Impact of Nosema pyrausta on field populations of Macrocentrus grandii, an introduced parasite of the European corn borer, Ostrinia nubilalis

In field populations of the European corn borer, Ostrinia nubilalis, there is a significant inverse correlation between the prevalence of corn borer infection with the microsporidian Nosema pyrausta and the prevalence of parasitism by the introduced braconid Macrocentrus grandii where infections with N. pyrausta exceed 45%. This relationship occurs geographically and from year to year. Corn borer infection with N. pyrausta, the source of infection for M. grandii, is significantly related to corn borer density in the cornfield. These findings strongly suggest that N. pyrausta has a significant adverse affect on field populations of M. grandii and may help explain the diminishing role of this and other introduced parasites as natural controls of the corn borer in the United States.     

Strong but opposing effects of associational resistance and susceptibility on defense phenotype in an African savanna plant

The susceptibility of plants to herbivores can be strongly influenced by the identity, morphology and palatability of neighboring plants. While the defensive traits of neighbors often determine the mechanism and strength of associational resistance and susceptibility, the effect of neighbors on plant defense phenotype remains poorly understood. We used field surveys and a prickle‐removal experiment in a semi‐arid Kenyan savanna to evaluate the efficacy of physical defenses against large mammalian herbivores in a common understory plant, Solanum campylacanthum. We then quantified the respective effects of spinescent Acacia trees and short‐statured grasses on browsing damage and prickle density in S. campylacanthum. We paired measurements of prickle density beneath and outside tree canopies with long‐term herbivore‐exclusion experiments to evaluate whether associational resistance reduced defense investment by decreasing browsing damage. Likewise, we compared defense phenotype within and outside pre‐existing and experimentally created clearings to determine whether grass neighbors increased defense investment via associational susceptibility. Removing prickles increased the frequency of browsing by ~25%, and surveys of herbivory damage on defended leaves suggested that herbivores tended to avoid prickles. As predicted, associational resistance and susceptibility had opposing effects on plant phenotype: individuals growing beneath Acacia canopies (or, analogously, within large‐herbivore exclosures) had a significantly lower proportion of their leaves browsed and produced ~ 70–80% fewer prickles than those outside refuges, whereas plants in grass‐dominated clearings were more heavily browsed and produced nearly twice as many prickles as plants outside clearings. Our results demonstrate that associational resistance and susceptibility have strong, but opposing, effects on plant defense phenotype, and that variable herbivore damage is a major source of intraspecific variation in defense phenotype in this system.     

Tests of associational defence provided by hairy plants for glabrous plants of Arabidopsis halleri subsp. gemmifera against insect herbivores

1. Trichome‐producing (hairy) and trichomeless (glabrous) plants of Arabidopsis halleri subsp. gemmifera were investigated to test whether plant resistance to herbivory depends on the plants' phenotypes and/or the phenotypes of neighbouring plants (associational effects). 2. A common garden experiment was conducted in which the relative frequency of hairy and glabrous plants was manipulated. Two species of leaf‐chewing insects (larvae of a white butterfly and a cabbage sawfly) were found less often on hairy plants than on glabrous plants. By contrast, the numbers of aphids and flea beetles did not differ significantly between hairy and glabrous plants. For none of these insects did abundance depend on the frequency of the two plant morphs. 3. A field survey was conducted in two natural populations of A. halleri. In the first population, a species of white butterfly was the dominant herbivore, and hairy plants incurred less leaf damage than glabrous plants across 2 years. By contrast, in the other population, where flea beetles were dominant, there were no consistent differences in leaf damage between the two types of plants. In neither of the two populations was any evidence found of associational effects. 4. This study did not provide any conclusive evidence of associational effects of anti‐herbivore resistance, but it was discovered that trichomes can confer resistance to certain herbivores. Given the results of previous work by the authors on associational effects against a flightless leaf beetle, such associational effects of the trichome dimorphism of A. halleri were herbivore‐specific.     

Do distances among host patches and host density affect the distribution of a specialist parasitoid?

The effect of spatial habitat structure and patchiness may differ among species within a multi-trophic system. Theoretical models predict that species at higher trophic levels are more negatively affected by fragmentation than are their hosts or preys. The absence or presence of the higher trophic level, in turn, can affect the population dynamics of lower levels and even the stability of the trophic system as a whole. The present study examines different effects of spatial habitat structure with two field experiments, using as model system the parasitoid Cotesia popularis which is a specialist larval parasitoid of the herbivore Tyria jacobaeae. One experiment examines the colonisation rate of the parasitoid and the percentage parasitism at distances occurring on a natural scale; the other experiment examines the dispersal rate and the percentage parasitism in relation to the density of the herbivore and its host plant. C. popularis was able to reach artificial host populations at distances up to the largest distance created (at least 80 m from the nearest source population). Also, the percentage parasitism did not differ among the distances. The density experiment showed that the total number of herbivores parasitised was higher in patches with a high density of hosts, regardless of the density of the host plant. The percentage parasitism, however, was not related to the density of the host. The density of the host plant did have a (marginally) significant effect on the percentage parasitism, probably indicating that the parasitoid uses the host plant of the herbivore as a cue to find the herbivore itself. In conclusion, the parasitoid was not affected by the spatial habitat structure on spatial scales that are typical of local patches.     

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.     

Variation in flight response of the specialist parasitoidMacrocentrus grandii goidanich to odours from food plants of its European corn borer host

The specialist parasitoidMacrocentrus grandii Goidanich (Hymenoptera: Braconidae) appears to parasitize its polyphagous host, European corn borer (Ostrinia nubilalis (Hübner)) (Lepidoptera: Pyralidae), in only certain habitats. To determine whether it differed in its olfactory response to host-habitat odours, volatiles from four plants were isolated using Tenax. Wind tunnel bioassays of the extracts revealed that, besides corn which was tested in an earlier study, olfactory stimuli for attraction ofM. grandii females were present in potato and snap bean but not in pepper or soybean. To further characterize the response to pepper and soybean, these extracts were bioassayed in combination with an attractive extract. The results indicated that pepper volatiles evoked a neutral response inM. grandii while response to soybean volatiles appeared to be neutral or slightly negative. The innate response to soybean volatiles was altered to one of attraction after oviposition experience on soybean. Seven days after oviposition, experienced females continued to respond positively to soybean volatiles. Components of soybean volatiles responsible for the change in flight behaviour resulting from oviposition experience were eluted by nonpolar and slightly polar solvents. These results support the idea that plant odour may be a factor determining the range of plants on whichM. grandii parasitizes its host. The study indicates the occurrence of associative learning of plant-related volatiles during oviposition inM. grandii, and suggests the involvement of diverse plant compounds in the learning process.     

Nosema pyrausta infection in Macrocentrus grandii, a braconid parasite of the European corn borer, Ostrinia nubilalis

Macrocentrus grandii which develop within Nosema pyrausta-infected larvae of the European corn borer, Ostrinia nubilalis, develop direct systemic infections from the ingestion of spores at the time of larval emergence from the host. Infections adversely affect pupal development and adult longevity. Infected females are unable to transmit the microsporidian to additional corn borer hosts. Pathogen development in the parasite host appears identical to its development in the corn borer host and mature spores show no morphological differences in size or shape when observed at the ultrastructural level. The prevalence of infection in natural parasite populations is 53.8% and closely parallels the 56.7% prevalence of infection in corn borer populations. Results suggest N. pyrausta may play a significant role in limiting M. grandii populations when levels of N. pyrausta in corn borers are high.     

Evaluation of conventional resistance to European corn borer (Lepidoptera: Crambidae) and western corn rootworm (Coleoptera: Chrysomelidae) in experimental maize lines developed from a backcross breeding program

Plant resistance is a promising control method for the two most damaging insect pests of maize, Zea mays L.: the European corn borer, Ostrinia nubilalis (Hübner), and the western corn rootworm Diabrotica virgifera virgifera LeConte. Fifteen experimental lines of maize, derived from a backcross breeding program designed to introgress resistance to European corn borer from Peruvian maize into two U.S. Corn Belt adapted inbred lines, were evaluated for resistance to European corn borer and western corn rootwonrm. The experimental lines were in the second generation of backcrossing. All experimental lines were resistant to leaf blade feeding by European corn borer. These lines had low levels of 2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one, a chemical commonly associated with leaf blade feeding resistance, indicating that this was not the mechanism of resistance to leaf blade feeding in these lines. Eleven experimental lines were resistant to leaf sheath and collar feeding by European corn borer. Useful sources of European corn borer ovipositional nonpreference and root feeding resistance to western corn rootworm were not identified. Some of the lines evaluated in this study may provide useful sources of resistance to both leaf blade and leaf sheath and collar feeding by European corn borer.     

Benefits of self-superparasitism in a polyembryonic parasitoid

Macrocentrus grandii is a polyembryonic parasitoid, with embryos that divide asexually within the host (European corn borer, Ostrinia nubilalis) to produce broods of clonal offspring. From a biological control standpoint, polyembryony seems advantageous because each parasitized host yields multiple parasitoids with minimal time and egg investment. When we observed M. grandii in the field, however, we found that the parasitoid virtually always invested additional time and, if possible, stings into hosts that it had already stung, apparently reducing some of the advantages of polyembryony. We therefore investigated and found support for two potential benefits that can be gained by self-superparasitism in this system. First, multiple stings allowed production of mixed-sex broods: 27% of multiply-stung versus 0% of singly-stung hosts produced mixed-sex broods. Second, multiple stings increased mean parasitoid progeny produced per host, primarily by reducing the chance of complete brood failure. Our results indicate substantial benefit for a second sting, but little benefit for three or more stings, even though M. grandii was sometimes observed to invest more than two stings within a single host. However, we also found that within-host larval competition is prevalent, suggesting that supernumerary stings may pay off in competition against conspecific larvae. Such additional investment within a single host would be particularly beneficial when hosts, rather than eggs, are limiting, but would decrease the overall efficacy of M. grandii as a biological control agent.     

Transgenic corn for control of the European corn borer and corn rootworms: a survey of Midwestern farmers' practices and perceptions

In 2001, a self-administered questionnaire was sent to 1000 corn, Zea mays L., farmers in each of five states (Illinois, Indiana, Iowa, Minnesota, and Nebraska) to evaluate their perceptions of transgenic corn designed to control the European corn borer, Ostrinia nubilalis (Hübner), and corn rootworms, Diabrotica spp. Respondents returned 1,313 surveys (26.2%). Farmers with small acreages planted a greater portion of their corn (54.5%) with transgenic corn for control of European corn borer than farmers with large farms (39.2%). The majority (75.2%) of farmers use crop rotation to control the corn rootworm. Nine insecticides comprised 92.2% of the commercial soil insecticides used for control of corn rootworm larvae. More than one-third of the farmers in Illinois (33.5%) and Indiana (39.4%) treated first-year corn for corn rootworm, primarily due to western corn rootworm egg laying in soybean, Glycine max (L.). When asked whether they would plant transgenic corn protected against the corn rootworm, 35.0% of farmers responded they would, whereas 40.5% said they were unsure. The two greatest farmer concerns about transgenic corn were the ability to sell harvested grain (59.3%) and additional technology fees (54.8%). Respondents indicated that less farmer exposure to insecticide (69.9%) and less insecticide in the environment (68.5%) were the primary benefits of transgenic corn. Farmers who had no concerns about transgenic corn for rootworm control were more likely to purchase the product (46.8%). The most common refuge-planting options farmers favored were adjacent fields (30.9%) and split fields (29.9%). Farmers (21.1%) observed a yield increase (23.7 bu/ha [9.6 bu/acre]) when using transgenic corn for European corn borer control compared with non-transgenic corn. These data can help in understanding farmers' knowledge and concerns regarding transgenic corn. This information may be of value to guide researchers, extension specialists, and policy makers in designing insect resistance management and integrated pest management programs.     

Plant genetic differences influence herbivore community structure: evidence from a hybrid willow system

To determine the influence of plant genetic variation on community structure of insect herbivores, we examined the abundances of 14 herbivore species among six genetic classes of willow: Salix eriocephala, S. sericea, their F₁ and F₂ interspecific hybrids, and backcross hybrids to each parental species. We placed 1-year-old plants, grown from seeds generated from controlled crosses, in a common garden. During the growing season, we censused gall-inducing flies and sawflies, leaf-mining insects, and leaf-folding Lepidoptera to determine the community structure of herbivorous insects on the six genetic classes. Our results provided convincing evidence that the community structure of insect herbivores in this hybrid willow system was shaped by genetic differences among the parental species and the hybrid genetic classes. Using MANOVA, we detected significant differences among genetic classes for both absolute and relative abundance of herbivores. Using canonical discriminant analysis, we found that centroid locations describing community structure of the insect herbivores differed for each genetic class. Moreover, the centroids for the four hybrid classes were located well outside of the range between the centroids for the parental species, suggesting that more than additive genetic effects of the two parental species influenced community formation on hybrid classes. Line-cross analysis suggested that plant genetic factors responsible for structuring the herbivore community involved epistatic effects, as well as additive and dominance effects. We discuss the ramifications of these results in regard to the structure of insect herbivore communities on plants and the implications of our findings for the evolution of interspecific interactions.     

Belowground resistance to western corn rootworm in lepidopteran-resistant maize genotypes

Several maize, Zea mays L., inbred lines developed from an Antiguan maize population have been shown to exhibit resistance to numerous aboveground lepidopteran pests. This study shows that these genotypes are able to significantly reduce the survival of two root feeding pests, western corn rootworm, Diabrotica virgifera virgifera LeConte, and southern corn rootworm, Diabrotica undecimpunctata howardi Barber. The results also demonstrated that feeding by the aboveground herbivore fall armyworm, Spodoptera frugiperda (J. E. Smith), before infestation by western corn rootworm reduced survivorship of western corn rootworm in the root tissues of some, but not all, genotypes. Likewise, the presence of western corn rootworm in the soil seemed to increase resistance to fall armyworm in the whorl in several genotypes. However, genotypes derived from the Antiguan germplasm with genetic resistance to lepidopterans were still more resistant to the fall armyworm and both rootworm species than the susceptible genotypes even after defense induction. These results suggest that there may be intraplant communication that alters plant responses to aboveground and belowground herbivores.     

Effects of host plant architecture on colonization by galling insects

Abstract: To study the abundance and occurrence of herbivore insects on plants it is important to consider plant characteristics that may control the insects. In this study the following hypotheses were tested: (i) an increase of plant architecture increases species richness and abundance of gall‐inducing insects and (ii) plant architecture increases gall survival and decreases parasitism. Two hundred and forty plants of Baccharis pseudomyriocephala Teodoro (Asteraceae) were sampled, estimating the number of shoots, branches and their biomass. Species richness and abundance of galling insects were estimated per module, and mortality of the galls was assessed. Plant architecture influenced positively species richness, abundance and survival of galls. However, mortality of galling insects by parasitoids was low (13.26%) and was not affected by plant architecture, thus suggesting that other plant characteristics (a bottom‐up pressure) might influence gall‐inducing insect communities more than parasitism (a top‐down pressure). The opposite effect of herbivore insects on plant characteristics must also be considered, and such effects may only be assessed through manipulative experiments.     

Effects of genotypic variation in stem solidity on parasitism of a grass-mining insect

Host plant traits can play a significant role in influencing the importance, direction and intensity of tri-trophic interactions by both direct and indirect pathways. A major goal in applied tri-trophic research has been to determine whether breeding for host plant resistance traits can be combined with biological control to develop a more comprehensive control strategy. An important component of developing such a strategy is understanding how host resistance traits affect natural enemy–prey interactions for important pest insects. Here we examine the influence of genotypic variation in stem solidity, the primary trait conferring resistance against the wheat stem sawfly, Cephus cinctus, on parasitism of this major pest of wheat by its native braconid parasitoids. To do so, we conducted a field experiment in which we established replicate plots of 23 wheat genotypes that varied in levels of stem solidity, and quantified herbivore abundance and levels of parasitism across three sites in two years. Increasing stem solidity was associated with an approximately four-fold reduction in average parasitism rates, both across experimental plots and across wheat genotypes. Our analyses suggest that these effects were primarily direct, rather than indirectly mediated via effects of stem solidity on herbivore infestation levels or density. Interestingly, wheat genotype also had a significant influence on levels of parasitism, independent of its effects on stem solidity. Overall, our results suggest that although increasing stem solidity generally reduces parasitism, significant genotypic variability in average parasitism levels exist within solidity categories. Thus it may be possible to select resistant solid stemmed genotypes that also maintain relatively high parasitism levels. To our knowledge, our study is among the first to demonstrate a strong direct effect of genotypic variation in stem solidity on parasitism of grass mining insects, with important applied implications.     

Landscape diversity slows the spread of an invasive forest pest species

According to the associational resistance hypothesis, diverse habitats provide better resistance to biological invasions than monocultures. Host‐plant abundance has been shown to affect the range expansion of invasive pests, but the effect of landscape diversity (i.e. density of host/non‐host patches and diversity of forest habitat patches) on invasions remains largely untested. We used boundary displacement models and boosted regression tree analyses to investigate the effects of landscape diversity on the invasion of Corsica by the maritime pine bast scale Matsucoccus feytaudi over an 18‐yr period. Taking the passive wind dispersal of the scale into account, we showed that open habitats and connectivity between host patches accelerated spread by up to 13%, whereas landscapes with high tree diversity and a high density of non‐host trees decreased scale spread by up to 14%. We suggest a new mechanism for such associational resistance to pest invasion at the landscape level, which we term ‘the pitfall effect’.     

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.     

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.     

Plant population size and isolation affect herbivory of <Emphasis Type="Italic">Silene latifolia</Emphasis> by the specialist herbivore <Emphasis Type="Italic">Hadena bicruris</Emphasis> and parasitism of the herbivore by parasitoids

Habitat fragmentation can affect levels of herbivory in plant populations if plants and herbivores are differentially affected by fragmentation. Moreover, if herbivores are top–down controlled by predators or parasitoids, herbivory may also be affected by differential effects of fragmentation on herbivores and their natural enemies. We used natural Silene latifolia populations to examine the effects of plant population size and isolation on the level of herbivory by the seed predating noctuid Hadena bicruris and the rate of parasitism of the herbivore by its parasitoids. In addition, we examined oviposition rate, herbivory and parasitism in differently sized experimental populations. In natural populations, the level of herbivory increased and the rate of parasitism decreased with decreasing plant population size and increasing degree of isolation. The number of parasitoid species also declined with decreasing plant population size. In the experimental populations, the level of herbivory was also higher in smaller populations, in accordance with higher oviposition rates, but was not accompanied by lower rates of parasitism. Similarly, oviposition rate and herbivory, but not parasitism rate, increased near the edges of populations. These results suggests that in this system with the well dispersing herbivore H. bicruris, habitat fragmentation increases herbivory of the plant through a behavioural response of the moth that leads to higher oviposition rates in fragmented populations with a reduced population size, increased isolation and higher edge-to-interior ratio. Although the rate of parasitism and the number of parasitoid species declined with decreasing population size in the natural populations, we argue that in this system it is unlikely that this decline made a major contribution to increased herbivory.     

Tritrophic interactions and trade-offs in herbivore fecundity on hybridising host plants

Abstract. 1. Interspecific plant hybridisation can have important evolutionary consequences for hybridising plants and for the organisms that they interact with on multiple trophic levels. In this study the effects of plant hybridisation on the abundance of herbivores and on the levels of herbivore parasitism were investigated. 2. Borrichia frutescens, B. arborescens, and their hybrid (B. × cubana) were censused for Asphondylia borrichiae galls and Pissonotus quadripustulatus plant hoppers in the Florida Keys. Levels of egg parasitism were determined by dissecting parental and hybrid stems and galls for herbivore and parasite eggs and larvae. Stem toughness and gall size are plant‐mediated modes of protection from parasitism and these were also measured. For gall midges, fly size was measured as an estimate of fecundity. 3. Field censuses indicated that herbivore abundances varied on hybrid hosts relative to parent plant species and that the different herbivore species exhibited different patterns of abundance. Asphondylia borrichiae gall numbers followed the additive pattern of abundance while P. quadripustulatus numbers most closely resembled the dominance pattern. 4. Parasitism of P. quadripustulatus eggs was high on B. frutescens and the hybrids, and low on B. arborescens, which also had significantly tougher stems. Asphondylia borrichiae suffered the highest levels of parasitism on B. frutescens, the host plant which produced the smallest galls. On B. arborescens, which produced the largest galls, levels of A. borrichiae parasitism were lowest. Both parasitism and gall size were intermediate on the hybrid plants. Galls on B. arborescens and hybrid plants produced significantly smaller flies then those from B. frutescens suggesting that, when selecting hosts from among parent species and hybrids, gall flies may face a trade‐off between escape from natural enemies and maximising fecundity.     

Observational evidence of herbivore‐specific associational effects between neighboring conspecifics in natural,dimorphic populations of Datura wrightii

Associational effects—in which the vulnerability of a plant to herbivores is influenced by its neighbors—have been widely implicated in mediating plant–herbivore interactions. Studies of associational effects typically focus on interspecific interactions or pest–crop dynamics. However, associational effects may also be important for species with intraspecific variation in defensive traits. In this study, we observed hundreds of Datura wrightii—which exhibits dimorphism in its trichome phenotype—from over 30 dimorphic populations across California. Our aim was to determine whether a relationship existed between the trichome phenotype of neighboring conspecifics and the likelihood of being damaged by four species of herbivorous insects. We visited plants at three timepoints to assess how these effects vary both within and between growing seasons. We hypothesized that the pattern of associational effects would provide rare morphs (i.e., focal plants that are a different morph than their neighbors) with an advantage in the form of reduced herbivory, thereby contributing to the negative frequency‐dependent selection previously documented in this system. We found the best predictor of herbivory/herbivore presence on focal plants was the phenotype of the focal plant. However, we also found some important neighborhood effects. The total number of plants near a focal individual predicted the likelihood and/or magnitude of herbivory by Tupiochoris notatus, Lema daturaphila, and Manduca sexta. We also found that velvety focal plants with primarily sticky neighbors are more susceptible to infestation by Tupiochoris notatus and Lema daturaphila. This does not align with the hypothesis that associational effects at the near‐neighbor scale contribute to a rare‐morph advantage in this system. Overall, the results of our study show that the number and trichome‐morph composition of neighboring conspecifics impact interactions between D. wrightii and insect herbivores.