Influence of plant genotype and environment on oviposition preference and offspring survival in a gallmaking herbivore

Summary Plant resistance to insect herbivores may derive from traits influencing herbivore preference, traits influencing the suitability of the plant as a host, or both. However, the plant traits influencing host-plant selection by ovipositing insect herbivores may not completely overlap those traits that affect larval survival, and distinct traits may exhibit different levels of genetic vs. environmental control. Therefore, resource supply to the host plant could affect oviposition preference and larval performance differently in different plant genotypes. To test this hypothesis, the effects of resistance level, plant genotype, and resource supply to the host plant on oviposition preference and larval performance of a gallmaking herbivore, and on various plant traits that could influence these, were examined. Replicates of four genotypes of Solidago altissima, grown under low, medium, or high levels of nutrient supply in full sun or with medium levels of nutrients in shade, were exposed to mass-released Eurosta solidaginis. The number of plants ovipunctured was significantly affected by plant genotype and the interaction between genotype and nutrient supply to the host plant: one susceptible and one resistant genotype were more preferred, and preference tended to increase with nutrient supply in the more-preferred genotypes. The growth rate of ovipunctured plants during the oviposition period was significantly greater than that of unpunctured plants. Bud diameter (which was strongly correlated with plant growth rate), leaf area, and leaf water content were significant determinants of the percentage of plants ovipunctured, explaining 74% of the variance. The number of surviving larvae was significantly affected by plant genotype, but no effect of nutrient or light supply to the host plant was detected. The ratio of bud diameter to bud length was positively related to the percentage of ovipunctured plants that formed galls, suggesting that the accurate placement of eggs near the apical meristem by ovipositing females may be easier in short, thick buds. No significant correlation was observed between oviposition preference and larval survival at the population level. These results suggest that the plant traits affecting oviposition preference may exhibit different magnitudes of phenotypic plasticity than those affecting larval survival, and that the degree of phenotypic plasticity in plant traits affecting oviposition preference may differ among genotypes within a species.     

Floral trait expression and plant fitness in response to below- and aboveground plant–animal interactions

Although plant–animal interactions like pollination and herbivory are obviously interdependent, ecological investigations focus mainly on one kind of interaction ignoring the possible significance of the others. Plants with flowers offer an extraordinary possibility to study such mutualistic and antagonistic interactions since it is possible to measure changes in floral traits and fitness components in response to different organisms or combinations of them. In a three factorial common garden experiment we investigated single and combined effects of root herbivores, leaf herbivores and decomposers on flowering traits and plant fitness of Sinapis arvensis. Leaf herbivory negatively affected flowering traits indicating that it could significantly affect plant attractiveness to pollinators. Decomposers increased total plant biomass and seed mass indicating that plants use the nutrients liberated by decomposers to increase seed production. We suggest that S. arvensis faced no strong selection pressure from pollen limitation, for two reasons. First, reduced nutrient availability through leaf herbivory affected primarily floral traits that could be important for pollinator attraction. Second, improved nutrient supply through decomposer activity was invested in seed production and not in floral traits. This study indicates the importance of considering multiple plant–animal interactions simultaneously to understand selection pressures underlying plant traits and fitness.     

Tall herb herbivory resistance reflects historic exposure to leaf beetles in a boreal archipelago age-gradient

In this paper, we introduce the coevolution-by-coexistence hypothesis which predicts that the strength of a coevolutionary adaptation will become increasingly apparent as long as the corresponding selection from an interacting counterpart continues. Hence, evolutionary interactions between plants and their herbivores can be studied by comparing discrete plant populations with known history of herbivore colonization. We studied populations of the host plant, Filipendula ulmaria (meadow sweet), on six islands, in a Bothnian archipelago subject to isostatic rebound, that represent a spatio-temporal gradient of coexistence with its two major herbivores, the specialist leaf beetles Galerucella tenella and Altica engstroemi. Regression analyses showed that a number of traits important for insect-plant interactions (leaf concentrations of individual phenolics and condensed tannins, plant height, G. tenella adult feeding and oviposition) were significantly correlated with island age. First, leaf concentrations of condensed tannins and individual phenolics were positively correlated with island age, suggesting that plant resistance increased after herbivore colonization and continued to increase in parallel to increasing time of past coexistence, while plant height showed a reverse negative correlation. Second, a multi-choice experiment with G. tenella showed that both oviposition and leaf consumption of the host plants were negatively correlated with island age. Third, larvae performed poorly on well-defended, older host populations and well on less-defended, younger populations. Thus, no parameter assessed in this study falsifies the coevolution-by-coexistence hypothesis. We conclude that spatio-temporal gradients present in rising archipelagos offer unique opportunities to address evolutionary interactions, but care has to be taken as abiotic (and other biotic) factors may interact in a complicated way.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Fecundity and feeding of <Emphasis Type="Italic">Galerucella calmariensis</Emphasis> and <Emphasis Type="Italic">G. pusilla</Emphasis> on <Emphasis Type="Italic">Lythrum salicaria</Emphasis>

Fecundity and feeding of two introduced sibling biological control species, Galerucella calmariensis and G. pusilla (Coleoptera: Chrysomelidae) on purple loosestrife, Lythrum salicaria L. (Lythraceae) were compared at constant temperatures of 12.5, 15, 20, 25, and 27.5 °C. Larval feeding was also carried out at 30 °C, but at this temperature, larvae developed only to the L2 stage and none pupated. Thus, data for this temperature were not used in the analysis. There were significant species × temperature interactions in fecundity. Of the two species, Galerucella pusilla laid more eggs. Although egg production of both species was lowest at 12.5 °C and increased to 20 °C, at higher temperatures, the two species reacted differently. From 25 to 27.5 °C, egg production decreased for G. pusilla, but G. calmariensis fecundity peaked at 27.5 °C. Significant temperature × species × life-stage interactions were also observed in feeding. For each species, the amount of feeding varied with temperature and stage of development. Galerucella pusilla adults consumed more foliage at 15, 20, and 27.5 °C. However, at 12.5 °C G. calmariensis adults fed more than G. pusilla. G. pusilla larvae consumed an average of 25% less foliage than G. calmariensis. The lower larval consumption of G. pusilla suggests that when food is limited, G. pusilla larvae may have a higher survival rate because of its ability to complete larval development with less food and produce more progeny due to its greater fecundity. When food is not limited neither species would have a competitive advantage and both species could coexist temporally and spatially. However, since G. calmariensis larvae consumed more leaf material, the larval stage of this species would have a greater impact on purple loosestrife than G. pusilla.     

Feeding damage by larvae of the mustard leaf beetle deters conspecific females from oviposition and feeding

Herbivorous insects may be informed about the presence of competitors on the same host plant by a variety of cues. These cues can derive from either the competitor itself or the damaged plant. In the mustard leaf beetle Phaedon cochleariae (Coleoptera, Chrysomelidae), adults are known to be deterred from feeding and oviposition by the exocrine glandular secretion of conspecific co-occurring larvae. We hypothesised that the exocrine larval secretion released by feeding larvae may adsorb to the surface of Chinese cabbage leaves, and thus, convey the information about their former or actual presence. Further experiments tested the influence of leaves damaged by conspecific larvae, mechanically damaged leaves, larval frass and regurgitant on the oviposition and feeding behaviour of P. cochleariae. Finally, the effect of previous conspecific herbivory on larval development and larval host selection was assessed. Our results show that (epi)chrysomelidial, the major component of the exocrine secretion from P. cochleariae larvae, was detectable by GC-MS in surface extracts from leaves upon which larvae had fed. However, leaves exposed to volatiles of the larval secretion were not avoided by female P. cochleariae for feeding or oviposition. Thus, we conclude that secretion volatiles did not adsorb in sufficient amounts on the leaf surface to display deterrent activity towards adults. By contrast, gravid females avoided to feed and lay their eggs on leaves damaged by second-instar larvae for three days when compared to undamaged leaves. Mechanical damage of leaves and treatment of artificially damaged leaves with larval frass or regurgitant did not affect oviposition and feeding of P. cochleariae. Since no adverse effects of previous herbivory on larval development were detected, we suggest that female P. cochleariae avoid Chinese cabbage leaves damaged by feeding larvae for other reasons than escape from competition or avoidance of direct negative effects that result from consuming induced plant material.     

Does mother know best? The preference–performance hypothesis and parent–offspring conflict in aboveground–belowground herbivore life cycles

1. A substantial amount of research on host‐plant selection by insect herbivores is focused around the preference–performance hypothesis (PPH). To date, the majority of studies have primarily considered insects with aboveground life cycles, overlooking insect herbivores that have both aboveground and belowground life stages, for which the PPH could be equally applicable. 2. This study investigated the factors influencing the performance of the root‐feeding vine weevil (Otiorhynchus sulcatus) larvae and whether this was linked to the oviposition behaviour of the maternal adult living aboveground. 3. Maternal insects feeding aboveground reduced root biomass by 34% and increased root carbon by 4%. Larvae feeding on plants subjected to aboveground herbivory had reduced mass. Irrespective of the presence of maternal herbivory, larval mass was positively correlated with root biomass. 4. Larval mass was also reduced by conspecific larvae, previously feeding on roots (19% reduction). However, the mechanism underpinning this effect remains unclear, as in contrast to maternal herbivory aboveground, prior larval feeding did not significantly affect root biomass or root carbon concentrations. 5. Maternal insects did not distinguish between plants infested with larvae and those that were free of larvae, in terms of their egg‐laying behaviour. Conversely, maternal insects tended to lay eggs on plants with smaller root systems, a behaviour that is likely to negatively affect offspring performance. 6. The PPH is not supported by our findings for the polyphagous vine weevil feeding on the host plant raspberry (Rubus idaeus), and in fact our results suggest that there is the potential for strong parent–offspring conflict in this system.     

Influence of leaf herbivory,root herbivory,and pollination on plant performance in Cucurbita moschata

Abstract 1. Plants experience herbivory on many different tissues that can affect reproduction directly by damaging tissues and decreasing resource availability, or indirectly via interactions with other species such as pollinators. 2. This study investigated the combined effects of leaf herbivory, root herbivory, and pollination on subsequent damage, pollinator preference, and plant performance in a field experiment using butternut squash (Cucurbita moschata). Leaf and root herbivory were manipulated using adult and larval striped cucumber beetles (Acalymma vittatum F.), a cucurbit specialist. 3. Leaf herbivory reduced subsequent pistillate floral damage and powdery mildew (Sphaerotheca fuliginea) infection. In spite of these induced defences, the overall effect of leaf herbivory on plant reproduction was negative. Leaf herbivory reduced staminate flower production, fruit number, and seed weight. In contrast, root herbivory had a minimal impact on plant reproduction. 4. Neither leaf nor root herbivory altered pollinator visitation or floral traits, suggesting that reductions in plant performance from herbivory were as a result of direct rather than indirect effects. In addition, no measured aspect of reproduction was pollen limited. 5. Our study reveals that although leaf herbivory by the striped cucumber beetle can protect against subsequent damage, this protection was not enough to prevent the negative impacts on plant performance.     

Polyphagy and primary host plants: oviposition preference versus larval performance in the lepidopteran pest Helicoverpa armigera

Oviposition preference and several measures of offspring performance of Helicoverpa armigera (Hübner) were investigated on a subset of its host plants that were selected for their reputed importance in the field in Australia. They included cotton, pigeon pea, sweet corn, mungbean, bean and common sowthistle. Plants were at their flowering stage when presented to gravid female moths. Flowering pigeon pea evoked far more oviposition than did the other plant species and was the most preferred plant for neonate larval feeding. It also supported development of the most robust larvae and pupae, and these produced the most fecund moths. Common sowthistle and cotton were equally suitable to pigeon pea for larval development, but these two species received far fewer H. armigera eggs than did pigeon pea. Mungbean also received relatively few eggs, but it did support intermediate measures of larval growth and survival. Fewest eggs were laid on bean and it was also the least beneficial in terms of larval growth. Among the host plant species tested, only flowering pigeon pea supported a good relationship between oviposition preference of H. armigera and its subsequent offspring performance. Australian H. armigera moths are thus consistent with Indian H. armigera moths in their ovipositional behaviour and larval performance relative to pigeon pea. The results suggest that the host recognition and acceptance behaviour of this species is fixed across its geographical distribution and they support the theory that pigeon pea might be one of the primary host plants of this insect. These insights, together with published results on the sensory responses of the females to volatiles derived from the different host plant species tested here, help to explain why some plant species are primary targets for the ovipositing moths whereas others are only secondary targets of this polyphagous pest, which has a notoriously broad host range. Handling Editor: Joseph Dickens     

Plant genotype and induced responses affect resistance to herbivores on evening primrose (Oenothera biennis)

Abstract. 1. Although both genotype and induced responses affect a plant's resistance to herbivores, little is known about their relative and interactive effects. This study examined how plant genotype of a native plant (Oenothera biennis) and induced plant responses to herbivory affect resistance to, and interactions among, several herbivores. 2. In a field experiment, genetic and environmental variation among habitats led to variation in the amount of early season damage and plant quality. The pattern of variation in early season infestation by spittlebugs (Philaenus spumarius, a piercing–sucking herbivore) negatively correlated with oviposition preference by a later feeding specialist weevil (Tyloderma foveolatum, a leaf‐chewer). 3. To determine if plant genotype and induced responses to herbivory might be responsible for these field patterns, we performed no‐choice and choice bioassays using four genotypes of O. biennis that varied in resistance. Plants were induced by either spittlebugs or weevils and assays measured the responses of the same specialist weevil as well as a generalist caterpillar (Spodoptera exigua). 4. Resistance to adult weevils was largely unaffected by plant genotype, while they experienced induced resistance following damage by conspecific weevils in no‐choice assays. Caterpillars were more strongly affected by plant genotype than induced responses in both no‐choice and choice assays, but they also fed less and experienced higher mortality on plants previously damaged by weevils. In contrast to the pattern suggested by the field experiment, spittlebugs did consistently induce resistance against either weevils or caterpillars in the bioassay experiment. 5. These results support recent findings that show herbivore species can compete via induced plant responses. Additionally, a quantitative review of the literature demonstrates that plant genotype tends to be more important than interspecific competition among herbivores (plant‐mediated or otherwise) in affecting herbivore preference and performance.     

Biological control of the wetlands weed purple loosestrife (Lythrum salicaria) in the Pacific northwestern United States

Purple loosestrife (Lythrum salicaria) is an Eurasian perennial hydrophyte that has become naturalized in wetlands and in and along waterways throughout temperate North America. The ecological integrity of such areas is threatened by rapidly forming monotypic infestations that displace valued flora and diminish critical fish and wildlife habitat. The inability of physical, cultural, and chemical methods to provide adequate control of the weed has led to the development of an insect-based biological control program. The first field releases of the bud and leaf feeding beetles, Galerucella calmariensis and G. pusilla, and a root-mining weevil, Hylobius transversovittatus, were made in the United States and Canada in 1992. A total of 4740 Galerucella spp. adults were released in central Washington during 1992 and 1993 at eight sites and 471 H. transversovittatus egg inoculations were made in 1993 at three locations. Establishment of both Galerucella spp. was confirmed and Hylobius colonization was achieved.     

Leaf quality and insect herbivory in model tropical plant communities after long-term exposure to elevated atmospheric CO2

Results from laboratory feeding experiments have shown that elevated atmospheric carbon dioxide can affect interactions between plants and insect herbivores, primarily through changes in leaf nutritional quality occurring at elevated CO₂. Very few data are available on insect herbivory in plant communities where insects can choose among species and positions in the canopy in which to feed. Our objectives were to determine the extent to which CO₂-induced changes in plant communities and leaf nutritional quality may affect herbivory at the level of the entire canopy. We introduced equivalent populations of fourth instar Spodoptera eridania, a lepidopteran generalist, to complex model ecosystems containing seven species of moist tropical plants maintained under low mineral nutrient supply. Larvae were allowed to feed freely for 14 days, by which time they had reached the seventh instar. Prior to larval introductions, plant communities had been continuously exposed to either 340 l CO₂ l^–1 or to 610 l CO₂ l^–1 for 1.5 years. No major shifts in leaf nutritional quality [concentrations of N, total non-structural carbohydrates (TNC), sugar, and starch; ratios of: C/N, TNC/N, sugar/N, starch/N; leaf toughness] were observed between CO₂ treatments for any of the species. Furthermore, no correlations were observed between these measures of leaf quality and leaf biomass consumption. Total leaf area and biomass of all plant communities were similar when caterpillars were introduced. However, leaf biomass of some species was slightly greater-and for other species slightly less (e.g. Cecropia peltata)-in communities exposed to elevated CO₂. Larvae showed the strongest preference for C. peltata leaves, the plant species that was least abundant in all communites, and fed relatively little on plants species which were more abundant. Thus, our results indicate that leaf tissue quality, as described by these parameters, is not necessarily affected by elevated CO₂ under relatively low nutrient conditions. Hence, the potential importance of CO₂-induced shifts in leaf nutritional quality, as determinants of herbivory, may be overestimated for many plant communities growing on nutrient-poor sites if estimates are based on traditional laboratory feeding studies. Finally, slight shifts in the abundance of leaf tissue of various species occurring under elevated CO₂ will probably not significantly affect herbivory by generalist insects. However, generalist insect herbivores appear to become more dependent on less-preferred plant species in cases where elevated CO₂ results in reduced availability of leaves of a favoured plant species, and this greater dependency may eventually affect insect populations adversely.     

Oviposition and chemosensory stimulation of the root flies Delia radicum and D. floralis in response to plants and leaf surface extracts from resistant and susceptible Brassica genotypes

In Brassica crops differences in susceptibility to root fly attack can be largely attributed to antixenotic resistance. Plants of four genotypes (two swedes and two kales) with widely differing resistance in field trials, were compared in laboratory choice assays for their susceptibility to oviposition by the root flies Delia radicum (L.) and D. floralis (Fallen) (Diptera, Anthomyiidae). For both species the preference among the genotypes corresponded to the susceptibility of the genotypes in the field. The preference ranking in response to surrogate leaves treated with methanolic surface extracts of the four genotypes was identical to the preference among potted plants, demonstrating that chemical factors on the leaf surface mediate host preference for oviposition in these species.For both species of fly, glucosinolates are major oviposition stimulants and for D. radicum an additional, nonglucosinolate oviposition stimulant, presently called CIF, is known. We describe a procedure for chromatographic separation of glucosinolates from CIF in leaf surface extracts. In oviposition-choice assays with D. radicum, the CIF-fractions of the two swede genotypes applied to surrogate leaves received a 1.8 and 4.6 times higher proportion of eggs than the respective glucosinolate-fractions, confirming the major importance of CIF as an oviposition stimulant. The genotype of swede that was preferred by both fly species in tests with plants and methanolic leaf surface extracts, also stimulated oviposition more in tests with the glucosinolate-fractions or the CIF-fractions derived from the surface extracts, respectively. Thus, glucosinolates and CIF together account for the observed preference among the genotypes and may also be responsible for their susceptibility under field conditions. In the two kale genotypes the preference for plants or surface extracts differed from the preference among the corresponding glucosinolate- and CIF-fractions, indicating that additional, as yet unknown chemical factors may also be involved.For both groups of stimulants tarsal chemoreceptors allow electrophysiological monitoring of glucosinolate- and CIF-activity in fractionated surface extracts. For D. radicum the chemosensory activity of both glucosinolate- and CIF-fractions corresponded to the respective behavioural activity in the oviposition preference tests, suggesting that preference for oviposition among genotypes can be predicted from the electrophysiological activity of their fractions. The chemosensory response of D. floralis, in particular to the CIF-fractions, was less pronounced than the response of D. radicum, indicating interspecific differences in the perception of the major oviposition stimulants. We discuss the potential application of electrophysiological techniques in support of other screening methods used in breeding for root fly resistance in Brassica crops.     

Preference for plants damaged by conspecific larvae in ovipositing cabbage root flies: influence of stimuli from leaf surface and roots

In laboratory dual-choice assays females of the cabbage root fly, Delia radicum, prefer for oviposition plants with roots damaged by conspecific larvae to undamaged controls. Cauliflower and kale plants were inoculated with root fly eggs (25 per plant) and the hatching larvae were allowed to feed on the roots for various periods of time (1–17 days). After 4 (cauliflower) or 5 (kale) days of larval feeding the oviposition preference was most pronounced and flies laid between 64% and 68% of their eggs near plants with damaged roots. Later, with increasing damage but fewer surviving, and thus actively feeding, larvae, the magnitude of the preference declined. The preference for plants already damaged by conspecific larvae may contribute to the previously observed aggregated distribution of D. radicum eggs in Brassica crop fields.Further experiments revealed that the sensory cues inducing this oviposition preference originate from the complex consisting of the damaged roots, the surrounding substrate (soil) and associated microbes, rather than from the aerial plant parts. In choice assays using the root-substrate complex of damaged and control plants (aerial parts removed), the observed preference for damaged roots was similar to that found for the entire plant but was more pronounced. The damaged roots alone, compared to control roots, received up to 72% (cauliflower) and 75% (kale) of the eggs. By contrast, surrogate leaves sprayed with methanolic leaf surface extracts from the most preferred plants which had been damaged were not discriminated from surrogate leaved sprayed with extracts of the respective control plants. Analysis of glucosinolate levels in methanolic leaf surface extracts revealed that root damage resulted in enhanced concentrations of indole-glucosinolates on the leaf surface in kale but not in cauliflower. Although indole-glucosinolates are oviposition stimulants for the cabbage root fly, the induced changes were apparently too small to influence oviposition behaviour.     

Effects of above- and belowground herbivory on growth,pollination, and reproduction in cucumber

Plants experience unique challenges due to simultaneous life in two spheres, above- and belowground. Interactions with other organisms on one side of the soil surface may have impacts that extend across this boundary. Although our understanding of plant–herbivore interactions is derived largely from studies of leaf herbivory, belowground root herbivores may affect plant fitness directly or by altering interactions with other organisms, such as pollinators. In this study, we investigated the effects of leaf herbivory, root herbivory, and pollination on plant growth, subsequent leaf herbivory, flower production, pollinator attraction, and reproduction in cucumber (Cucumis sativus). We manipulated leaf and root herbivory with striped cucumber beetle (Acalymma vittatum) adults and larvae, respectively, and manipulated pollination with supplemental pollen. Both enhanced leaf and root herbivory reduced plant growth, and leaf herbivory reduced subsequent leaf damage. Plants with enhanced root herbivory produced 35% fewer female flowers, while leaf herbivory had no effect on flower production. While leaf herbivory reduced the time that honey bees spent probing flowers by 29%, probing times on root-damaged plants were over twice as long as those on control plants. Root herbivory increased pollen limitation for seed production in spite of increased honey bee preference for plants with root damage. Leaf damage and hand-pollination treatments had no effect on fruit production, but plants with enhanced root damage produced 38% fewer fruits that were 25% lighter than those on control plants. Despite the positive effect of belowground damage on honey bee visitation, root herbivory had a stronger negative effect on plant reproduction than leaf herbivory. These results demonstrate that the often-overlooked effects of belowground herbivores may have profound effects on plant performance.     

Herbivory and seedling performance in a fragmented temperate forest of Chile

Forest fragmentation alters plant-animal interactions, including herbivory. Relying manipulative experiments, we test if the reduction in insect herbivory associated with forest fragmentation translates into increased seedling growth and survival of three tree common species (Aristotelia chilensis, Cryptocarya alba and Persea lingue) in forest fragments and continuous forests in coastal Maulino forest, central Chile. Furthermore, we test if after protecting seedlings from herbivorous insects, plant performance is increased regardless of forest fragmentation. Nursery grown seedlings were transplanted into four forest fragments and a continuous forest during 2002. Insects, important herbivores in this forest, were excluded from half the seedlings by repeated applications of insecticides. Compared to continuous forests, in forest fragments, herbivory was reduced in all three species, seedling growth was greater in A. chilensis and C. alba but not in P. lingue, and survivorship was unaffected by herbivory or fragmentation in all three species. Protecting seedlings from insects reduced herbivory in the continuous forest to similar levels attained in the forest fragments. No change in herbivory results from by protecting seedlings in forest fragments. These results confirm that insects are important herbivores in the Maulino forest and also support the hypothesis that fragmentation can have strong indirect effects on plant communities as mediated through trophic interactions.     

Cost of trichome production and resistance to a specialist insect herbivore in <Emphasis Type="Italic">Arabidopsis lyrata</Emphasis>

Theory predicts that trade-offs between resistance to herbivory and other traits positively affecting fitness can maintain genetic variation in resistance within plant populations. In the perennial herb Arabidopsis lyrata, trichome production is a resistance trait that exhibits both qualitative and quantitative variation. Using a paternal half-sib design, we conducted two greenhouse experiments to ask whether trichomes confer resistance to oviposition and leaf herbivory by the specialist moth Plutella xylostella, and to examine potential genetic constraints on evolution of increased resistance and trichome density. In addition, we examined whether trichome production is induced by insect herbivory. We found strong positive genetic and phenotypic correlations between leaf trichome density and resistance to leaf herbivory, demonstrating that the production of leaf trichomes increases resistance to leaf damage by P. xylostella. Also resistance to oviposition tended to increase with increasing leaf trichome density, but genetic and phenotypic correlations were not statistically significant. Trichome density and resistance to leaf herbivory were negatively correlated genetically with plant size in the absence of herbivores, but not in the presence of herbivores. There was no evidence of increased trichome production after leaf damage by P. xylostella. The results suggest that trichome production and resistance to leaf herbivory are associated with a cost and that the direction of selection on resistance and trichome density depends on the intensity of herbivory.     

Herbivory below ground and biological weed control: life history of a root-boring weevil on purple loosestrife

The life history of the root-boring weevil Hylobius transversovittatus was studied in north and central Europe. The weevil develops on Lythrum salicaria, a perennial marshland plant that has become a problem weed in North America. It was found in all habitats of its host plant with the exception of permanently flooded sites. It also attacked L. salicaria in an early successional stage, devastating large parts of the storage tissue. The beetle was found in two-thirds of the field populations examined with a mean attack rate of 76.3%. Larvae developed according to a 1- or 2-year generation cycle depending on the time of oviposition. Adult beetles developing within 1 year emerged between July and October, whereas beetles with a 2-year larval period emerged within 3 weeks in July and August. Adults were long-lived and could hibernate several times. Beetles of the new generation mated right on the spot and some oviposited prior to overwintering. Hibernated females had an oviposition period from May to early September and produced 3–4 eggs/day during the peak oviposition period. Specific mortality factors were scarce. Dispersal flights ensure the regular occurrence of H. transversovittatus in scattered L. salicaria populations. The severe impact of the weevil is expected to reduce the competitive ability of its host plant after introduction into North America.     

Behavioural and chemical mechanisms of plant‐mediated deterrence and attraction among frugivorous insects

1. Herbivory often induces systemic plant responses that affect the host choice of subsequent herbivores, either deterring or attracting them, with implications for the performance of both herbivore and host plant. Combining measures of herbivore movement and consumption can efficiently provide insights into the induced plant responses that are most important for determining choice behaviour. 2. The preferences of two frugivorous stink bug species, Nezara viridula and Euschistus servus between cotton plants left undamaged or damaged by Helicoverpa zea and Heliothis virescens larvae were investigated. A novel consumer movement model was used to investigate if attraction rates or leaving rates determined preferences. Stink bug consumption rates were measured using salivary sheath flanges. Finally, the systemic induction of selected phenolic‐based and terpenoid secondary metabolites were measured from heliothine herbivory on developing cotton bolls, to investigate if they explained stink bug feeding responses. 3. Heliothine herbivory did not affect the N. viridula feeding preference. However, we found opposing effects of H. zea and H. virescens herbivory on the behaviour of E. servus. Avoidance of H. zea‐damaged plants is not obviously related to phenolic or terpenoid induction in cotton bolls; whereas a preference for H. virescens‐damaged plants may be related to reductions in chlorogenic acid in boll carpel walls. 4. The present results highlight the inferential power of measuring both consumer movement and consumption in preference experiments and combining behavioural responses with phytochemical responses. Furthermore, while plant‐mediated interactions among herbivorous insects are well studied, interactions among frugivorous species specifically have been poorly documented.     

Effects of aphid-tending Argentine ants, nitrogen enrichment and early-season herbivory on insects hosted by a coastal shrub

The presence of the exotic Argentine ant, Linepithema humile Mayr (Hymenoptera: Dolichoderinae), nitrogen enrichment, and early-season herbivory by the specialist beetle Trirhabda bacharidis (Coleoptera: Chrysomelidae) have been shown, through separate experiments, to affect the densities of insect herbivores of the coastal shrub Baccharis halimifolia (Asteraceae), in Florida. Using a fully-factorial field experiment, we examined the relative importance of all three of these factors to the six most common insect herbivore species utilizing this host plant in a West Central Florida coastal habitat. The presence of ants affected more herbivore species than either early-season herbivory by larval T. bacharidis or nitrogen enrichment. Experimental reductions of L. humile resulted in reductions of an aphid, its coccinellid predators, and adult T. bacharidis, and increases of two species of leafminers and one species of stemborer. Due to the strong negative effects of stemborer herbivory on host plant survival, the increase in stemborer abundance led to increased host plant mortality. Early-season herbivory by larval T. bacharidis only affected the abundance of aphids and their predators, both of which were more abundant on trees with reduced early-season herbivory. Nitrogen fertilization had the most limited effects and only T. bacharidis larvae achieved higher densities on fertilized trees. Our results indicate that aphid tending by the exotic L. humile affects other insects on B. halimifolia more so than herbivory by the exploitative competitor T. bacharidis or nitrogen as a limiting nutrient.     

Impact of foliar herbivory on the development of a root-feeding insect and its parasitoid

The majority of studies exploring interactions between above- and below-ground biota have been focused on the effects of root-associated organisms on foliar herbivorous insects. This study examined the effects of foliar herbivory by Pieris brassicae L. (Lepidoptera: Pieridae) on the performance of the root herbivore Delia radicum L. (Diptera: Anthomyiidae) and its parasitoid Trybliographa rapae (Westwood) (Hymenoptera: Figitidae), mediated through a shared host plant Brassica nigra L. (Brassicaceae). In the presence of foliar herbivory, the survival of D. radicum and T. rapae decreased significantly by more than 50%. In addition, newly emerged adults of both root herbivores and parasitoids were significantly smaller on plants that had been exposed to foliar herbivory than on control plants. To determine what factor(s) may have accounted for the observed results, we examined the effects of foliar herbivory on root quantity and quality. No significant differences in root biomass were found between plants with and without shoot herbivore damage. Moreover, concentrations of nitrogen in root tissues were also unaffected by shoot damage by P. brassicae larvae. However, higher levels of indole glucosinolates were measured in roots of plants exposed to foliar herbivory, suggesting that the development of the root herbivore and its parasitoid may be, at least partly, negatively affected by increased levels of these allelochemicals in root tissues. Our results show that foliar herbivores can affect the development not only of root-feeding insects but also their natural enemies. We argue that such indirect interactions between above- and below-ground biota may play an important role in the structuring and functioning of communities.