Matrix habitat and plant damage influence colonization of purple loosestrife patches by specialist leaf-beetles

The characteristics of the matrix, that is, the unsuitable habitat connecting host-plant patches may facilitate or limit herbivore movement thus affecting their population dynamics. We evaluated the effect of matrix habitat, distance between patches, and plant damage on movement of two leaf-beetles (Galerucella calmariensis Linnaeus and G. pusilla Duft) introduced to North America as biocontrol agents of the invasive purple loosestrife (Lythrum salicaria Linnaeus). Mark-recapture/resight experiments indicated (1) that leaf-beetles are more likely to colonize purple loosestrife patches surrounded by meadow than forest; (2) that previously attacked purple loosestrife plants are more likely to be colonized by Galerucella spp. than unattacked plants, especially in the forest habitat; and (3) that leaf beetle colonization of purple loosestrife decreased with distance from release point. Low colonization rates of purple loosestrife patches embedded in forests suggest either insufficient detection or active avoidance of such habitats. Biological control programs intend to manage dispersal of specialized insect herbivores for the purpose of sufficient and sustained control of their host plants. Such management needs to be informed by knowledge of interactions of habitat structure, plant damage, and dispersal capabilities of herbivores to facilitate release programs and control at the local and regional level.     

Interactions of root and leaf herbivores on purple loosestrife (<Emphasis Type="Italic">Lythrum salicaria</Emphasis>)

Interspecific interactions of herbivores sharing a host plant may be important in structuring herbivore communities. We investigated host plant-mediated interactions of root (Hylobius transversovittatus) and leaf herbivores (Galerucella calmariensis), released to control purple loosestrife (Lythrum salicaria) in North America, in field and potted plant experiments. In the potted plant experiments, leaf herbivory by G. calmariensis reduced H. transversovittatus larval survival (but not larval development) but did not affect oviposition preference. Root herbivory by H. transversovittatus did not affect either G. calmariensis fitness or oviposition preference. In field cage experiments, we found no evidence of interspecific competition between root and leaf herbivores over a 4-year period. Our data suggest that large populations of leaf beetles can negatively affect root-feeding larvae when high intensity of leaf damage results in partial or complete death of belowground tissue. Such events may be rare occurrences (or affected by experimental venue) since field data differed from data obtained from potted plant experiments, particularly at high leaf beetle densities. Interspecific interactions between G. calmariensis and H. transversovittatus are possible and may negatively affect either species, but this is unlikely to occur unless heavy feeding damage results in partial or complete plant death.     

Strong effects of hydrologic environment and weak effects of elevated CO<Subscript>2</Subscript> on the invasive weed <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> and the biocontrol beetle <Emphasis Type="Italic">Agasicles hygrophila</Emphasis>

Global change, such as elevated CO₂, may alter interactions between invasive plants and biocontrol agents, impacting biocontrol efficacy. Here, we conducted four experiments in Texas, USA to test how elevated CO₂ influences an invasive plant (Alternanthera philoxeroides) and its interactions with an introduced biocontrol beetle (Agasicles hygrophila) in terrestrial (well-watered) and flooded environments. We grew plants for 9 months in ambient or elevated CO₂ (800 ppm) chambers in continuously flooded or well-watered conditions. In no-choice trials, flooding increased leaf toughness and decreased beetle consumption but beetles only oviposited on ambient CO₂ leaves. In choice trials, beetles preferred to feed and oviposit on terrestrial plants but were also less likely to damage elevated CO₂ leaves. Caged beetle populations were larger in terrestrial conditions than aquatic conditions for a second set of plants grown in the chambers. With a third set of plants grown in the ambient or elevated CO₂ chambers, damage for plants placed in the field (aquatic setting) was higher for plants grown in terrestrial conditions vs. flooded conditions at ambient CO₂. Our results suggest that elevated CO₂ will have minor effects on the efficacy of this biocontrol agent by decreasing oviposition and number of leaves damaged, and hydrologic environment may affect invasive plant performance by altering herbivore oviposition and feeding preferences. A broader understanding of the effects of global change on biocontrol will help prevent and manage future spread of invasive plants.     

Impact of Galerucella birmanica (Coleoptera: Chrysomelidae) on growth and seed production of Trapa natans

Success in biological weed control programs depends upon the ability of host-specific herbivores to suppress populations of their host plant. While pre-release predictions of field host range (i.e., specificity) appear widely accurate, predictions about which agent or agent combination may suppress plant populations have lately been compared to predictions in a lottery. The history of weed biocontrol does not offer immediately obvious approaches to improve the lottery model, however, pre-release assessments of the impact of different herbivore densities on the invasive plant may provide an opportunity to improve predictions of success. In this paper, we report on the impact of the leaf beetle Galerucella birmanica on growth and reproduction of water chestnut, Trapa natans, in the native range in China. At low herbivore densities (10–50 larvae/rosette), plants compensated for leaf herbivory by increasing leaf production at the expense of reproductive effort. Inoculating >50 first instar larvae per rosette greatly suppressed biomass production and plants were unable to grow when three or more G. birmanica pairs were released per seven rosettes. In the native range, similar densities are found in the field, resulting in complete defoliation of T. natans. Our study indicates that G. birmanica feeding has significant negative impacts on T. natans. This chrysomelid species appears to be a promising biological control agent and we would predict that the species will be able to attain sufficiently high populations to control its host plant—if approved for release in North America.     

Dispersal of Galerucella pusilla and G. calmariensis via passive water transport in the Columbia River Estuary

Dispersal of biological control agents and their subsequent population growth can be a major determinant of the success of landscape-scale weed control programs. Biocontrol agents must be able to disperse across the distances between patches of host plants in order to colonize and control their targets. The presence of three species of biocontrol agents for purple loosestrife (Lythrum salicaria L.): Galerucella calmariensis L. (Coleoptera: Chrysomelidae), Galerucella pusilla Duftschmid (Coleoptera: Chrysomelidae), and Nanophyes marmoratus Goeze (Coleoptera: Brentidae), on relatively remote islands in the Columbia River Estuary (CRE) indicate that these organisms have the ability to disperse across large expanses of open flowing water to colonize remote sites. Previous studies suggest that colonization of these islands by active flight is highly unlikely; therefore, some other dispersal mechanism must be responsible for colonization. A spatial database of all known biocontrol agent release sites for purple loosestrife within 68 river kilometers of our CRE study area was developed and field surveys for biocontrol agents were conducted. A GIS was used to model dispersal distances between biocontrol agent recovery sites and the nearest conspecific release site. Tidal water flow within the CRE was assessed as a potential dispersal mechanism across the modeled distances. The ability of the biocontrol agents to withstand submersion was evaluated in field tests. Our results indicate that it is highly likely that passive water transport has been responsible for some of the long-distance open-water dispersal that would have been necessary for colonization of the remote islands where biocontrol agents were recovered.     

Quantifying the effects of distance and conspecifics on colonization: experiments and models using the loosestrife leaf beetle, Galerucella calmariensis

The ability of an insect to disperse to new habitat patches is difficult to quantify, but key to the establishment and persistence of populations. In this study, we examined dispersal of the phytophagous chrysomelid beetle, Galerucella calmariensis, which is currently being introduced into North America for the biological control of purple loosestrife (Lythrum salicaria), an aggressive wetland weed. We used a mark, release, and recapture approach to determine how rates of colonization of host patches by this beetle are influenced by the distance of the patch from the source of dispersers, and by the presence of conspecifics at the patch. We released color-coded beetles at six distances from a long, linear patch of purple loosestrife that was divided into segments with and without conspecifics. We observed initial flight directions as beetles left the release points and collected all beetles that settled at the target patch. We found a bias in initial flight toward the target for distances up to 50 m. Over the 7 days of the experiment, beetles arrived at the target from all release points, including the farthest release point, 847 m away. G. calmariensis was strongly attracted to conspecifics when settling after dispersal; 86% of the 582 recovered beetles came from the segments inhabited by conspecifics. The probability of an individual arriving at the patch declined steeply with release distance. This relationship fits a model in which beetles move in a random direction and stop if they intercept the target patch, and where beetles are lost at a constant rate with distance travelled. The dispersal and patch-colonizing behavior of G. calmariensis is likely to have important consequences for the biological control program against purple loosestrife. Received: 23 January 1996 / Accepted:30 September 1996     

Experimental Invasions Using Biological Control Introductions: The Influence of Release Size on the Chance of Population Establishment

Introductions of biological control organisms offer a unique opportunity to experimentally study the process of invasion by exotic species. I used two chrysomelid beetles, Galerucella calmariensis and Galerucella pusilla, which are currently being introduced into North America for the biological control of purple loosestrife (Lythrum salicaria), to determine how the initial size of a release affects the probability that the introduced population grows and persists. I released both species into stands of their host plant at 36 sites scattered throughout central New York State using four release sizes: 20, 60, 180, and 540. I returned to these sites over the next 3 years to census the populations. For both species, the probability of population establishment increased with release size. Population growth rates also depended positively on release size. The implication from these results is that the demographic factors whose influence depends on population size or density such as demographic stochasticity, Allee effects, and genetics play important roles in the establishment of invading populations. A second set of releases was used to determine if it was at all possible for a single gravid female to found a population. Out of twenty individual females released, one female (a G. calmariensis) founded a population that persisted until the end of the study (3 generations). This revised version was published online in July 2006 with corrections to the Cover Date.     

Herbivore community promotes trait evolution in a leaf beetle via induced plant response

Several recent studies have emphasised that community composition alters species trait evolution. Here, we demonstrate that differences in composition of local herbivore communities lead to divergent trait evolution of the leaf beetle Plagiodera versicolora through plant‐mediated indirect interactions. Our field surveys, genetic analyses and community‐manipulation experiments show that herbivore community composition determines the degree of herbivore‐induced regrowth of willows (Salicaceae), which in turn, promotes the divergent evolution of feeding preference in the leaf beetle from exclusive preference for new leaves to a lack of preference among leaf‐age types. Regrowth intensity depends both on the differential response of willows to different herbivore species and the integration of those herbivore species in the community. Because herbivore‐induced regrowth involves phenological changes in new leaf production, leaf beetle populations develop divergent feeding preferences according to local regrowth intensity. Therefore, herbivore community composition shapes the selection regime for leaf beetle evolution through trait‐mediated indirect interactions.     

Effects of plant quality and ant defence on herbivory rates in a neotropical ant‐plant

1. Understanding the degree to which populations and communities are limited by both bottom‐up and top‐down effects is still a major challenge for ecologists, and manipulation of plant quality, for example, can alter herbivory rates in plants. In addition, biotic defence by ants can directly influence the populations of herbivores, as demonstrated by increased rates of herbivory or increased herbivore density after ant exclusion. The aim of this study was to evaluate bottom‐up and top‐down effects on herbivory rates in a mutualistic ant‐plant. 2. In this study, the role of Azteca alfari ants as biotic defence in individuals of Cecropia pachystachya was investigated experimentally with a simultaneous manipulation of both bottom‐up (fertilisation) and top‐down (ant exclusion) factors. Four treatments were used in a fully factorial design, with 15 replicates for each treatment: (i) control plants, without manipulation; (ii) fertilised plants, ants not manipulated; (iii) unfertilised plants and excluded ants and (iv) fertilised plants and ants excluded. 3. Fertilisation increased the availability of foliar nitrogen in C. pachystachya, and herbivory rates by chewing insects were significantly higher in fertilised plants with ants excluded. 4. Herbivory, however, was more influenced by bottom‐up effects – such as the quality of the host plant – than by top‐down effects caused by ants as biotic defences, reinforcing the crucial role of leaf nutritional quality for herbivory levels experienced by plants. Conditionality in ant defence under increased nutritional quality of leaves through fertilisation might explain increased levels of herbivory in plants with higher leaf nitrogen.     

Bottom‐up effects of glucosinolate variation on aphid colony dynamics in wild cabbage populations

1. There is an ongoing debate about the relative importance of top‐down and bottom‐up regulation of herbivore dynamics in the wild. Secondary metabolites, produced by plants, have negative effects on survival and growth of some herbivore species, causing bottom‐up regulation of population dynamics. Herbivore natural enemies may use plant secondary metabolites as cues to find their prey, but their survival and reproduction can also be influenced by the upward cascade of secondary metabolites through the food web. Thus plant chemistry might also affect herbivore populations by mediating top‐down regulation. 2. We investigated the influence of heritable variation in aliphatic glucosinolates, a class of secondary metabolites produced by Brassica plants, on the relative importance of top‐down and bottom‐up regulation of Brevicoryne brassicae (mealy cabbage aphid) colonies in natural Brassica oleracea (wild cabbage) populations. We manipulated natural enemy pressure on plants differing in their glucosinolate profiles, and monitored aphid colony growth and disperser production. 3. Aphid colony sizes were significantly smaller on plants producing sinigrin, compared with plants producing alternative aliphatic glucosinolates. Aphid natural enemy numbers correlated with aphid colony size, but there was no additional effect of the plants' chemical phenotype on natural enemy abundance. Furthermore, experimental reduction of natural enemy pressure had no effect on aphid colony size or production of winged dispersers. 4. Our results provide evidence for glucosinolate‐mediated, bottom‐up regulation of mealy cabbage aphid colonies in natural populations, but we found no indication of top‐down regulation. We emphasise that more studies of these processes should focus on tritrophic interactions in the wild.     

Population Establishment, Dispersal, and Impact of Galerucella pusilla and G. calmariensis, Introduced to Control Purple Loosestrife in Central Ontario

As part of a biological control program against purple loosestrife (Lythrum salicaria), two species of exotic leaf-grazing beetles, Galerucella pusilla and G. calmariensis, have been introduced throughout North America. To assess the potential of these species to establish in an area approaching the northern limit of L. salicaria, we studied beetle population characteristics and feeding impact on L. salicaria at wetlands near North Bay, Ontario, Canada, where each beetle species was released on a separate site in 1995. Weekly counts of adults and larvae of each Galerucella species were made on tagged L. salicaria plants throughout the growing season from 1996 to 1998. At the G. pusilla release site (Site 1), the abundance and frequency of beetles decreased significantly (P < 0.05) from 1996 to 1998, life history and development patterns were inconsistent and incomplete, and dispersal activity was low. Over the same time period at the G. calmariensis release site (Site 2), the abundance of beetles increased significantly (P < 0.05), frequency remained stable, life history and development patterns were consistent and complete, and adults dispersed actively. In the third year following release, G. calmariensis exhibited a 17-week active period and produced three generations of adults at Site 2. Leaf area analysis on plants harvested in 1998 suggested that the impact of both species on L. salicaria was low. Feeding produced a 4.5 and 3.5% leaf tissue loss at Sites 1 and 2, respectively. The frequency of shot-hole feeding (specific to Galerucella spp.) was 32% at Site 2, where G. calmariensis was established, compared to 10% at Site 1 (where G. pusilla failed to establish) and at a control site. Based on these results, it appears that there is a potential for effective control of L. salicaria in central Ontario using G. calmariensis.     

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

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

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

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

Intra- and interspecific density dependence mediates weather effects on the population dynamics of a plant–insect herbivore system

Temperature and precipitation are two major factors determining arthropod population densities, but the effects from these climate variables are seldom evaluated in the same study system and in combination with inter- and intraspecific density dependence. In this study, I used a 19 year time series on plant variables (shoot height and flowering incidence) and insect density in order to understand direct and indirect effects of climatic fluctuations on insect population densities. The study system includes two closely related leaf beetle species (Galerucella spp.) and a flower feeding weevil Nanophyes marmoratus attacking the plant purple loosestrife Lythrum salicaria. Results suggest that both intraspecific density dependence and weather variables affected Galerucella population densities, with interactive effects of rain and temperature on insect densities that depended on the timing relative to insect life cycles. In spring, high temperatures increased Galerucella densities only when combined with high rain, as low rain implies a high drought risk. Low temperatures are only beneficial if combined with little rain, as high rain cause chilly and wet conditions that are bad for insects. In summer, interactive effects of rain and temperature are different because high temperatures and little rain cause drought that induce wilting in plants, thus reducing food availability for the leaf feeding larvae. In contrast, the density of the flower feeding weevil was less affected by temperature and precipitation directly, and more indirectly interspecific density dependent effects through reduced resource availability caused by previous Galerucella damage.     

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.     

Risk to non-target plants from Charidotis auroguttata (Chrysomelidae: Coleoptera), a potential biocontrol agent for cat’s claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia

The potential of the leaf beetle Charidotis auroguttata as a biocontrol agent for cat’s claw creeper Macfadyena unguis-cati (Bignoniaceae), an environmental weed in Australia, and risk to non-target plants was evaluated under quarantine conditions. In no-choice tests, C. auroguttata adults and larvae fed on many plant species across different families, but egg to adult development occurred only on the target weed. However, when neonate larvae from the target weed were transferred onto Myoporum boninense australe (Myoporaceae), a non-target native plant, 11.7% completed development, as compared to 95% of larvae that completed development on the target weed. Larval development on this non-target species also took twice as long as on the target weed. No larvae completed development on other test plants. In choice tests, leaf area consumption by adults and larvae was significantly more on the target weed than on other plants, and oviposition occurred only on the target weed. In the no-choice demography trials, adults laid eggs from the second week after emergence on the target weed, with an average of 0.286 eggs/female/day, resulting in an 18-fold increase in the adult population over 16 weeks. On My. boninense australe adult survival remained high, but oviposition commenced only from the 10th week after emergence with an average of 0.023 eggs/female/day, and none of the eggs developed into adults. In the choice demography trials, oviposition on the target weed was evident from the fourth week onwards, while on the non-target plant oviposition commenced only from the 14th week. Only 10% of total adults and 11.3% of total eggs were found on the non-target plant, and none of these eggs developed into adults. Although the biocontrol agent can ‘spill-over’ from the target weed to the non-target native plant and cause adult feeding damage, the non-target plant could not sustain a viable insect population on its own. This agent was not approved for field release in Australia due to perceived risk to non-target species.     

Natural Enemies ofTrapaspp. in Northeast Asia and Europe

Trapa natansis an aquatic plant native to the Old World. In the late 1880s this plant was found to be naturalized in the eastern United States where it has become an important weed. To evaluate the potential of biological control to reduce this weed, surveys for natural enemies ofTrapaspp. were made in Northeast Asia and Europe. In addition, populations ofTrapa japonicain South Korea were monitored for one season to evaluate the occurrence and impact of insect natural enemies. Among the insects found, the leaf beetleGalerucella birmanicawas the most common and damaging species in Asia. It can cause complete defoliation of whole populations of plants. Nymphuline pyralid moths were also common and at times damaging. Both the beetle and the moths feed and develop on unrelated plants and so have no potential as biological control agents ofT. natansin North America. TwoNanophyesweevils were found in Asia which feed in petiole floats of the leaves. They are thought to be specific toTrapabut were not observed to be damaging. Low level populations of polyphagous Homoptera were common. Chironomid midges were also frequently associated with the plants, but for the most part were filter feeders, not herbivores. The greatest impact on the monitored plants in South Korea was by the leaf beetleG. birmanicawhich defoliated most of the plants, causing the mats of plants to sink. In Europe, a similar and related insect fauna was found, but none of the insects was very damaging to the plants. One Italian weevil,Bagous rufimanus,feeds within the fruit stems and might be more damaging at higher than observed population levels. Some diseases were seen on some plants in both regions. European populations ofT. natanshave declined significantly, stimulating conservation efforts. Literature records ofTrapainsects from warm areas are presented. These could be of interest ifT. natansspreads into warmer areas of North America.     

Effects of local vegetation and plantation age for the parasitoid Asecodes mento– a biocontrol agent in organic strawberry fields

Abstract The parasitoid Asecodes mento (Walker, 1839) (Hymenoptera: Eulophidae) is the most important biocontrol agent of the strawberry leaf beetle Galerucella tenella (L.) (Coleoptera: Chrysomelidae) in northern Europe. Here, I investigated whether natural parasitism in organic strawberry plantations was affected by the presence of the alternative host plant meadowsweet (Filipendula ulmaria), and whether parasitism rates differed between plantations of different ages (6 to 79 years). I also investigated whether parasitoid brood size, body size and sex ratio differed between the two host plants in the field. Parasitism was very low (0%) in newly established plantations and increased to a plateau (～40%) in fields where strawberries had been grown for approximately 20 years or longer. Such an extended colonization process is unacceptable for commercial growers. It would thus be desirable to find a method to catalyze parasitoid population buildup in young plantations. Parasitoid brood sizes were larger in beetles collected from meadowsweet, while body size and sex ratio did not differ between parasitoids collected from the two plants. These findings suggest that meadowsweet can export parasitoids to neighboring strawberry fields. Although this is a possibility, I did not find any significant differences in parasitism rates between isolated strawberry fields and fields adjacent to meadowsweet stands, indicating that effects of local vegetation are small on parasitism rates. Releasing parasitoids in newly established strawberry plantations may be a better strategy for quickly obtaining high parasitism than intercropping with meadowsweet.     

Integrating Management Techniques to Restore Sites Invaded by Mile‐a‐Minute Weed,Persicaria perfoliata

Efforts to suppress an invasive weed are often undertaken with the goal of facilitating the recovery of a diverse native plant community. In some cases, however, reduction in the abundance of the target weed results in an increase in other exotic weeds. Mile‐a‐minute weed (Persicaria perfoliata (L.) H. Gross (Polygonaceae)) is an annual vine from Asia that has invaded the eastern United States, where it can form dense monocultures. The host‐specific Asian weevil Rhinoncomimus latipes Korotyaev (Coleoptera: Curculionidae) was first released in the United States in 2004 as part of a classical biological control program. At three sites invaded by mile‐a‐minute weed, biological control was integrated with pre‐emergent herbicide use and two densities of native plantings. After 2 years, native plant cover differed significantly and was greater than 80% in the plots with plantings and pre‐emergent herbicide but less than 30% in the planting treatments without herbicide. Where mile‐a‐minute cover decreased at the two sites with the greatest pressure from exotic plants, plots were dominated by another exotic weed, Microstegium vimineum (Trin.) A. Camus, Japanese stiltgrass. The combination of biocontrol, pre‐emergent herbicide, and revegetation with native plants suppressed mile‐a‐minute weed, prevented invasion by Japanese stiltgrass, and increased the abundance of native plants. The selection of the management strategies used to control mile‐a‐minute weed determined the extent of recovery of the native plant community.     

Spatial structure-mediated indirect effects of an aquatic plant, Trapa japonica, on interaction between a leaf beetle, Galerucella nipponensis, and a water strider, Gerris nepalensis

The spatial structure-mediated indirect effects of an aquatic host plant, Trapa japonica (Trapaceae), on survival of a leaf beetle, Galerucella nipponensis (Coleoptera: Chrysomelidae), with regard to predation by a water strider, Gerris nepalensis (Hemiptera: Gerridae), were investigated in an irrigation pond and under indoor conditions. Beetle density, particularly in the first-instar stage of the larvae, became extremely low in the second generation in the pond, even though food resources were abundant. The biological check method, which excludes natural enemies by caging, suggested that a causal factor of the low density was predation by Gerris nepalensis in midsummer and thereafter. Normally growing rosettes extend their leaves horizontally on the water surface. However, some rosettes of T. japonica grew leaves vertically in the later seasons, and such overgrown rosettes were distributed patchily in the ponds. Beetle density was higher on the latter rosettes than on the former ones. The indoor experiment showed that the water strider can feed on the eggs of beetles on normally grown rosettes but not on the overgrown rosettes. Furthermore, female adults of Galerucella nipponensis preferred the overgrown rosettes over the normally grown rosettes for oviposition when water striders were present. These facts suggest that the spatial structure-mediated indirect effects of the host plant modify the interaction between herbivore and predator. Received: November 22, 2001 / Accepted: February 26, 2002