfested twigs. In contrast, heavy infestation rendered elm odours less attractive to the beetles. Feeding and oviposition bioassays revealed an oviposition preference for leaves from uninfested twigs when compared to locally infested leaves. However, beetles preferred to feed upon systemically induced leaves compared to uninfested ones. The different preferences of the elm leaf beetle during host plant approach might be explained by a strategy that accounts for both gaining access to high quality nutrition and avoiding competition or parasitism.     

The role of olfactory cues in the sequential radiation of a gall‐boring beetle,Mordellistena convicta

1. Herbivorous insects often have close associations with specific host plants, and their preferences for mating and ovipositing on a specific host‐plant species can reproductively isolate populations, facilitating ecological speciation. Volatile emissions from host plants can play a major role in assisting herbivores to locate their natal host plants and thus facilitate assortative mating and host‐specific oviposition. 2. The present study investigated the role of host‐plant volatiles in host fidelity and oviposition preference of the gall‐boring, inquiline beetle, Mordellistena convicta LeConte (Coleoptera: Mordellidae), using Y‐tube olfactometers. Previous studies suggest that the gall‐boring beetle is undergoing sequential host‐associated divergence by utilising the resources that are created by the diverging populations of the gall fly, Eurosta solidaginis Fitch (Diptera: Tephritidae), which induces galls on the stems of goldenrods including Solidago altissima L. (Asteraceae) and Solidago gigantea Ait. 3. Our results show that M. convicta adults are attracted to galls on their natal host plant, avoid the alternate host galls, and do not respond to volatile emissions from their host‐plant stems. 4. These findings suggest that the gall‐boring beetles can orient to the volatile chemicals from host galls, and that beetles can use them to identify suitable sites for mating and/or oviposition. Host‐associated mating and oviposition likely play a role in the sequential radiation of the gall‐boring beetle.     

The effect of a foliar disease (rust) on the development of Gastrophysa viridula (Coleoptera: Chrysomelidae)

Abstract.

• 1 Gastrophysa viridula Degeer (Coleoptera: Chrysomelidae) and the pathogenic rust fungus Uromyces rumicis (Schum.) Wint. both occur on leaves of Rumex crispus L. and R.obtusifolius L. Individual stages of beetle development, and egg laying, were compared on healthy and infected leaves of each plant species in the laboratory. Oviposition choice was investigated in the field and laboratory.
• 2 Beetles reared on infected leaves of each species had greater larval mortality and slower development than those reared on healthy leaves. Although larvae feeding on infected leaves consumed up to 2.5 times more dry weight than those reared on healthy leaves, they had a lower relative growth rate and pupated at a lower weight. These changes were consistent with the reduced nutritive quality of rust-infected Rumex leaves.
• 3 Fecundity of beetles reared on infected leaves of both species was considerably reduced. Eggs laid by beetles feeding on infected R.crispus leaves also had a reduced viability.
• 4 The beetle developed consistently poorer on healthy R.crispus than on healthy R.obtusifolius throughout its life-cycle. Differences in larval performance were greater between host species than between infected and healthy leaves.
• 5 Oviposition was similar on infected and healthy R.crispus in both the laboratory and field. However, adults consumed less, and laid fewer eggs on infected than on healthy R.obtusifolius. The pattern of egg laying on different aged leaves was affected by rust infection: a greater proportion of eggs was laid on the older, infected leaves, than on the equivalent aged leaves on the healthy plants. Few larvae survived from eggs laid on rusted leaves in the field.

     

Choice,performance and heritability of performance of specialist and generalist insect herbivores towards cacalol and seneciphylline,two allelochemicals of Adenostyles alpina (Asteraceae)

We compared the effects of a sesquiterpene (ST, cacalol) and a pyrrolizidine alkaloid (PA, seneciphylline), both occurring in Adenostyles alliariae, on food choice and performance of specialist and generalist insect herbivores which are all known to feed or live on A. alliariae. In choice experiments we investigated whether the compounds were preferred, deterrent or had no effect. All specialist species Aglaostigma discolor (Hymenoptera, Tenthredinidae), Oreina cacaliae (Coleoptera, Chrysomelidae) and O. speciosissima avoided feeding when confronted with the combination of compounds. Only larvae of A. discolor avoided the single ST treatment as well. Larvae of the generalist species Callimorpha dominula (Lepidoptera, Arctiidae), Cylindrotoma distinctissima (Diptera, Tipulidae) and Miramella alpina (Caelifera, Acrididae) generally avoided feeding from PA, ST and PAST treatments. The only exception were caterpillars of C. dominula which were indiscriminate towards PA when naive, and preferred to feed on the PA treatment when they had experienced the compound before. Performance, measured as the growth of larvae on the different treatments in a no choice situation over a period of 10–17 days, was not different between treatments in the specialist leaf beetles O. cacaliae and O. speciosissima. Their avoidance of the combination treatment in the choice experiments had no obvious effect on growth when forced to feed from the treatment. In the generalist C. dominula only the high concentration combination treatment (PAST) reduced growth of the larvae due to decreased consumption. In C. distinctissima we found reduced growth in all treatments except one (PA3%). Poor growth performance in C. distinctissima was due to postingestive physiological effects of all treatments and additionally to consumption reduction in high‐dose ST treatments. Genetic variability (broad sense heritability) of growth performance metabolism varied in accordance with the specialization degree of the species. O. cacaliae, the most specialized species, had no significant heritability; O. speciosissima, the less specialized specialist, had a heritability of 0.46; C. dominula, the PA adapted generalist species, had a heritability of 0.64; C. distinctissima, the generalist with no apparent adaptations, had a heritability of 0.84.     

Performance of leaf beetle larvae on sympatric host and non-host plants

Studies asking the ability of insects to utilize novel host plants often use novel hosts that are allopatric with the insect population under investigation. However, since the outcomes of species interactions are often site-specific, such studies cannot tell us whether a plant would actually be used by a given insect population if the plant grew sympatrically with it. We therefore performed a quantitative genetics experiment to analyse the performance of larvae of the leaf beetle Oreina elongata Suffrian (Coleoptera: Chrysomelidae, Chrysomelinae) on two host and three non-host plants, collected from a site where insects and plants co-occur in the Western Alps. When raised on the non-host Petasites albus (L.), larvae were able to survive equally well as on the two hosts, Adenostyles alliariae (Gouan) and Cirsium spinosissimum (L.), whereas they did not survive on the two other non-hosts, Peucedanum ostruthium (L.) and Rumex alpinus L. On P. albus, growth rate was slightly lower and development time slightly longer than on the two hosts. We found a genotype by environment interaction only for growth rate but not for development time and survival. However, the shape of the reaction norms of growth rates suggests that it is unlikely that selection could favour the inclusion of P. albus into the host range of the study population.     

Behavioral responses of Ceratitis capitata flies to bait spray droplets and natural food

We usedF-statistics to quantify the population structure of two sympatric species of leaf beetles,Oreina cacaliae andO. speciosissima (Chrysomelinae, Coleoptera), which share the same microhabitat since they feed on the same herbaceous host plants. We measured genetic differentiation at six allozyme loci 1) among populations separated by relatively small distances (40 to 250 kilometers), 2) within each population, and 3) between sexes within populations. For both species, the populations studied are not panmictic. For each population, heterozygosities are relatively high, but the observed heterozygosities are generally lower than the expected values. Overall, within-population differentiation is high and similar for both species (F _is=0.326 forO. cacaliae and 0.332 forO. speciosissima). Additionally, populations of both species are highly differentiated (F _st=0.234 versus 0.051 forO. speciosissima). ForO. cacaliae,F _is andF _st are greater among females than among males, while forO. speciosissima,F _st is sustantially greater among the males whileF _is is slightly greater among males. Differences in gene frequency among the sexes were statistically tested using a modifiedF _st with sex as the defining category, and the sexes differed significantly with the exception of one population inO. cacaliae. Possible explanations for this difference are discussed.     

Flight dimorphism is related to survival,reproduction and mating success in the leaf beetle Oreina cacaliae

1. Alternative life histories may be maintained in populations due to variation in the costs and benefits of the underlying strategies. In this study, potential costs of dispersal by flight were investigated as an alternative life‐history strategy in the mountain‐living chrysomelid beetle Oreina cacaliae. 2. In this species, previous mark–recapture studies showed a dispersal dimorphism in both males and females. While a fraction of the population engages in flight in autumn and spring (in the following referred to as ‘flyers’), the other part does not fly (non‐flyers). Flyers emerge earlier than non‐flyers and feed on a spring host plant before the emergence of the main host plant. 3. In this study, the overwintering and dispersal locations were recorded over 7 years in the field, flyers from the spring host plant were collected, and morphology and lifetime reproductive output and survival of collected flyers and non‐flyers were compared. 4. A potential trade‐off between flight and life‐history traits was observed: flyers were smaller in size, lighter in body mass, had a lower lifetime fecundity and a higher mortality. 5. Mating experiments of field‐caught beetles in the laboratory showed that larger beetles had a higher (multiple) mating success, but there was no evidence for size‐assortative mating. It is hypothesized that one reason for small beetles to disperse by flight might be to escape competition for mates with larger non‐flyers. 6. The overwhelming quantity of beetles found on the spring host every year reveals that the flying strategy is successful, despite the costs and risks.     

Oviposition site selection on oats: the effect of plant architecture,plant and leaf age,tissue toughness,and hardness on cereal leaf beetle,Oulema melanopus

In sequentially planted oat stands, the cereal leaf beetle (CLB), Oulema melanopus (L.) (Coleoptera: Chrysomelidae: Lemini), is found in greater numbers, and lays more eggs, on later planted (younger) oats (Avena sativa L.) (Poaceae). Plant characteristics that could explain this ovipositional preference were examined in a series of experiments. Cage and open field whole plant preference tests confirmed the attraction of ovipositing females to younger oats. A cage effect illustrated the role of plant architecture (plant height) in CLB host selection. Two multiple‐choice and one no‐choice excised leaf experiments determined that characteristics of individual leaves associated with leaf insertion level (leaf number from base to apex) and age influence ovipositional site selection. Leaves of higher insertion level have higher nitrogen content, but fewer eggs are laid on those leaves. Two experiments examining the interaction between total leaf nitrogen and leaf insertion level showed that only leaf insertion level affected oviposition choice. Published literature suggests variation in secondary plant compounds cannot explain O. melanopus ovipositional preference among leaves. Grass leaves of higher insertion level have more extensively developed cells associated with tissue toughness and hardness. The data and supporting literature suggest tissue toughness and hardness are deterring oviposition on oat leaves of higher insertion level. However, newly eclosed larvae are able to feed on leaves usually avoided as oviposition sites. The explanation for this result may be a lack of correlation between host suitability and ovipositional preference.     

A quantitative genetic analysis of leaf beetle larval performance on two natural hosts: including a mixed diet

Published quantitative genetic studies of larval performance on different host plants have always compared performance on one host species or genotype vs. performance on another species or genotype. The fact that some insects may feed on more than one plant species during their development has been neglected. We executed a quantitative genetic analysis of performance with larvae of the leaf beetle Oreinaelongata, raised on each of two sympatric host plants or on a mixture of them. Growth rate was higher for larvae feeding on Adenostylesalliariae, intermediate on the mixed diet and lowest on Cirsium spinosissimum. Development time was shortest on A. alliariae, intermediate on mixed diet and longest on C. spinosissimum. Survival was higher on the mixed diet than on both pure hosts. Genetic variation was present for all three performance traits but a genotype by host interaction was found only for growth rate. However, the reaction norms for growth rate are unlikely to evolve towards an optimal shape because of a lack of heritability of growth rate in each single environment. We found no negative genetic correlations for performance traits among hosts. Therefore, our results do not support a hypothesis predicting the existence of between‐host trade‐offs in performance when both hosts are sympatric with an insect population. We conclude that the evolution of host specialized genotypes is unlikely in the study population.     

Individual variation in two host plants of the ladybird beetle,Epilachna pustulosa (Coleoptera: Coccinellidae)

Individual variation in two species of host plants (thistle,Cirsium kamtschaticum, and blue cohosh,Caulophyllum robustum) of the herbivorous ladybird beetleEpilachna pustulosa was examined under laboratory conditions for their acceptability to adult beetles as a food resource, for adult preference and for larval performance. When clones of these plants were subjected to non-choice tests using posthibernating female beetles, there was found to be significant intraspecific variation among clones in terms of their acceptability, but interspecific variation was not detected. Significant intraspecific as well as interspecific variation were frequently detected in the two host plants when clones of these plants were subjected to choice tests using posthibernating female beetles; the magnitude of interspecific plant variation for beetle preference is not necessarily larger than that of intraspecific plant variation. Individual variation across plant species with respect to beetle larval performance was also significant. A positive correlation between adult preference and larval performance is suggested across the two taxonomically remote host plant species, thistle and blue cohosh, although this needs further investigation.     

Reproductive biology of viviparous and oviparous species of the leaf beetle genus Oreina

In five species of the genus Oreina Chevrolat (Coleoptera, Chrysomelidae) we compared the size of offspring, the fecundity of the females, the timing of offspring production and female investment over the season. Two of the species, O. elongata and O. luctuosa, laid eggs, while O. cacaliae, O. gloriosa and O. variabilis gave birth to larvae. Offspring size corrected for female size was similar in the two oviparous species and in the viviparous O. cacaliae. In the two other viviparous species the larvae were two to three times bigger in relation to the female. The greater size of the offspring was not traded off for lower fecundity in these latter two species, yet the production of bigger larvae was associated with a longer laying period and thereby a spreading of reproductive investment over the season. The prediction of life history theory that higher investment in individual offspring should be traded off for lower fecundity could not be confirmed. The investigation of egg and larval development showed that in one of the oviparous species, O. luctuosa, the length of the egg stage was more variable. This corroborates the view that in this species the eggs can be retained for varying times before being laid. Greater size at birth does not necessarily lead to shortened developmental times: the larval periods of O. cacaliae, O. elongata, O. gloriosa and O. variabilis were all comparable although the larvae of the first two species were relatively smaller when laid; only the small larvae of O. luctuosa needed significantly longer for their development. For all growth parameters examined the differences between species were larger than the differences between populations. A comparison of larval growth of the oligophagous species O. cacaliae on three plant genera showed that larval growth rate is influenced by the food plant. However, the plant on which the larvae grew worst is apparently not chosen for oviposition in the field. A comparison with a phylogeny of the species based on allozymes suggests that species with similar reproductive parameters are closely related, yet that viviparity evolved independently in O. cacaliae on one hand and O. variabilis and O. gloriosa on the other.     

Development of a positive preference–performance relationship in an oligophagous beetle: adaptive learning?

The relationship between oviposition preference and larval performance is a central topic in insect–plant biology. In this study, we investigate whether the oligophagous flea beetle, Altica fragariae Nakane (Coleoptera: Chrysomelidae), exhibits a positive preference–performance relationship, and whether oviposition preference develops over time. We tested the beetles using four sympatric plant species: Duchesnea indica (Andrews) Focke (the normal host plant), Agrimonia pilosa Ledeb. (a secondary host plant), and Potentilla chinensis Ser. and Sanguisorba officinalis L. (host plants of two related Altica species) (all Rosaceae). In no‐choice experiments, both oviposition rate and offspring fitness parameters (eclosion rate, development time, and body mass) were highest on D. indica. Oviposition rate was much lower on P. chinensis than on A. pilosa, whereas offspring fitness parameters did not differ significantly between these two host plants. Offspring fitness were lowest for S. officinalis, and adult females refused to oviposit on this acceptable non‐host in a no‐choice situation. Repeated two‐choice experiments showed that the proportion of oviposition on one of the novel host plants decreased significantly over time when the alternative host plant was D. indica. In repeated two‐choice experiments using A. pilosa and P. chinensis, females mainly fed on A. pilosa but distributed their eggs equally over the two host plants, in accordance with the lack of difference in offspring fitness on those hosts. Together, these results showed that A. fragariae females develop a positive preference–performance relationship over time. We suggest that A. fragariae achieves this through adaptive learning of oviposition preference: not only does the female learn to discriminate among the host plants when there is a fitness difference for her offspring, but the female also fails to discriminate when there is no fitness difference.     

Movement patterns of an Australian chrysomelid beetle in a stand of two Eucalyptus host species

Summary We examined the movements of Chrysophtharta hectica, a eucalypt-feeding chrysomelid beetle in New South Wales, Australia, in relation to the beetle's sex, age and life-history, and to attributes of its Eucalyptus host plants. Beetle movements within the site were not influenced by beetle age or sex, but may be related to generation. Beetle distributions on the two host plant species, Eucalyptus stellulata and E. pauciflora, were generally clumped. Some of this clumping resulted from preference for E. stellulata over E. pauciflora. Clumping of beetles also occurred within host plant species; some plant individuals were consistently heavily used by beetles over the course of three years. We examined nutritional, spatial and biomass attributes of plants and found plant height and foliage production to be the best predictors of beetle numbers.     

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.