Trade-off in investment between dispersal and ingestion capability in phytophagous insects and its ecological implications

In population ecology, dispersal plays a fundamental role, but is potentially costly. Traditionally, studies of phenotypic trade-offs involving dispersal focus on resource allocation differences between flight and reproduction. However, investments in dispersal may also result in reduced allocation to other “third-party traits” (e.g. compensatory feeding) that are not directly associated with reproduction. Such traits remain largely uninvestigated for any phytophagous insect despite their importance for performance and survival. Using two wing-dimorphic, phloem-feeding planthoppers, Prokelisia dolus and Prokelisia marginata that differ dramatically in dispersal abilities, we sought evidence for a trade-off between investments in dispersal (flight apparatus) and ingestion capability (allocation to the esophageal musculature governing ingestion). Dispersal allows species to meet nutrient demands by moving to higher-quality resources. In contrast, enhanced investment in esophageal musculature increases ingestion capacity and allows phloem feeders to compensate for deteriorating plant nutrition on site. Our objectives were to compare differences in flight and feeding investment between P. dolus and P. marginata and between the wing forms of both species, and to compare ingestion capacity between the two species and wing forms. Morphometric and gravimetric measures of investment in flight versus feeding indicate that the sedentary P. dolus allocates more muscle mass to feeding whereas P. marginata invests more heavily in flight. Likewise, brachypters invest more in feeding and less in flight than macropters. The greater esophageal investment in P. dolus is associated with enhanced ingestion capacity compared to P. marginata. As a consequence, P. dolus is better equipped to meet on-site nutrient demands when faced with deteriorating plant quality than P. marginata, which must migrate elsewhere to do so. Notably, such third-party trade-offs place constraints on how insect herbivores cope with changing resources and set the stage for fundamental differences in population dynamics.     

The influence of salt and nitrogen on herbivore abundance: direct and indirect effects

We report on the influence of experimentally increased interstitial salinity and plant nitrogen on the abundance of the delphacid planthopper, Prokelisia marginata (Van Duzee) (Homoptera: Delphacidae), which feeds on salt marsh cordgrass, Spartina alterniflora. We also report the effects of these treatments on parasitism of P. marginata eggs by the fairyfly parasitoid, Anagrus sophiae (Hymenoptera: Mymaridae). Soil salinity was significantly elevated following the addition of salt pellets broadcast over the ground and plant foliar nitrogen was significantly increased after the addition of fertilizer. The addition of fertilizer increased P. marginata densities on Spartina but addition of salt did not. Neither treatment significantly affected levels of egg parasitism by A. sophiae. In this system direct effects of plants on their herbivores via changes in plant chemistry appear more important than indirect effects of plants on herbivores via their natural enemies. Received: 1 August 1997 / Accepted: 29 September 1997     

Competition as a factor underlying the abundance of an uncommon phytophagous insect, the salt-marsh planthopper Delphacodes penedetecta

Abstract. 1. Recent reviews of experimental studies provide compelling evidence that competition should be retained as a potential factor influencing the success of phytophagous insects. In this context, the objective of the study was to determine the role of interspecific and intraspecific competition, both contemporaneous and plant mediated (feeding‐induced resistance), in limiting the population density of a consistently rare insect in a guild of abundant potential competitors. 2. Competitive interactions were assessed experimentally between two phloem‐feeding planthoppers, the abundant Prokelisia dolus and the rare Delphacodes penedetecta (Hemiptera: Delphacidae). Both species are monophagous on the cordgrass Spartina alterniflora and overlap broadly in their use of habitats in the intertidal salt marshes along the Atlantic coast of North America. 3. The two planthoppers partition their cordgrass host plant, with D. penedetecta feeding more on the basal stems (particularly females) and P. dolus occurring most often on the canopy leaves. Notably, there was no evidence for niche shifting in D. penedetecta because its distribution on the plant did not change in the presence or absence of P. dolus. 4. Interspecific interactions with P. dolus had very little effect on the performance (development time and body size) and survival of D. penedetecta, a result demonstrated in both the laboratory and field. This result occurred both in contemporaneous interactions and on plants fed on previously by P. dolus. Only the males of D. penedetecta experienced weak competitive effects from P. dolus, as evidenced by reduced body size and slightly protracted development. 5. By contrast, there were strong adverse effects of intraspecific crowding (both from contemporaneous interactions and on plants fed on previously by conspecifics), whereby the survival, development time, and body size of D. penedetecta were affected very adversely. 6. These results suggest that interspecific competition is a weak force influencing the abundance of D. penedetecta in the field. Rather, strong intraspecific competition, a high requirement for plant nitrogen, and intrinsically low lifetime fecundity combine to explain the rarity of D. penedetecta.     

Is host-plant quality responsible for the populational pulses of salt-marsh planthoppers (Homoptera: Delphacidae) in northwestern Florida?

Abstract.

• 1 Rare populational pulses of the sibling species of plant-hoppers Prokelisia dolus Wilson and P.marginata (Van Duzee) occur along the Gulf Coast of Florida. Densities at least three standard deviations ahove the average number of adult planthoppers per stem host plant are exhibited during these pulses.
• 2 This context of rare local irruptions provides a basis for testing a component of T. C. R. White's hypothesis, which states that the trigger for insect outbreaks is an increase in the nutritional quality, in terms of available nitrogen, of host-plant tissue.
• 3 In the field, over approximately five generations of these hoppers (9 months), we elevated foliar nitrogen in host plants substantially by fertilizing, six times, salt-marsh plots of 10 m² with ammonium nitrate. No densities approaching a populational pulse occurred in either fertilized or unfertilized plots during the course of this study or in the year following. Fertilization did increase adult densities in the spring but this increase deteriorated over the summer months and was lost in the autumn. Host plants in fertilized plots retained high levels of nitrogen for at least the second year.
• 4 Several very dense pulses, of more than thirty adult hoppers per stem, occurred at other sites during the course of this study. Host-plant nitrogen in a pulse site did not differ from that found in host plants immediately adjacent to the pulse. We conclude that increased foliar nitrogen is not the proximate cause of the populational pulse behaviour exhibited by these planthoppers.
• 5 We speculate that these pulses result from behavioural aggregation of adult hoppers.

     

Plant–herbivore interactions and ecological stoichiometry: when do herbivores determine plant nutrient limitation?

Recent studies on plant–herbivore indirect interactions via nutrient recycling have led to the hypothesis that herbivores with a low nitrogen: phosphorus ratio, feeding on plants with a higher nitrogen: phosphorus ratio, recycle relatively more nitrogen, driving plants into phosphorus limitation. We demonstrate in this paper that such a hypothesis is valid only under restricted conditions, i.e. the nitrogen: phosphorus ratio of inorganic nutrients supplied to the system must be neither too high nor too low compared with the nitrogen: phosphorus ratio of the whole plant + herbivore biomass. If plants have a greater affinity for phosphorus than for nitrogen, low herbivore nitrogen: phosphorus ratio can even promote nitrogen limitation. These results are qualitatively robust, whether grazing functions are donor-controlled or recipient-controlled. We present a graphical analysis of these conditions based on the Zero Net Growth Isocline method.     

THE INFLUENCE OF INTRASPECIFIC VARIATION IN DISPERSAL STRATEGIES ON THE GENETIC STRUCTURE OF PLANTHOPPER POPULATIONS

The hypothesis that levels of gene flow among populations are correlated with dispersal ability has typically been tested by comparing gene flow among species that differ in dispersal abilities, an approach that potentially confounds dispersal ability with other species-specific differences. In this study, we take advantage of geographic variation in the dispersal strategies of two wing-dimorphic planthopper species, Prokelisia marginata and P. dolus, to examine for the first time whether levels of gene flow among populations are correlated with intraspecific variation in dispersal ability. We found that in both of these coastal salt marsh–inhabiting species, population-genetic subdivision, as assessed using allozyme electrophoresis, parallels geographic variation in the proportion of flight-capable adults (macropters) in a population; in regions where levels of macroptery are high, population genetic subdivision is less than in regions where levels of macroptery are low. We found no evidence that geographic variation in dispersal capability influences the degree to which gene flow declines with distance in either species. Thus, both species provided evidence that intraspecific variation in dispersal strategies influences the genetic structure of populations, and that this effect is manifested in population-genetic structure at the scale of large, coastal regions, rather than in genetic isolation by distance within a region. This conclusion was supported by interspecific comparisons revealing that: (1) population-genetic structure (G_ST) of the two Prokelisia species correlated negatively with the mean proportion of flight-capable adults within a region; and (2) there was no evidence that the degree of isolation by distance increased with decreasing dispersal capability. Populations of the relatively sedentary P. dolus clustered by geographic region (using Nei's distances), but this was not the case for the more mobile P. marginata. Furthermore, gene flow among the two major regions we surveyed (Atlantic and Gulf Coasts) has been substantial in P. marginata, but relatively less in P. dolus. The results for P. marginata suggest that differences in the dispersal strategies of Atlantic and Gulf Coast populations occur despite extensive gene flow. We argue that gene flow is biased from Atlantic to Gulf Coast populations, indicating that selection favoring a reduction in flight capability must be intense along the Gulf. Together, the results of this study provide the first rigorous evidence of a negative relationship within a species between dispersal ability and the genetic structure of populations. Furthermore, regional variation in dispersal ability is apparently maintained by selective differences that outweigh high levels of gene flow among regions.     

Does anthropogenic nitrogen deposition induce phosphorus limitation in herbivorous insects?

Anthropogenic nitrogen deposition has shifted many ecosystems from nitrogen (N) limitation to phosphorus (P) limitation. Although well documented in plants, no study to date has explored whether N deposition exacerbates P limitation at higher trophic levels, or focused on the effects of induced plant P limitation on trophic interactions. Insect herbivores exhibit strict N : P homeostasis, and should therefore be very sensitive to variations in plant N : P stoichiometry and prone to experiencing deposition‐induced P limitation. In the current study, we investigated the effects of N deposition and P availability on a plant‐herbivorous insect system. Using common milkweed (Asclepias syriaca) and two of its specialist herbivores, the monarch caterpillar (Danaus plexippus) and milkweed aphid (Aphis asclepiadis) as our study system, we found that experimental N deposition caused P limitation in milkweed plants, but not in either insect species. However, the mechanisms for the lack of P limitation were different for each insect species. The body tissues of A. asclepiadis always exhibited higher N : P ratios than that of the host plant, suggesting that the N demand of this species exceeds P demand, even under high N deposition levels. For D. plexippus, P addition increased the production of latex, which is an important defense negatively affecting D. plexippus growth rate. As a result, we illustrate that P limitation of herbivores is not an inevitable consequence of anthropogenic N deposition in terrestrial systems. Rather, species‐specific demands for nutrients and the defensive responses of plants combine to determine the responses of herbivores to P availability under N deposition.     

The use of molecular assays to identify plant pathogenic organisms vectored by biological control agents

The planthopper Prokelisia marginata VanDuzee (Homoptera: Delphacidae) has beenconsidered for the biological control of theweed Spartina alterniflora Loisel(Poaceae) in Willapa Bay, Washington, U.S.A. Prokelisia marginata is a stenophagousphloem-feeding insect with the potential totransmit bacterial plant diseases that could bemoved by less-specific vectors to other plantspecies. Initial assays with PCR primers thatare putatively specific for phytoplasmas gavepositive results in Spartina. However,subsequent analyses did not indicate thetransmission of the pathogen by theplanthopper. We sequenced the 16S ribosomalRNA (rRNA) gene of the bacterial species thatgave positive results in PCR. Comparisons withsequences available in GenBank suggested thatthe positive results using the putativelyspecific PCR primers were due to the presenceof such bacteria as Pseudomonas, Holomonas, Vibrio, and Acinetobacter. We did not find phytoplasmasin either Spartina or the planthopperP. marginata.     

Nitrogen addition does not reduce belowground competition in a salt marsh clonal plant community in Mississippi (USA)

Previous studies have suggested that belowground competition for nutrients influences plant zonation in salt marshes. In this study, I tested the hypothesis that competition for nitrogen structured a clonal plant community in a nitrogen-limited salt marsh in coastal Mississippi, USA. In contrast to most previous field studies that have investigated mechanisms of competition, I examined clonal growth responses of established genets of a nitrogen-demanding low-intertidal species (Spartina alterniflora) to nitrogen addition and the removal of a nitrogen-conserving high-intertidal species (Juncus roemerianus). Nitrogen addition stimulated clonal invasion of the Juncus zone by Spartina but did not reduce the significant competitive effects of Juncus on Spartina. Simulated Juncus shade did not reduce invasion of the Juncus zone by Spartina, indicating that belowground competition reduced clonal invasion. In the last year of the study, the border shifted unexpectedly towards the Spartina zone, resulting in competitive displacement of Spartina by Juncus. Nitrogen addition did not prevent or slow this displacement, further contradicting the nitrogen competition hypothesis. Although growth rates were much more strongly limited by nitrogen in Spartina than in Juncus, nitrogen addition did not cause the displacement of Juncus by Spartina after three growing seasons. I conclude that zonation of Spartina and Juncus is maintained by preemption of space and greater tolerance of low nitrogen supplies by Juncus in the high marsh. These results contrast sharply with findings of reduced belowground competition with nutrient addition in previous studies and highlight the important role of nutrient-mediated competition for space between clonal plants.     

Nutritional suitability and ecological relevance of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> and <Emphasis Type="Italic">Brassica oleracea</Emphasis> as foodplants for the cabbage butterfly, <Emphasis Type="Italic">Pieris rapae</Emphasis>

The thale cress, Arabidopsis thaliana, is considered to be an important model species in studying a suite of evolutionary processes. However, the species has been criticized on the basis of its comparatively small size at maturity (and consequent limitations in the amount of available biomass for herbivores) and on the duration and timing of its life cycle in nature. In the laboratory, we studied interactions between A. thaliana and the cabbage butterfly, Pieris rapae, in order to determine if plants are able to support the complete development of the herbivore. Plants were grown in pots from seedlings in densities of one, two, or four per pot. In each treatment, one, two, or five newly hatched larvae of P. rapae were placed on fully developed rosettes of A. thaliana. In a separate experiment, the same densities of P. rapae larvae were reared from hatching on single mature cabbage (Brassica oleracea) plants. Pupal fresh mass and survival of P. rapae declined with larval density when reared on A. thaliana but not on B. oleracea. However, irrespective of larval density and plant number, some P. rapae were always able to complete development on A. thaliana plants. A comparison of the dry mass of plants in different treatments with controls (= no larvae) revealed that A. thaliana partially compensated for plant damage when larval densities of P. rapae were low. By contrast, single cress plants with 5 larvae generally suffered extensive damage, whereas damage to B. oleracea plants was negligible. Rosettes of plants that were monitored in spring, when A. thaliana naturally grows, were not attacked by any insect herbivores, but there was often extensive damage from pulmonates (slugs and snails). Heavily damaged plants flowered less successfully than lightly damaged plants. Small numbers of generalist plant-parasitic nematodes were also recovered in roots and root soil. By contrast, plants monitored in a sewn summer plot were heavily attacked by insect herbivores, primarily flea beetles (Phyllotreta spp.). These results reveal that, in natural populations of A. thaliana, there is a strong phenological mismatch between the plant and most of its potential specialist insect herbivores (and their natural enemies). However, as the plant is clearly susceptible to attack from non-insect generalist invertebrate herbivores early in the season, these may be much more suitable for studies on direct defense strategies in A. thaliana.     

Genetic and soil-nutrient effects on the abundance of herbivores on willow

The effects of soil-nutrient environment, plant genotype, and the interaction between the two on the resistance of the willow, Salix sericea, to insect species in a diverse herbivore community was measured. We found that soil-nutrient environment influenced plant growth and the abundance of most herbivores of S. sericea. However, environmental effects on herbivore abundance were often modified by plant genetics; the abundance of four of seven herbivores exhibited significant genotypeby-environment interaction effects. Pure genotype effects were mostly small and non-significant. The effects of fertilization differed among herbivores. Several herbivores were more abundant on fertilized plants, one was less abundant, and the abundance of others did not change. We found that feeding guild was a poor predictor of herbivore response. Finally we found significant phenotypic and genetic correlations among growth rate, internode length, and the abundances of several herbivores.     

Effects of different leaf traits on growth rates of insect herbivores on willows

We examined relative effects of traits of leaf quality of ten willow species (Salix: Salicaceae) on growth rates of five species of insect herbivores found in interior Alaska (a willow sawfly, Nematus calais; the tiger swallowtail butterfly, Papilio canadensis; and three species of chrysomelid beetles, Gonioctena occidentalis, Calligrapha verrucosa, and Chrysomela falsa). Leaf traits examined were water content, toughness, total nitrogen contnet, pubescence, and presence or absence of phenolic glycosides. Of ten Salix species, four species contain phenolic glycosides in their leaves. We examined relative effects of water content, toughness, and nitrogen content of the Salix leaves on larval growth rates at three different levels, i.e., on a single host species, between different host species, and between herbivore species. The within-host analyses showed that effects of water content, toughness and/or nitrogen content on herbivore growth rates were generally significant in early-season herbivores but not in late-season herbivores. For each herbivore species, differences in growth rates between hosts were not explained by differences in water content, toughness, or nitrogen content. The between-herbivore analysis showed that the interspecific difference in larval growth rates were related to difference in water and nitrogen content of the hosts. Pubescence of Salix leaves had little effects on herbivore growth rates. Presence of phenolic glycosides had a positive effects on growth rates of a specialist, N. calais, but no effect on the other specialist, Ch. falsa. Presence of phenolic glycosides had, in general, negative effects on growth rates of nonspecialists, G. occidentalis, C. verrucosa, and P. canadensis.     

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.     

Direct and ecological costs of resistance and tolerance in the stinging nettle

Plant resistance and tolerance to herbivores, parasites, pathogens, and abiotic factors may involve two types of costs. First, resistance and tolerance may be costly in terms of plant fitness. Second, resistance and tolerance to multiple enemies may involve ecological trade-offs. Our study species, the stinging nettle ( Urtica dioica L.) has significant variation among seed families in resistance and tolerance as well as costs of resistance and tolerance to the holoparasitic plant Cuscuta europaea L. Here we report on variation among seed families (i.e. genetic) in tolerance to nutrient limitation and in resistance to both mammalian herbivores (i.e. number of stinging trichomes) and an invertebrate herbivore (i.e. inverse of the performance of a generalist snail, Arianta arbustorum). Our results indicate direct fitness costs of snail resistance in terms of host reproduction whereas we did not detect fitness costs of mammalian resistance or tolerance to nutrient limitation. We further tested for ecological trade-offs among tolerance or resistance to the parasitic plant, herbivore resistance, and tolerance to nutrient limitation in the stinging nettle. Tolerance of nettles to nutrient limitation and resistance to mammalian herbivores tended to correlate negatively. However, there were no significant correlations among resistance and tolerance to the different natural enemies (i.e. parasitic plants, snails, and mammals). The results of this greenhouse study thus suggest that resistance and tolerance of nettles to diverse enemies are free to evolve independently of each other but not completely without direct costs in terms of plant fitness.     

Deciduous sapling responses to season and large herbivores in a semi‐arid African savanna

Impacts of large herbivores (>5 kg) on woody plants in African savannas are potentially most severe among plants shorter than 1.6 m. It is well established that severe browsing leads to longer shoots, yet prevents saplings from recruiting into adult size‐classes in African savannas. Increased shoot length, indicating faster shoot growth, is often associated with reduced concentrations of tannins and increased nutrient concentrations, suggesting carbon limitation. We hypothesized that, on average, large herbivores suppress stem height or circumference, but increase shoot length. We also hypothesized that if there were concomitant positive effects on nutrients, or negative effects on tannin concentrations, they would be greatest early in the wet season. We sampled saplings of four deciduous woody species (Acacia grandicornuta, Dichrostachys cinerea, Combretum apiculatum and Grewia flavescens) at different stages of the wet season in a large‐scale, long‐term herbivore exclusion experiment in Kruger National Park, South Africa. Plant height, shoot length and stem circumference were generally not adversely affected by large herbivores, suggesting C limitation is rarely present among deciduous saplings in semi‐arid African savannas, allowing them to tolerate browsing. Time since first rainfall emerged as a predominant factor consistently affecting nutrient and tannin concentrations, rather than large herbivores. Nitrogen and phosphorus generally decreased (by 20–50%), while condensed tannin concentration increased (150–350%) during the wet season, except for one species. We postulate that A. grandicornuta is less prone than other species to accumulating tannins during the wet season because of high investment of C in spines. Although nutrient and tannin concentrations were generally not affected by large herbivores, species‐specific responses were evident very early in the wet season, which is when herbivore populations are most likely to be affected by differential forage quality among plants.     

The influence of species identity and herbivore feeding mode on top-down and bottom-up effects in a salt marsh system

In this study we investigated the potential importance of species identity and herbivore feeding mode in determining the strengths of top-down and bottom-up effects on phytophagous insect densities. In 1998, we conducted two factorial field experiments in which we manipulated host plant quality and intensity of parasitoid attack on three salt marsh herbivores, the planthoppers Prokelisia marginata and Pissonotus quadripustulatus (Homoptera: Delphacidae), which feed only on Spartina alterniflora and Borrichia frutescens, respectively, and the gall fly Asphondylia borrichiae (Diptera: Cecidomyiidae), which feeds only on B. frutescens. We increased plant quality through addition of nitrogen fertilizer, and decreased parasitism by trapping hymenopteran parasitoids continuously throughout the study. Herbivore densities were censused biweekly. Increasing plant quality through fertilization increased the density of all three herbivores within 2 weeks of treatment application, and higher densities were maintained for the duration of the study. Reduction of top-down pressure had no effect on either planthopper species, possibly because of compensatory mortality affecting the two species. In contrast, reduction of parasitism significantly increased the density of A. borrichiae galls, perhaps because development within gall tissue reduces the sources of compensatory mortality affecting this species. The results of this study show that the bottom-up effects of plant quality were strong and consistent for all three species, but the strength of top-down effects differed between the two feeding guilds. Thus, even for herbivores feeding on the same host plant, conclusions drawn regarding the relative importance of top-down and bottom-up effects may vary depending upon the feeding mode of the herbivore.     

Host plant nutritional quality affects the performance of the parasitoid Diadegma insulare

The well documented biochemical profile of Brassicaceae, oligophagy of the herbivore Plutella xylostella (L.) (Lepidoptera: Plutellidae), and host specialization of the parasitoid Diadegma insulare (Cresson) (Hymenoptera: Ichneumonidae) provide an ideal system for investigating tritrophic interactions mediated by nutritional quality of plants. We evaluated the bottom-up effects of five soil fertility regimes on nutritional quality of canola (Brassica napus L.) and then on several fitness correlates of female and male D. insulare as mediated through P. xylostella. Variation in soil fertility influenced the nutritional quality of host plants and this in turn affected the performance of D. insulare. In general, D. insulare performed best on plants grown with 3.0 g fertilizer pot⁻¹; these plants had 2.06-, 3.77-, and 1.02-fold more nitrogen, phosphorous and potassium, respectively than ones grown without any added fertilizer. P. xylostella escape from D. insulare was highest (32%) on plants grown at 1.0 g fertilizer, and this could be attributed to both physical and physiological defense mechanisms mediated by host plant nutritional quality. Plant stress and plant vigor are competing paradigms pertaining to the performance of herbivorous insects on their host plants. These hypotheses were originally proposed to predict responses of herbivores, but may also explain the effects of plant quality on koinobiont parasitoids, such as D. insulare.     

Induced resistance in intertidal macroalgae modifies feeding behaviour of herbivorous snails

Herbivory in terrestrial and marine systems can induce changes in plant chemistry affecting the foraging behaviour of herbivores. A model based on terrestrial plant-herbivore interactions predicts herbivory-induced changes in leaf chemistry to be manifested in (1) increased herbivore mobility, (2) increased feeding dispersal and (3) reduced tissue consumption by herbivores. This study is the first to demonstrate that herbivory-induced changes in the tissue chemistry of the brown seaweed Ascophyllum nodosum elicit the same response in the feeding behaviour of the gastropod Littorina obtusata as predicted for herbivorous insects, providing good evidence for the models validity across different ecosystems. The potential benefit of increased feeding dispersal to terrestrial plants as suggested by the model is the prevention of concentrated damage to apical tissues thereby preserving the plants ability to compete for light; A. nodosum does not conform to these predictions. Increased dispersal of feeding damage on A. nodosum away from primary frond tissues would reduce the likelihood of frond breakage implying a fitness benefit of induced resistance.     

Herbivore-herbivore interactions complicate links between soil fertility and pest resistance

Soil fertility is tightly linked with herbivore pressure because it affects the nutritional status of host plants as well as the production of anti-herbivore defenses. This in turn can influence whether herbivores in different feeding guilds render plants more or less susceptible to one another. Thus, growers’ fertility management choices may impact herbivores through a variety of indirect channels. We examined relationships between soil fertility and interactions between phloem-feeding and leaf-chewing herbivores on broccoli (Brassica oleracea) plants in the greenhouse, taking advantage of natural variation in nitrogen (N) and phosphorus (P) in soils from 20 working organic vegetable farms. Next, we experimentally fertilized soil in a field trial with N and/or P to examine the consequences of these nutrients for growth of and interactions between specialist and generalist herbivores. Soils on our cooperating farms varied widely in P and N concentrations, with 40% exceeding recommended pre-plant N concentrations and 90% exceeding P recommendations. In single-herbivore infestations, augmenting N in the soil increased caterpillar (Pieris rapae) growth, augmented N and P additively enhanced generalist green peach aphid (Myzus persicae) colonization, and augmented P (but not N) increased specialist cabbage aphid (Brevicoryne brassicae) growth. In dual-guild herbivore infestations, caterpillars facilitated specialist cabbage aphid growth in the absence of fertilizer, but this pattern disappeared under augmented N, and reversed under augmented P. We found that a complex web of indirect effects linked soil fertility to herbivore performance, depending on the identity of the nutrients being altered, the ecological roles of responding herbivore species (i.e., specialist versus generalist), and indirect interactions between chewing and sucking herbivores. More generally, we highlight that successful use of fertility management to improve pest resistance requires careful consideration of herbivore feeding niches and herbivore-herbivore interactions.