Effects of pepper (Capsicum chinense) genotypic diversity on insect herbivores

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Do larvae of species in macrolepidopteran assemblages share traits that influence susceptibility to parasitism?

Research on how morphology, behavior, and life histories of insects determine their susceptibility to parasitism has primarily focused on the traits of single host species. Very little research has been conducted attempting to determine if similarities or differences in the traits of co-occurring species, on a single plant or in a habitat, influence levels of parasitism imposed on any member of an assemblage. In this study, we use categorical and regression tree analyses to determine which traits of the larvae of macrolepidopteran species are associated with highest levels of parasitism. Of a variety of morphological, behavioral, and ecological traits of 72 species (representing eight families), caterpillar color was the trait that had the greatest influence on susceptibility to parasitism. The highest levels of parasitism were associated primarily with green larvae. These results suggest that the herbivore species composition and, specifically, the traits possessed by the species may influence community or assemblagewide patterns of parasitism. That is, sharing of traits that enhance host finding and successful parasitism may result in associational susceptibility to parasitism, whereas co-occurrence with species that differ morphologically, behaviorally, or ecologically may reduce the likelihood of parasitism and thus result in associational resistance. The concepts of associational resistance and susceptibility have historically been restricted to plant-plant interactions. The extension of these concepts to herbivores is novel. If found to be widespread, these interactions can have significant impacts on our expectations of the effectiveness of biological control agents.     

Herbivore regulation in urban agroecosystems: Direct and indirect effects

Urban agroecosystems can provide habitat for biodiversity and can benefit human communities through urban food provisioning. Moreover, urban agroecosystems could be managed so as to optimize ecosystem services like natural pest control provided by trophic interactions between natural enemies and herbivores. As in other ecosystems, predation and parasitism regulate herbivores in urban settings, but less is known about the relative importance of direct and indirect effects at local and landscape scales in highly managed urban agroecosystems. We collected data on herbivore (cabbage aphid) density and parasitism ratios (proportion of parasitized aphid “mummies”) in 25 community gardens in three counties in the California central coast, USA. We used structural equation modeling to examine the effects of direct factors (host plant characteristics and parasitism) and indirect factors (soil, garden, and landscape characteristics) on herbivore density changes at two time points in the growing season (June and August). Aphid density, but not parasitism, varied across counties over the season, and there was a strong negative relationship between aphid density and parasitism. Direct effects were strong drivers of aphid density but not parasitism. In June, aphid density increased with host plant volume but decreased with greater floral density, while parasitism was only influenced by aphid density. In August, host plant volume similarly positively affected aphid density, and soil water holding capacity increased host plant volume. In addition, host plant density had a strong negative effect on parasitism. Urban gardeners may be able to reduce aphid pest densities by increasing floral resource density and strategically spatially distributing host plants throughout garden beds, though these processes depend on the season. The indirect effects of soil water holding capacity on aphid densities further suggest a critical role of human management on pest populations and pest control services through soil amendments and irrigation.     

Costs of aggregation: shadow competition in a sit-and-wait predator

The role of top-down (e.g. parasitism) and bottom-up (e.g. resource competition) processes is of fundamental importance for the stability and persistence of insect herbivore populations. Although emphasis has often focused on single regulatory agents, the processes underpinning tri-trophic interactions may actually be more pluralistic. Recently, further complexities involved in the regulation of tri-trophic systems have been highlighted. In particular, life history characteristics may have a concomitant role when coupled with the regulatory effects of resource competition and/or parasitism. Here we present an age-structured model to investigate the effects of larval development period, parasitism and resource competition on the stability and persistence of herbivore–parasitoid interactions. We show that the influence of weak density dependent parasitism is sufficient to stabilise the interaction when the period of host susceptibility to parasitism is short. For longer periods of host susceptibility, parasitism needs to be highly non-linear to overcome the destabilising effects of the time delays. In systems where host development is protracted through the season we predict that resource competition is likely to be the dominant process for herbivore regulation. We use this age-structured approach to explore the population dynamics of two field studies from temperate ecosystems. Predictions from these case studies show 1) that both the strength and type of competition and parasitism are important for the stability and persistence of the particular system, and 2) that the length of the developmental period of the vulnerable host is critical to understand the influence of different regulatory processes. Host demography is of overriding importance in determining whether herbivores show outbreaks, and which particular ecological processes and mechanisms are responsible for generating such overcompensatory dynamics.     

农业景观格局与麦蚜密度对其初寄生蜂与重寄生蜂种群及寄生率的影响

农业景观格局与过程能够强烈影响寄生蜂对寄主的寻找及寄生作用,寄主密度亦是影响寄生蜂分布的重要因素,然而农业景观的格局和寄主密度对寄生蜂寄生率的相互影响是一项值得研究的工作.在简单与复杂2种麦田农业景观结构下,调查了麦蚜的分布格局与2种寄主密度下麦蚜的初寄生率与重寄生率,分析了景观结构对麦蚜密度的影响、景观格局与麦蚜密度对寄生蜂寄生率与重寄生率的影响及交互作用.结果表明:景观结构的复杂性对麦蚜分布和寄生蜂初寄生率与重寄生率的影响均不明显,但寄主密度与景观结构的复杂性对寄生蜂的影响存在着明显的交互作用,寄主密度与寄生率呈正相关,寄主密度较低时烟蚜茧蜂为优势种,寄主密度较高时燕麦蚜茧蜂为优势种.麦蚜初寄生蜂与重寄生蜂对寄主密度的反应与其形态学、体型大小以及生活史特征相关,初寄生蜂与重寄生蜂的群落组成显著影响其对麦蚜的寄生率,而与景观结构的复杂性关系不大.     

Maize Domestication and Anti-Herbivore Defences: Leaf-Specific Dynamics during Early Ontogeny of Maize and Its Wild Ancestors

As a consequence of artificial selection for specific traits, crop plants underwent considerable genotypic and phenotypic changes during the process of domestication. These changes may have led to reduced resistance in the cultivated plant due to shifts in resource allocation from defensive traits to increased growth rates and yield. Modern maize (Zea mays ssp. mays) was domesticated from its ancestor Balsas teosinte (Z. mays ssp. parviglumis) approximately 9000 years ago. Although maize displays a high genetic overlap with its direct ancestor and other annual teosintes, several studies show that maize and its ancestors differ in their resistance phenotypes with teosintes being less susceptible to herbivore damage. However, the underlying mechanisms are poorly understood. Here we addressed the question to what extent maize domestication has affected two crucial chemical and one physical defence traits and whether differences in their expression may explain the differences in herbivore resistance levels. The ontogenetic trajectories of 1,4-benzoxazin-3-ones, maysin and leaf toughness were monitored for different leaf types across several maize cultivars and teosinte accessions during early vegetative growth stages. We found significant quantitative and qualitative differences in 1,4-benzoxazin-3-one accumulation in an initial pairwise comparison, but we did not find consistent differences between wild and cultivated genotypes during a more thorough examination employing several cultivars/accessions. Yet, 1,4-benzoxazin-3-one levels tended to decline more rapidly with plant age in the modern maize cultivars. Foliar maysin levels and leaf toughness increased with plant age in a leaf-specific manner, but were also unaffected by domestication. Based on our findings we suggest that defence traits other than the ones that were investigated are responsible for the observed differences in herbivore resistance between teosinte and maize. Furthermore, our results indicate that single pairwise comparisons may lead to false conclusions regarding the effects of domestication on defensive and possibly other traits.     

Host‐plant genotypic diversity and community genetic interactions mediate aphid spatial distribution

Genetic variation in plants can influence the community structure of associated species, through both direct and indirect interactions. Herbivorous insects are known to feed on a restricted range of plants, and herbivore preference and performance can vary among host plants within a species due to genetically based traits of the plant (e.g., defensive compounds). In a natural system, we expect to find genetic variation within both plant and herbivore communities and we expect this variation to influence species interactions. Using a three‐species plant‐aphid model system, we investigated the effect of genetic diversity on genetic interactions among the community members. Our system involved a host plant (Hordeum vulgare) that was shared by an aphid (Sitobion avenae) and a hemi‐parasitic plant (Rhinanthus minor). We showed that aphids cluster more tightly in a genetically diverse host‐plant community than in a genetic monoculture, with host‐plant genetic diversity explaining up to 24% of the variation in aphid distribution. This is driven by differing preferences of the aphids to the different plant genotypes and their resulting performance on these plants. Within the two host‐plant diversity levels, aphid spatial distribution was influenced by an interaction among the aphid's own genotype, the genotype of a competing aphid, the origin of the parasitic plant population, and the host‐plant genotype. Thus, the overall outcome involves both direct (i.e., host plant to aphid) and indirect (i.e., parasitic plant to aphid) interactions across all these species. These results show that a complex genetic environment influences the distribution of herbivores among host plants. Thus, in genetically diverse systems, interspecific genetic interactions between the host plant and herbivore can influence the population dynamics of the system and could also structure local communities. We suggest that direct and indirect genotypic interactions among species can influence community structure and processes.     

Exploiting physical defence traits for crop protection: leaf trichomes of Rubus idaeus have deterrent effects on spider mites but not aphids

Plants possess anti‐herbivore defences that could be exploited for crop protection. The potential for deploying physical defence traits for more sustainable pest management (i.e. reduced pesticide application) has not been fully realised. Using a perennial crop (red raspberry, Rubus idaeus), we take the novel approach of quantifying within‐ and between‐genotype variation in a resistance trait, leaf trichome density, to determine precisely the effect of trichomes on host plant preference and suitability for two shoot‐feeding arthropods, the European large raspberry aphid (Amphorophora idaei) and two‐spotted spider mite (Tetranychus urticae). Additionally, we tested whether this trait influenced searching behaviour of a generalist herbivore predator (lacewing larvae, Chrysoperla carnea). Although there was no consistent genotypic variation in R. idaeus suitability for T. urticae, our hypothesis that T. urticae would avoid high leaf trichome density was supported on certain genotypes. The deterrent effect was mainly on egg deposition rather than leaf selection by adults, with up to sixfold differences in leaf preference depending on the genotypes offered. By contrast, there was significant genotypic variation in R. idaeus suitability for A. idaei (10‐fold variation in aphid abundance), but, contrary to our prediction, aphid preference and infestation levels were unrelated to leaf trichome density. Instead, A. idaei performed best on vigorous genotypes, indicating that plant tolerance traits contributed to R. idaeus suitability for aphids. Leaf trichomes had little effect on the behaviour of the beneficial control agent C. carnea larvae. We conclude that physical anti‐herbivore defences, specifically leaf trichomes, could be deployed to deter particular arthropod pests. However, the mechanistic approach adopted here is necessary to avoid antagonistic effects on other pests or on natural enemies.     

Plant genotypic diversity reduces the rate of consumer resource utilization

While plant species diversity can reduce herbivore densities and herbivory, little is known regarding how plant genotypic diversity alters resource utilization by herbivores. Here, we show that an invasive folivore—the Japanese beetle (Popillia japonica)—increases 28 per cent in abundance, but consumes 24 per cent less foliage in genotypic polycultures compared with monocultures of the common evening primrose (Oenothera biennis). We found strong complementarity for reduced herbivore damage among plant genotypes growing in polycultures and a weak dominance effect of particularly resistant genotypes. Sequential feeding by P. japonica on different genotypes from polycultures resulted in reduced consumption compared with feeding on different plants of the same genotype from monocultures. Thus, diet mixing among plant genotypes reduced herbivore consumption efficiency. Despite positive complementarity driving an increase in fruit production in polycultures, we observed a trade-off between complementarity for increased plant productivity and resistance to herbivory, suggesting costs in the complementary use of resources by plant genotypes may manifest across trophic levels. These results elucidate mechanisms for how plant genotypic diversity simultaneously alters resource utilization by both producers and consumers, and show that population genotypic diversity can increase the resistance of a native plant to an invasive herbivore.     

Consequences for a host–parasitoid interaction of host-plant aggregation, isolation, and phenology

Abstract.  1. Spatial habitat structure can influence the likelihood of patch colonisation by dispersing individuals, and this likelihood may differ according to trophic position, potentially leading to a refuge from parasitism for hosts.
2. Whether habitat patch size, isolation, and host-plant heterogeneity differentially affected host and parasitoid abundance, and parasitism rates was tested using a tri-trophic thistle–herbivore–parasitoid system.
3.  Cirsium palustre thistles ( n = 240) were transplanted in 24 blocks replicated in two sites, creating a range of habitat patch sizes at increasing distance from a pre-existing source population. Plant architecture and phenological stage were measured for each plant and the numbers of the herbivore Tephritis conura and parasitoid Pteromalus elevatus recorded.
4. Mean herbivore numbers per plant increased with host-plant density per patch, but parasitoid numbers and parasitism rates were unaffected. Patch distance from the source population did not influence insect abundance or parasitism rates. Parasitoid abundance was positively correlated with host insect number, and parasitism rates were negatively density dependent. Host-plant phenological stage was positively correlated with herbivore and parasitoid abundance, and parasitism rates at both patch and host-plant scales.
5. The differential response between herbivore and parasitoid to host-plant density did not lead to a spatial refuge but may have contributed to the observed parasitism rates being negatively density dependent. Heterogeneity in patch quality, mediated by variation in host-plant phenology, was more important than spatial habitat structure for both the herbivore and parasitoid populations, and for parasitism rates.     

Scale-dependent responses in cabbage herbivores affect attack rates in spatially heterogeneous systems

Herbivorous insects face a dilemma when selecting suitable hosts in a complex environment, and their sensory capability may often reduce the female capacity for proper selection. As a consequence, eggs are often deposited on inferior hosts, affecting both insect and host plant fitness. We examined the attack rates of three cabbage herbivores in monocultures and biculture plots of different Brassica oleracea genotypes, with different spatial heterogeneity. The main goals of the study were to improve our understanding of the spatial scales involved in herbivore search processes and to examine the possibility of using spatial heterogeneity for manipulating pest attack rates in cabbage cropping systems. The results showed that the host selection behaviour of the small white butterfly (Pieris rapae) was strongly dependent on spatial heterogeneity. The difference in egg density between plant genotypes was larger when contrasting plants were growing in close proximity than in monoculture. This suggests that P. rapae is able to differentiate among genotypes from a small distance, while selection is compromised at larger spatial scales. The two other herbivores in the study (Mamestra brassicae and Delia radicum) did not respond to heterogeneity at any spatial scale, but showed a constant preference hierarchy. This suggests that host selection in these species occurs after direct plant contact. The difference in species’ responses to spatial heterogeneity has consequences both for selection gradients in natural communities and for the potential to reduce pest attack in polyculture systems.     

Interaction of Anthonomus grandis and cotton genotypes: biological and behavioral responses

The boll weevil, Anthonomus grandisBoheman (Coleoptera: Curculionidae), is a key pest of cotton, Gossypium hirsutumL. (Malvaceae). Knowledge about boll weevil feeding and oviposition behavior and its response to plant volatiles can underpin our understanding of host plant resistance, and contribute to improved monitoring and mass capture of this pest. Boll weevil oviposition preference and immature development in four cotton genotypes (CNPA TB90, TB85, TB15, and BRS Rubi) were investigated in the laboratory and greenhouse. Volatile organic compounds (VOCs) produced by TB90 and Rubi genotypes were obtained from herbivore‐damaged and undamaged control plants at two phenological stages – vegetative (prior to squaring) and reproductive (during squaring) – and four collection times – 24, 48, 72, and 96 h following herbivore damage. The boll weevil exhibited similar feeding and oviposition behavior across the four tested cotton genotypes. The chemical profiles of herbivore‐damaged plants of both genotypes across the two phenological stages were qualitatively similar, but differed in the amount of volatiles produced. Boll weevil response to VOC extracts was studied using a Y‐tube olfactometer. The boll weevil exhibited similar feeding and oviposition behavior at the four tested cotton genotypes, although delayed development and production of smaller adults was found when fed TB85. The chemical profile of herbivore‐damaged plants of both genotypes at the two phenological stages and time periods (24–96 h) was similar qualitatively, with 30 identified compounds, but differed in the amount of volatiles produced. Additionally, boll weevil olfactory response was positive to herbivory‐induced volatiles. The results help to understand the interaction between A. grandis and cotton plants, and why it is difficult to obtain cotton genotypes possessing constitutive resistance to this pest.     

Increased nutrient availability decreases insect resistance in cranberry

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Top‐down,bottom‐up,or side to side? Within‐trophic‐level interactions modify trophic dynamics of a salt marsh herbivore

Many factors can influence the top‐down and bottom‐up dynamics of phytophagous insects. Although interactions between herbivore species have been frequently shown to be ecologically important, the effects of such horizontal trophic interactions on the relative roles of top‐down and bottom‐up forces have gone largely unstudied. In this paper we report on the results of a factorial field experiment in which we examined the effects of within‐trophic‐level interactions on the top‐down and bottom‐up dynamics of a salt marsh planthopper.
We manipulated the bottom‐up effects of plant quality by increasing soil salinity, and manipulated top‐down effects by decreasing the intensity of parasitoid attack with yellow sticky traps that removed hymenopteran parasitoids. We applied these treatments to plots in two patches of the host plant, one with low densities of lepidopteran stem borer larvae, and one with high densities of stem borers. We maintained the treatments and monitored planthopper density for ten months, from March through December 1999. Increased salinity significantly increased planthopper density within one month of the first application of salt. The rapid response of the planthopper to salt treatments suggested a chemical mechanism, perhaps mobilization of bound nitrogen. Yellow sticky traps, although significantly reducing parasitism of planthopper eggs, had little impact on hopper density. The density of lepidopteran stem borers, however, had an even greater impact on planthopper density than did salt treatments, with high stem borer plots supporting much lower densities of hoppers. Stem borer density also reduced the response of the planthopper to other treatments, especially salt supplementation. The results of this study show that the impact of within‐trophic‐level interactions can significantly change herbivore trophic dynamics and can be even more important than either top‐down or bottom‐up effects in determining herbivore density.     

EVOLUTION OF AN APHID-PARASITOID INTERACTION: VARIATION IN RESISTANCE TO PARASITISM AMONG APHID POPULATIONS SPECIALIZED ON DIFFERENT PLANTS

The evolution of associations between herbivorous insects and their parasitoids is likely to be influenced by the relationship between the herbivore and its host plants. If populations of specialized herbivorous insects are structured by their host plants such that populations on different hosts are genetically differentiated, then the traits affecting insect-parasitoid interactions may exhibit an associated structure. The pea aphid (Acyrthosiphon pisum) is a herbivorous insect species comprised of genetically distinct groups that are specialized on different host plants (Via 1991a, 1994). Here, we examine how the genetic differentiation of pea aphid populations on different host plants affects their interaction with a parasitoid wasp, Aphidius ervi. We performed four experiments. (1) By exposing pea aphids from both alfalfa and clover to parasitoids from both crops, we demonstrate that pea aphid populations that are specialized on alfalfa are successfully parasitized less often than are populations specialized on clover. This difference in parasitism rate does not depend upon whether the wasps were collected from alfalfa or clover fields. (2) When we controlled for potential differences in aphid and parasitoid behavior between the two host plants and ensured that aphids were attacked, we found that pea aphids from alfalfa were still parasitized less often than pea aphids from clover. Thus, the difference in parasitism rates is not due to behavior of either aphids or wasps, but appears to be a physiologically based difference in resistance to parasitism. (3) Replicates of pea aphid clones reared on their own host plant and on a common host plant, fava bean, exhibited the same pattern of resistance as above. Thus, there do not appear to be nutritional or secondary chemical effects on the level of physiological resistance in the aphids due to feeding on clover or alfalfa, and therefore the difference in resistance on the two crops appears to be genetically based. (4) We assayed for genetic variation in resistance among individual pea aphid clones collected from clover fields and found no detectable genetic variation for resistance to parasitism within two populations sampled from clover. This is in contrast to Henter and Via's (1995) report of abundant genetic variation in resistance to this parasitoid within a pea aphid population on alfalfa. Low levels of genetic variation may be one factor that constrains the evolution of resistance to parasitism in the populations of pea aphids from clover, leading them to remain more susceptible than populations of the same species from alfalfa.     

Host-plant effects the expression of resistance to Bacillus thuringiensis kurstaki in Trichoplusia ni (Hubner): an important factor in resistance evolution

Pathogens are thought to exert strong selection on their hosts leading to increased host resistance. Bacillus thuringiensis kurstaki (Bkt) is a ubiquitous entomopathogen that has become the mainstay of nonchemical control of Lepidopteran pests and thus, the potential exists for the evolution of resistance in targeted host insects. We have studied the expression of Btk resistance in the cabbage looper, Trichoplusia ni (Hubner). For this generalist insect herbivore, three common host plants, tomato, pepper and cucumber, vary in suitability for larval growth and development. Here we show that the host plant also affects the overall toxicity of Btk, the relative expression of resistance between a resistant and a susceptible line and their F(1) reciprocal crosses, and importantly, the dominance of the resistance trait. This study demonstrates that tri-trophic interactions involving an insect, host plants and a pathogen have the potential to strongly influence the evolutionary response of an insect host to a pathogen.     

Effects of tree species diversity and genotypic diversity on leafminers and parasitoids in a tropical forest plantation

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Geographical variation and sexual dimorphism of phenoloxidase levels in Japanese beetles (Popillia japonica)

Phenoloxidase (PO) production can be used as an indicator of pathogen defence in insects. The geographical mosaic of coevolution implies that traits associated with pathogen defence will vary across a geographical range. Bateman's principle implies that the benefit of increased defence levels is greater for females than males. To test both of these hypotheses, we sampled four populations of Japanese beetles, Popillia japonica, across Vermont: two were from locations using biological pest control, and two locations did not use biocontrol. We quantified defence levels (PO) and parasite resistance in males and females from each population. Populations from areas with biocontrol exhibited greater PO production, females produced more PO, populations with higher levels of PO suffered less mortality from pathogen exposure, and PO levels positively correlate with increased mortality in the absence of parasitism. Results support the geographical mosaic of coevolution and the implications of Bateman's principle.     

Effect of vegetation density, height, and connectivity on the oviposition pattern of the leaf beetle Galeruca tanaceti

Vegetation structure can profoundly influence patterns of abundance, distribution, and reproduction of herbivorous insects and their susceptibility to natural enemies. The three main structural traits of herbaceous vegetation are density, height, and connectivity. This study determined the herbivore response to each of these three parameters by analysing oviposition patterns in the field and studying the underlying mechanisms in laboratory bioassays. The generalist leaf beetle, Galeruca tanaceti L. (Coleoptera: Chrysomelidae), preferentially deposits its egg clutches on non‐host plants such as grasses. Earlier studies revealed that oviposition within structurally complex vegetation reduces the risk of egg parasitism. Consequently, leaf beetle females should prefer patches with dense, tall, or connected vegetation for oviposition in order to increase their reproductive success. In the present study, we tested the following three hypotheses on the effect of stem density, height, and connectivity on oviposition: (1) Within habitats, the number of egg clutches in areas with high stem densities is disproportionately higher than in low‐density areas. The number of egg clutches on (2) tall stems or (3) in vegetation with high connectivity is higher than expected for a random distribution. In the field, stem density and height were positively correlated with egg clutch presence. Moreover, a disproportionately high presence of egg clutches was determined in patches with high stem densities. Stem height had a positive influence on oviposition, also in a laboratory two‐choice bioassay, whereas stem density and connectivity did not affect oviposition preferences in the laboratory. Therefore, stem height and, potentially, density, but not connectivity, seem to trigger oviposition site selection of the herbivore. This study made evident that certain, but not all traits of the vegetation structure can impose a strong influence on oviposition patterns of herbivorous insects. The results were finally compared with data on the movement patterns of the specialised egg parasitoid of the herbivore in comparable types of vegetation structure.     

Plant genotypic diversity increases population size of a herbivorous insect

It is critical to incorporate the process of population dynamics into community genetics studies to identify the mechanisms of the linkage between host plant genetics and associated communities. We studied the effects of plant genotypic diversity of tall goldenrod Solidago altissima on the population dynamics of the aphid Uroleucon nigrotuberculatum. We found genotypic variation in plant resistance to the aphid in our experiments. To determine the impact of plant genotypic diversity on aphid population dynamics, we compared aphid densities under conditions of three treatments: single-genotype plots, mixed-genotype plots and mixed-genotype-with-cages plots. In the latter treatment plants were individually caged to prevent natural enemy attack and aphid movement among plants. The synergistic effects of genotypes on population size were demonstrated by the greater aphid population size in the mixed-genotype treatment than expected from additive effects alone. Two non-exclusive hypotheses are proposed to explain this pattern. First, there is a source-sink relationship among plant genotypes: aphids move from plant genotypes where their reproduction is high to genotypes where their reproduction is low. Second, natural enemy mortality is reduced in mixed plots in a matrix of diverse plant genotypes.