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

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

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.     

Phenolic glucosides as feeding cues for willow-feeding leaf beetles

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

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.     

Drivers of host plant shifts in the leaf beetle Chrysomela lapponica: natural enemies or competition?

1. The leaf beetle, Chrysomela lapponica, originally uses the salicyl glucosides (SGs) of its host plants to sequester salicylaldehyde, which serves as a defence against generalist enemies but attracts specialist enemies. However, some populations of C. lapponica have shifted to SG‐poor hosts, and their secretions do not contain salicylaldehyde. 2. In was suggested that beetles shift to SG‐poor hosts to escape from specialist enemies. To test this hypothesis, we compared field mortality between two populations of C. lapponica that were associated with SG‐rich willow, Salix myrsinifolia (Kola Peninsula and Finland) and two populations that fed on SG‐poor willows, S. glauca (Ural) and S. caprea (Belarus). 3. Mortality from generalist enemies was significantly higher in Belarus than in three other populations, whereas mortality from specialists did not differ among populations. A specialist predator (syrphid fly larvae, Parasyrphus nigritarsis) and specialist parasitoids (phorid flies, Megaselia spp.) were attracted to the secretions of larvae reared on both SG‐rich and SG‐poor hosts. 4. Feeding on leaves of S. caprea and S. myrsinifolia both previously damaged by leaf puncturing and by the larvae of potentially competing species Chrysomela vigintipunctata, decreased the weight and prolonged the development of C. lapponica. 5. Thus, populations of C. lapponica that have shifted to SG‐poor willow species did not obtain enemy‐free space because specialist enemies have developed adaptations to herbivores that switched to a novel host plant. We suggest that in some populations host plant shift was favoured by interspecific competition with the early season SG‐using specialist, C. vigintipunctata.     

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

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

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.     

Chlorogenic acid as an antiherbivore defence of willows against leaf beetles

In the leaves of 13 Finnish willow species, the content of a phenolic, chlorogenic acid, was found to vary from 0 up to 18 mg g^–1 D.W. Effects of pure chlorogenic acid on insect feeding behaviour were tested using four common leaf beetle species which are in the field mainly found on willows with low-chlorogenic acid leaves. One species, Lochmaea capreae L., was invariably deterred by pure chlorogenic acid applied in naturally occurring concentrations on the willow leaves. Accordingly, in 2-choice laboratory feeding trials L. capreae was found to prefer low-chlorogenic acid leaves of four willow species over high-chlorogenic acid leaves of Salix pentandra L. and S. myrsinifolia Salisb. When presented on the leaves of S. phylicifolia L, pure chlorogenic acid inhibited also the feeding by Phratora polaris Sp.-Schn. Instead, chlorogenic acid had no significant effect on Ph. polaris when it was presented on the leaves of another willow S. cinerea L. In laboratory, Ph. polaris did not show general preference for willow species with low chlorogenic acid content in their leaves. Thus, the response of Ph. polaris to chlorogenic acid seems to depend on the plant species. Apparently variation in other traits such as leaf hairyness may easily override the potential effect of chlorogenic acid content on Ph. polaris. To two other leaf beetle species, Galerucella lineola F. and Plagiodera versicolora Laich., chlorogenic acid is an ineffective deterrent even at unnaturally high concentrations. In laboratory, G. lineola and P. versicolora did not prefer willows with low chlorogenic acid content in their leaves. Thus, among four studied leaf beetle species, only L. capreae seems to be clearly affected by this phenolic. Therefore, overall importance of chlorogenic acid as a defence against willow-feeding leaf beetles appears to be very limited.     

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.     

Host race formation in the leaf-mining moth Acrocercops transecta (Lepidoptera: Gracillariidae)

Genetic differentiation in ecological traits plays an important role in the reproductive isolation of phytophagous insects. The present study aims to elucidate the genetic changes involved during the process of host shifts, by combining analyses for (1) host adaptations, (2) pre‐ and postmating isolation, and (3) phylogeney among populations, using a leaf‐mining moth, Acrocercops transecta. This species is associated with Juglans ailanthifolia and Lyonia ovalifolia. Transplantation of the larvae demonstrated that the Juglans‐associated population completely failed to survive on Lyonia, whereas the Lyonia‐associated population survived on Juglans as well as on Lyonia. Females of respective host‐associated populations oviposited on their natal host plant only. An mtDNA‐based phylogeny clearly separated the Lyonia‐associated population from the Juglandaceae‐associated population, and indicated that the Lyonia‐associated population once evolved from the Juglandaceae‐associated population. These results indicate that the processes of host shifting from juglandaceous species to Lyonia involved genetic changes both in larval ability to use host plants and in host preference of females. The derived Lyonia‐associated population has retained the potential to assimilate the ancestral host, Juglandaceae. Mating between the two host‐associated populations was successful for both directions of crossing, and there were no significant differences in egg hatchability between hybrids and control crosses. No adults emerged when the F₁ hybrid larvae were maintained on Lyonia; however, on Juglans the F₁ hybrid larvae grew to adulthood as well as in the control, suggesting a lack of genomic incompatibilities between the two host‐associated populations. In conclusion, the results showed that the two host‐associated populations are host races that are partially reproductively isolated, and that the differences in performance and preference function as strong barriers against gene flow between the host races. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93 , 135–145.     

Size class structure and tree dispersion patterns in old-growth cedar-hemlock forests of the northern Rocky Mountains (USA)

Summary The relationship between the food selection of four leaf beetle species (Phratora vitellinae, Plagiodera versicolora, Lochmaea capreae, Galerucella lineola) and the phenolic glycosides of willow (Salix spp.) leaves was tested in laboratory food choice experiments. Four willow species native to the study area (Eastern Finland) and four introduced, cultivated willows were tested.The willow species exhibited profound differences in their phenolic glycoside composition and total concentration. The food selection patterns of the leaf beetles followed closely the phenolic glycoside spectra of the willow species. Both the total amount and the composition of phenolic glycosides affected the feeding by the beetles. Phenolic glycosides apparently have both stimulatory and inhibitory influences on leaf beetle feeding depending on the degree of adaptation of a particular insect. Very rare glycosides or exceptional combination of several glycoside types seem to provide certain willow species with high level of resistance against most herbivorous insects. Analogously the average absolute amount of leaf beetle feeding was lower on the introduced willows than on the native species to which the local herbivores have a good opportunity to become adapted.     

Oviposition preference and larval performance of monarch butterflies (Danaus plexippus) on two invasive swallow-wort species

The potential of two invasive herbaceous vines Vincetoxicum nigrum (L.) Moench and Vincetoxicum rossicum (Kleopow) Barbar. (Asclepiadaceae) to reduce monarch butterfly (Danaus plexippus L.) (Lepidoptera: Nymphalidae, Danainae) populations was investigated by evaluating oviposition selection in adult monarch butterflies and larval feeding preference in choice tests comparing the native host plant of monarch butterflies, Asclepias syriaca L. (Asclepiadaceae) and the two non‐indigenous Vincetoxicum species. In both choice and no‐choice tests, no eggs were oviposited on either of the two Vincetoxicum species whereas over 66 eggs per female were oviposited on A. syriaca plants. All first instar larvae allowed to feed on A. syriaca for 48 h survived while a significantly lower proportion survived on V. rossicum (44%) and V. nigrum (14%). Mean weight of larvae that did survive on the Vincetoxicum species was significantly lower than the mean weight of larvae that fed on A. syriaca. The mean weight of surviving larvae, however, did not differ between the two Vincetoxicum species. The mean proportion of leaves consumed by larvae feeding on A. syriaca was significantly greater than the mean proportion of leaves consumed by larvae feeding on either Vincetoxicum species. Findings from this research indicate that V. rossicum and V. nigrum are not viable hosts of monarch butterflies and are likely to pose little direct threat to their populations as oviposition sinks. The ability of these highly aggressive plants, however, to out‐compete and displace the native host of monarchs, A. syriaca, may pose a more serious threat. The potential of monarch populations to adapt to the two Vincetoxicum species as host plants over the long‐term is discussed.     

Temporal and host-related variation in frequencies of genes that enable Phyllotreta nemorum to utilize a novel host plant, Barbarea vulgaris

The flea beetle, Phyllotreta nemorum L. (Coleoptera: Chrysomelidae), is an intermediate specialist feeding on a small number of plants within the family Brassicaceae. The most commonly used host plant is Sinapis arvensis L., whereas the species is found more rarely on Cardaria draba (L.) Desv., Barbarea vulgaris R.Br., and cultivated radish (Raphanus sativus L.). The interaction between flea beetles and Barbarea vulgaris ssp. arcuata (Opiz.) Simkovics seems to offer a good opportunity for experimental studies of coevolution. The plant is polymorphic, as it contains one type (the P‐type) that is susceptible to all flea beetle genotypes, and another type (the G‐type) that is resistant to some genotypes. At the same time, the flea beetle is also polymorphic, as some genotypes can utilize the G‐type whereas others cannot. The ability to utilize the G‐type of B. vulgaris ssp. arcuata is controlled by major dominant genes (R‐genes). The present investigation measured the frequencies of flea beetles with R‐genes in populations living on different host plants in 2 years (1999 and 2003). Frequencies of beetles with R‐genes were high in populations living on the G‐type of B. vulgaris ssp. arcuata in both years. Frequencies of beetles with R‐genes were lower in populations living on other host plants, and declining frequencies were observed in five out of six populations living on S. arvensis. Selection in favour of R‐genes in populations living on B. vulgaris is the most likely mechanism to account for the observed differences in the relative abundance of R‐genes in flea beetle populations utilizing different host plants. A geographic mosaic with differential levels of interactions between flea beetles and their host plants was demonstrated.     

Adaptive change and conservatism in host specificity in two local populations of the thistle-feeding ladybird beetle Epilachna niponica

We investigated geographic differences in the host specificity of Epilachna niponica Lewis (Coleoptera: Coccinellidae). The Yuwaku population feeds mainly on Cirsium matsumurae Nakai (Asteraceae) and secondarily on Cirsrium kagamontanum Nakai. The Asiu population, located 150 km away from the Yuwaku, feeds exclusively on C. ashiuense Yokoyama et T. Shimizu. Under laboratory conditions, we examined the differences between the two populations in adult feeding acceptance, adult feeding preference, and larval performance, using several closely related thistle species and varieties, including their native hosts. In the Asiu population, adult beetles clearly avoided the host of the Yuwaku population, C. kagamontanum, and no larvae were able to complete their development, whereas in the Yuwaku population, adults accepted and even preferred it to some other thistle species, and about 10% of first instar larvae became adults. This indicates that the Yuwaku population evolved its feeding preference and physiological adaptation to C. kagamontanum through a utilization of this low‐ranked host under natural conditions. Apart from C. kagamontanum, the two populations showed a similar host susceptibility pattern, indicating that this ladybird beetle has a conserved hierarchy in feeding preference and growth performance. We also observed adult leaf choice behavior when given different thistle species, and found that difference in biting rate after palpation determined the leaf areas consumed, implying that factors on the leaf surface played an important role in the choice.     

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

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

Post hibernation dispersal of three leaf-eating beetles (Coleoptera: Chrysomelidae) colonising cultivated willows and poplars

1 We studied the spring dispersal of three common chrysomelids, from overwintering habitats into cultivated willow and poplar coppices at four sites in southern England over 2 years. 2 Adult Galerucella lineola, Phratora vulgatissima and P. vitellinae overwintered under the bark of mature trees or in other niches that simulated this habitat, within a few hundred metres of the coppice plantation. Relatively few beetles remained in the coppice fields during the winter. 3 Phratora vitellinae at the poplar sites emerged several weeks later than G. lineola and P. vulgatissima at the willow sites, reflecting the later leafing of poplar compared to willow. For all species, dispersal was by flight, with most activity during warm periods. Dispersal continued for several weeks for the willow- feeding species but was shorter for P. vitellinae. 4 All three species initially colonized the edge of the coppice field. Typically, 80% or more of the beetles colonising a plantation were within 8 m of the edge. Both Phratora spp. accumulated in the plantation edge zone for several weeks before leaving this area and colonising the crop interior. 5 The patterns of dispersal and colonization identified by this study may facilitate chrysomelid management practices in infested short rotation coppice that avoid the need for insecticide applications over the entire plantation. Keywords Chrysomelidae, Galerucella lineola, Phratora vitellinae, Phratora vulgatissima, poplar, short rotation coppice, willow, winter dispersal.     

Beetle assemblages on willow trees: do phenolic glycosides matter?

Herbivorous beetles were sampled in central Slovakia and in the Tatra Mountains of northern Slovakia from seven different Salix species which are partly characterized by smooth leaves containing phenolic glycosides and partly by hairy leaves containing tannins. The collection included about 8000 individuals representing 129 species. Of the 129 species, 77 species are able to use the willows as hosts; the remaining 52 ‘tourist’ species comprise less than 3% of the collected individuals. The data on species richness and abundance did not generally support the ‘feeding specialization’ hypothesis. The proportion of specialized (= monophagous and oligophagous) beetles feeding on willows of both morphological and biochemical groups was in the same range. Host plants of the two groups could support high diversity and high evenness values, even when leaf characteristics and plant chemicals largely influenced species assemblage. The region in which the willow trees grow had a considerable impact on host plant use. Generalist beetle species predominated in central Slovakia. By contrast, in the Tatra Mountains, specialist feeders which are able to use phenolic glycosides to their advantage were predominant. The number of species and the total density of individuals collected from willows containing phenolic glycosides (S. fragilis and S. purpurea) did not usually vary between the two regions. In contrast, fewer species and individuals were found in the Tatra Mountains when they settled on willow species containing tannins (S. caprea and S. cinerea). Also, the phylogenetic status of host plants affected species assemblages. In central Slovakia willow species of the subgenus Vetrix (S. purpurea, S. caprea and S. cinerea) generally showed a higher beetle diversity (H_s) than species of the subgenus Salix (S. fragilis, S. alba and S. triandra), although both subgenera comprise species of both morphological and biochemical groups. Furthermore, when the analysis was restricted to beetles of central Slovakia, which should be most adapted to their host plants (i.e. catkin feeders and phyllophages in the adult and larval stage), the phylogenetic status was found to be more important than any single leaf character measured.     

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.     

Host-plant preference of Brachypterolus pulicarius, an inadvertently introduced biological control insect of toadflaxes

Brachypterolus pulicarius (L.) (Coleoptera: Kateridae) is an inadvertently introduced biological control agent that can reduce seed set in two North American invasive species, yellow (Linaria vulgaris P. Mill.) (Scrophulariaceae) and Dalmatian toadflax (Linaria genistifolia (L.) P. Mill. ssp. dalmatica). The beetles are more common on yellow toadflax than on Dalmatian toadflax. To understand their distribution on the two host plants, we investigated whether they prefer one host to the other and whether individuals aggregate toward conspecifics. In field and laboratory experiments where beetles were presented with a choice of both toadflax species, B. pulicarius sampled from both host plants preferred yellow toadflax. However, in the laboratory experiment, beetles collected from Dalmatian toadflax showed a weaker preference for yellow toadflax than beetles collected from yellow toadflax. In the field experiment, all beetle populations sampled showed similar preferences. When given a choice between yellow toadflax plants with and without trapped adult B. pulicarius, beetles preferred plants with conspecifics, suggesting aggregation toward beetle pheromones or host‐plant volatiles induced by beetle activity. These results do not support the current practice of redistributing North American B. pulicarius onto Dalmatian toadflax because of their preference for yellow toadflax.     

Gene genealogies reveal cryptic species and host preferences for the pine fungal pathogen Grosmannia clavigera

Grosmannia clavigera is a fungal pathogen of pine forests in western North America and a symbiotic associate of two sister bark beetles: Dendroctonus ponderosae and D. jeffreyi. This fungus and its beetle associate D. ponderosae are expanding in large epidemics in western North America. Using the fungal genome sequence and gene annotations, we assessed whether fungal isolates from the two beetles inhabiting different species of pine in epidemic regions of western Canada and the USA, as well as in localized populations outside of the current epidemic, represent different genetic lineages. We characterized nucleotide variations in 67 genomic regions and selected 15 for the phylogenetic analysis. Using concordance of gene genealogies and distinct ecological characteristics, we identified two sibling phylogenetic species: Gc and Gs. Where the closely related Pinus ponderosa and P. jeffreyi are infested by localized populations of their respective beetles, Gc is present. In contrast, Gs is an exclusive associate of D. ponderosae mainly present on its primary host‐tree P. contorta; however, in the current epidemic areas, it is also found in other pine species. These results suggest that the host‐tree species and the beetle population dynamics may be important factors associated with the genetic divergence and diversity of fungal partners in the beetle‐tree ecosystems. Gc represents the original G. clavigera holotype, and Gs should be described as a new species.