Visual Active Space of the Milfoil Weevil, <Emphasis Type="Italic">Euhrychiopsis lecontei</Emphasis> Dietz (Coleoptera: Curculionidae)

Euhrychiopsis lecontei Dietz (Coleoptera: Curculionidae), a native weevil, is used as a biological control agent for the invasive aquatic macrophyte, Eurasian watermilfoil (Myriophyllum spicatum L.). Because E. lecontei overwinters on land in the adult stage and must find plants in lakes each spring, plant finding behaviors are essential to eventually understanding and predicting long term biological control. Our research showed that E. lecontei is visually attracted to M. spicatum at up to 17.5 cm, and is more attracted to plants than other visual stimuli within 15 cm. We also showed that turbidity may affect visual plant finding at 15 cm. Using available data from this and other previous studies involving chemical cues and other life history traits, we propose a testable conceptual model for how E. lecontei finds plants each year, especially while underwater. This model may also be used to explain plant finding by aquatic phytophagous insects in general.     

Oviposition specificity and behavior of the watermilfoil specialist Euhrychiopsis lecontei

Eurasian watermilfoil (Myriophyllum spicatum L.) is a nuisance aquatic weed, exotic to North America. The freshwater weevil Euhrychiopsis lecontei (Dietz) is a potential control agent of Eurasian watermilfoil and is a fully submersed aquatic specialist herbivore. Its presumed original host is the native northern watermilfoil (Myriophyllum sibiricum Komarov). We conducted a set of oviposition experiments to reveal first and second oviposition preference of Euhrychiopsis lecontei when presented with seven macrophytes. We tested differences between source (lake) populations of weevils, differences in behavior between weevils reared on the exotic Eurasian watermilfoil and the native northern watermilfoil and between weevils in the presence and absence of their preferred hostplant. Oviposition assays confirmed that E. lecontei is a watermilfoil specialist. Out of the 207 females that laid eggs, only three oviposited on a non-watermilfoil plant, Megalodonta beckii. The weevils' degree of specificity was influenced by the watermilfoil species on which they were reared. Weevils reared on Eurasian watermilfoil tended to oviposit on Eurasian watermilfoil, spent more time on Eurasian watermilfoil than on other plants, and spent more time off plants and took longer to oviposit when Eurasian watermilfoil was removed. Weevils reared on northern watermilfoil did not exhibit a preference for either watermilfoil species in oviposition or in time allocation, although they oviposited on and spent significantly more time on watermilfoils than on other species. Rearing of the two populations on their complementary watermilfoil hostplant resulted in responses typical of the rearing plant, not the original host. These results show that although both weevil populations are watermilfoil specialists, Eurasian-reared weevils prefer Eurasian watermilfoil in general host attraction and oviposition, whereas northern-reared weevils do not. The results support the contention that E. lecontei may be a good biocontrol agent for Eurasian watermilfoil because of its high specificity. The results also suggest that the current host range expansion of the weevil to Eurasian watermilfoil has the potential to become a host shift due to the increased specificity. Herbivory in freshwater systems is not well studied, and the E. lecontei-M. spicatum relationship is a rare example of submersed freshwater specialist herbivore-host-plant interactions.     

Effects of eutrophication and snails on Eurasian watermilfoil (Myriophyllum spicatum) invasion

Exotic species can invade and establish new habitats both as a result of their own traits, and as a result of the characteristics of the environment they invade. Here, we show that the abundance of the invasive submerged aquatic plant, Myriophyllum spicatum (Eurasian watermilfoil) is highly dependent on the conditions of the environment in a mesocosm experiment. M. spicatum is allelopathic towards epiphytic algae, and in the absence of algivorous snails, we found that the abundance of both algae and M. spicatum significantly increased with experimentally increased nutrient loading, while the abundance of native submerged macrophytes declined. However, when snails were present, snail biomass increased with increasing nutrient loading, and M. spicatum biomass was consistently low while native submerged macrophyte biomass was consistently high. Our results stress the importance of the interaction between species traits and environmental conditions when considering the invasiveness of certain exotic species and the invasibility of certain environments.     

Vegetative Propagule Pressure and Water Depth Affect Biomass and Evenness of Submerged Macrophyte Communities

Vegetative propagule pressure may affect the establishment and structure of aquatic plant communities that are commonly dominated by plants capable of clonal growth. We experimentally constructed aquatic communities consisting of four submerged macrophytes (Hydrilla verticillata, Ceratophyllum demersum, Elodea nuttallii and Myriophyllum spicatum) with three levels of vegetative propagule pressure (4, 8 and 16 shoot fragments for communities in each pot) and two levels of water depth (30 cm and 70 cm). Increasing vegetative propagule pressure and decreasing water level significantly increased the growth of the submerged macrophyte communities, suggesting that propagule pressure and water depth should be considered when utilizing vegetative propagules to re-establish submerged macrophyte communities in degraded aquatic ecosystems. However, increasing vegetative propagule pressure and decreasing water level significantly decreased evenness of the submerged macrophyte communities because they markedly increased the dominance of H. verticillata and E. nuttallii, but had little impact on that of C. demersum and M. spicatum. Thus, effects of vegetative propagule pressure and water depth are species-specific and increasing vegetative propagule pressure under lower water level can facilitate the establishment success of submerged macrophyte communities.     

Natural enemies from South Africa for biological control of <Emphasis Type="Italic">Lagarosiphon major</Emphasis> (Ridl.) Moss ex Wager (Hydrocharitaceae) in Europe

The non-native invasive plant, Lagarosiphon major (Hydrocharitaceae) is a submersed aquatic macrophyte that poses a significant threat to water bodies in Europe. Dense infestations prove difficult to manage using traditional methods. In order to initiate a biocontrol programme, a survey for natural enemies of Lagarosiphon was conducted in South Africa. Several phytophagous species were recorded for the first time, with at least three showing notable promise as candidate agents. Amongst these, a leaf-mining fly, Hydrellia sp. (Ephydridae) that occurred over a wide distribution causes significant leaf damage despite high levels of parasitism by braconid wasps. Another yet unidentified fly was recorded mining the stem of L. major. Two leaf-feeding and shoot boring weevils, cf. Bagous sp. (Curculionidae) were recorded damaging the shoot tips and stunting the growth of the stem. Several leaf-feeding lepidopteran species (Nymphulinae) were frequently recorded, but are expected to feed on a wide range of plant species and are not considered for importation before other candidates are assessed. The discovery of several natural enemies in the country of origin improves the biological control prospects of L. major in Europe.     

Morphological and Chemical Changes Induced by Herbivory in Three Common Aquatic Macrophytes

The Dry Matter Content (DMC), the total phenolic content, the production of new branches and the plant fragmentation were compared in three macrophyte species (Elodea canadensis, Elodea nuttallii and Myriophyllum spicatum) exposed or not to snail herbivory. Grazing significantly reduced the DMC of M. spicatum and E. canadensis, but had no effect on the DMC of E. nuttallii. The phenolic contents of Elodea species were not modified by snail herbivory, whereas that of M. spicatum significantly increased when exposed to grazers. The number of new branches produced by M. spicatum and E. canadensis plants, and the fragmentation of E. canadensis also increased in response to herbivory. Chemical defences are therefore probably constitutive in Elodea and induced in M. spicatum, and morphological changes can be related to species growth form and synthesis of phenolic compounds. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Responses of four submerged macrophytes to freshwater snail density (Radix swinhoei) under clear‐water conditions: A mesocosm study

Macrophytes play a key role in stabilizing clear‐water conditions in shallow freshwater ecosystems. Their populations are maintained by a balance between plant grazing and plant growth. As a freshwater snail commonly found in shallow lakes, Radix swinhoei can affect the growth of submerged macrophytes by removing epiphyton from the surface of aquatic plants and by grazing directly on macrophyte organs. Thus, we conducted a long‐term (11‐month) experiment to explore the effects of snail density on macrophytes with distinctive structures in an outdoor clear‐water mesocosm system (with relatively low total nitrogen (TN, 0.66 ± 0.27 mg/L) and total phosphorus (TP, 36 ± 20 μg/L) and a phytoplankton chlorophyll a (Chla) range of 14.8 ± 4.9 μg/L) based on two different snail densities (low and high) and four macrophyte species treatments (Myriophyllum spicatum, Potamogeton wrightii, P. crispus, and P. oxyphyllus). In the high‐density treatment, snail biomass and abundance (36.5 ± 16.5 g/m² and 169 ± 92 ind/m², respectively) were approximately twice that observed in the low‐density treatment, resulting in lower total and aboveground biomass and ramet number in the macrophytes. In addition, plant height and plant volume inhabited (PVI) showed species‐specific responses to snail densities, that is, the height of P. oxyphyllus and PVI of M. spicatum were both higher under low‐density treatment. Thus, compared with low‐density treatment, the inhibitory effects of long‐term high snail density on macrophytes by direct feeding may be greater than the positive effects resulting from epiphyton clearance when under clear‐water conditions with low epiphyton biomass. Thus, under clear‐water conditions, the growth and community composition of submerged macrophytes could be potentially modified by the manual addition of invertebrates (i.e., snails) to lakes if the inhibitory effects from predatory fish are minor.     

Herbivory in omnivorous fishes: effect of plant secondary metabolites and prey stoichiometry

1. Many animals that consume freshwater macrophytes are omnivorous (i.e., they include both plant and animal matter in their diet). For invertebrate omnivorous consumers, selection of macrophyte species depends partly on the presence of secondary metabolites in plants, plant carbon/nutrient balances and/or physical structure of plants. However, little is known about the mechanisms influencing consumption of macrophytes in aquatic vertebrates. 2. For two fish species, the omnivorous rudd (Scardinius erythrophthalmus) and herbivorous grass carp (Ctenopharyngodon idella), feeding preferences were determined in three choice experiments. We tested (i) whether the presence of secondary metabolites and macrophyte stoichiometry affects macrophyte species selection by fish, (ii) the importance of macrophyte stoichiometry by manipulating the macrophytes experimentally and (iii) the rate of herbivory when the most palatable macrophyte is offered simultaneously with a common animal prey. 3. In a choice experiment with five species of submerged macrophytes (Callitriche sp., Chara globularis, Elodea nuttallii, Myriophyllum spicatum and Potamogeton pectinatus), Myriophyllum was clearly consumed least by both fishes, which strongly correlated with the highest phenolic concentration of this macrophyte. Additionally, a significant negative relationship was found between consumption and C : N ratio of the five macrophytes. The two most consumed macrophytes also had the lowest dry matter concentration (DMC). 4. In a second choice experiment, the C : N ratio of the least (Myriophyllum) and most (Potamogeton) palatable plants was manipulated by growing the macrophytes under fertilised and unfertilised conditions and subsequently feeding them to rudd. The avoidance of consumption of the chemically defended Myriophyllum by rudd was partly alleviated by the lowered C : N ratio. 5. The third choice experiment showed that both fishes preferred animal prey (the amphipod Gammarus pulex) over the most palatable macrophyte (Potamogeton) when offered simultaneously. The C : N ratio of the amphipods was about half that of the lowest C : N ratio measured in the macrophytes. Consumption by the fishes could not clearly be related to C : P or N : P ratios of prey items in any of the experiments. 6. We conclude that omnivorous fish avoid macrophytes that are chemically defended. However, when these defences are only minor, stoichiometry (C : N ratio) in combination with DMC may be a determining factor for consumption by vertebrate facultative herbivores.     

Impact in quarantine of the galling weevil Lepidapion argentatum on shoot growth of French broom (Genista monspessulana), an invasive weed in the western U.S.

In weed biocontrol, there is a need for pre-release efficacy assessments for potential agents. Genista monspessulana ((L.) L. A. S. Johnson (Fabaceae), French broom) is an invasive perennial shrub in the western U.S. The galling weevil Lepidapion argentatum Gerstaecker is a potential biocontrol agent. The impact of increasing weevil density on galling damage, plant height, width, leaf damage, and relative growth rate (RGR) was assessed in greenhouse experiments on two to three-month-old seedlings infested with either one or three weevils. Infestation by three female weevils caused 48% more galls producing 27% more larvae than did infestation with one female while causing only 1% leaf damage and no difference in total leaf area. Infestation with multiple weevils caused a 55% and 29% decrease in plant height and canopy width respectively, while single-weevil infestation decreased height by 32% and width to the same degree as for multiple weevils. The RGR of seedlings infested with three weevils was three times slower than the controls, while growth was reduced 2-fold by single-weevil infestation. Reductions in plant size and growth rate induced by weevil galling could reduce plant competitive survival to reproduction and also plant population dispersal as seedlings. Our results suggest that L. argentatum has the potential to cause impact to French broom seedlings if released in the invasive range.     

Catchment context and the bottom‐up regulation of the abundance of specialist semi‐aquatic weevils on water hyacinth

1. Bottom‐up regulation is prevalent in plant–herbivore interactions and is thought to be particularly important in the case of aquatic plants and their specialist insect herbivores. 2. Recently published mesocosm studies have shown that the abundance of specialist Neochetina weevils, N. bruchi and N. eichhorniae, on water hyacinth (Eichhornia crassipes) are principally under the influence of nutrients in plant tissues. 3. We examined historical patterns of the abundance of these species of semi‐aquatic weevils in two water bodies from catchments with significantly different nutrient loads in subtropical Australia to test the validity of the published conceptual model of bottom‐up regulation. 4. Our results revealed that these weevils are indeed under bottom‐up regulation under field conditions. However, the nature of this regulation appears to be influenced by the broader catchment context of the water hyacinth‐infested water body, with plant tissue nutrients influencing weevil abundance more in the catchment with lower nutrient run‐offs. 5. Our findings reaffirm the importance of bottom‐up regulation in plant–insect interactions, add to the growing evidence of indirect effects spanning terrestrial and aquatic ecosystems, and inform management of water hyacinth using these weevils as biocontrol agents.     

Size-related auto-fragment production and carbohydrate storage in auto-fragment of <Emphasis Type="Italic">Myriophyllum spicatum</Emphasis> L. in response to sediment nutrient and plant density

Size-related asexual reproduction of submersed macrophytes is still poorly understood. Here, we investigate how size-related auto-fragmentation in Myriophyllum spicatum L. responds to sediment nutrients and plant density. An experiment was carried out with sediments containing two different nutrient levels and with two levels of plant density. The results show that sediment nutrients and plant density brought about a strong dependency of auto-fragment production and the amount of total non-structural carbohydrate (TNC) storage in auto-fragments on individual plant size (total plant biomass). However, these two factors acted differently on size dependency. Sediment nutrients positively affected auto-fragment production and the amount of TNC in auto-fragments of M. spicatum. High concentrations of sediment nutrients significantly increased these two traits in absolute value and the value relative to plant size. Although the auto-fragment biomass and the amount of TNC in auto-fragments did not differ between density treatments when plant size was considered, the absolute values of these two traits were much larger in the low plant density treatment than in the high plant density treatment, which suggested an indirect negative effect of plant density on the auto-fragmentation of M. spicatum. In addition, higher percentages of large auto-fragments (>100 mg) were produced by plants that grew in nutrient poor sediment and low plant density environment than plants in nutrient rich sediment and high plant density environment. These results do not solely highlight a size-dependent effect, but also a size-independent effect of auto-fragment production and the amount of TNC in auto-fragments of M. spicatum. Furthermore, such size-independent effects can be explained by the significant biomass partitioning differences and the similar TNC-concentrations in auto-fragments under different environmental conditions.     

Food preference of freshwater invertebrates: comparing fresh and decomposed angiosperm and a filamentous alga

1. Fresh and decomposed Mougeotia sp. (a filamentous green alga) and Elodea nuttallii (a vascular plant) were offered as food to three species of aquatic macroinvertebrates (Lymnaea peregra, Asellus meridianus and Endochironomus albipennis) to test: (i) if filamentous algae are preferred to aquatic higher plants (hereafter, called ‘macrophytes’) and (ii), as is known for higher plants, if decomposition also results in greater palatability of filamentous algae. 2. Compared with the alga, the macrophyte in both states was of higher nutritional value. Conditioning improved the nutritional value of both food types, but especially of the macrophyte. 3. Both fresh alga and fresh macrophyte were eaten little by all animals except A. meridianus feeding upon the macrophyte. Consumption was higher for both plants in their decomposed state. However, L. peregra consumed more decomposed macrophyte than the decomposed alga. Both decomposed plants were eaten most by E. albipennis followed by A. meridianus and L. peregra. 4. Digestibility of both plants, but especially of the macrophyte, increased significantly after decomposition. The assimilation efficiencies of all animals on the fresh E. nuttallii were higher than on fresh Mougeotia sp. After decomposition, the efficiency increased significantly only on the alga. Consequently, both decomposed plants were assimilated with similar efficiency by all test animals. 5. Amongst aquatic macrophytes, the increase of their consumption and digestibility upon decomposition has hitherto been known only for vascular plants but not for filamentous algae.     

Influence of host log age and refuge from natural enemies on colonization and survival of Phoracantha semipunctata

The pine cone weevil, Pissodes validirostris Gyll. (Coleoptera: Curculionidae), attacks seed cones of most Eurasian pine species, except these of Swiss stone pine (Pinus cembra L.). Behavioural responses of adult weevils to cone volatile emissions of Swiss stone pine and to those of a common host, mountain pine (Pinus uncinata Ram.), were compared in an olfactometer. Weevils were significantly attracted by the volatile blend emitted by mountain pine, but Swiss stone pine volatiles elicited an inverse response, with most weevils moving in the opposite direction to the odour source. However, the majority of second instar weevil larvae that were extracted from mountain pine cones and transferred into Swiss stone pine cones were capable of developing to the adult stage. This suggests that Swiss stone pine cones do not contain strong feeding deterrents that could prevent larval development. The possible factors involved in the absence of colonization of Swiss stone pine cones by cone weevils are discussed.     

Prior host feeding experience influences ovipositional but not feeding preference in a polyphagous insect herbivore

Black vine weevils, Otiorhynchus sulcatus (Fabricius) (Coleoptera: Curculionidae), are globally‐distributed polyphagous pests of many horticultural crops. We investigated how adult weevils were affected by host switching and, in particular, how host plant species nutritional and defensive chemistry affected subsequent host plant species selection and oviposition. Adults were fed one of three host plant species, blackcurrant [Ribes nigrum L. (Grossulariaceae)], raspberry [Rubus idaeus L. (Rosaceae)], or strawberry [Fragaria x ananassa Duchesne (Rosaceae)], throughout their pre‐reproductive periods and then subjected to behavioral choice assays with these plants. Foliar chemistry differed significantly among the three host plant species. Compared to raspberry and strawberry foliage, blackcurrant foliage was 13% lower in nitrogen, 3% higher in carbon, and 28% higher in phenolic compounds. Initial host plant species had a significant effect on weevil mortality, with more weevils dying when previously fed blackcurrant (12%) than strawberry (3%) or raspberry (0%) regardless of subsequent host. Initial host plant species also affected oviposition, with weevils laying only ca. two eggs per week when previously fed blackcurrant, compared to those on raspberry or strawberry (ca. 11 and 15 eggs per week, respectively). When given a choice, weevils discriminated among host plant species and tended to oviposit on plants on which they had previously fed, even when the plant was nutritionally inferior for egg production and adult survival. In contrast, feeding behavior was only affected by the current host plant species. Feeding and oviposition were related to leaf chemistry only in blackcurrant, as leaf consumption was negatively correlated with foliar carbon and zinc concentrations, and positively correlated with foliar phosphorus and potassium concentrations.     

Mutual benefit interactions between banded pine weevil Pissodes castaneus and blue‐stain fungus Leptographium serpens in maritime pine

• 1 Ecological interactions between banded pine weevil Pissodes castaneus and blue‐stain fungus Leptographium serpens, when simultaneously sharing the same host plant (maritime pine Pinus pinaster) in winter and spring, were investigated. Temporal components of the interaction were taken into account by either introducing the weevils and the pathogen simultaneously or sequentially, with the weevils being introduced 1 month after the fungal inoculation.
• 2 We measured larval mortality, development time, offspring number, sex ratio and body size of P. castaneus. Phloem phosphorus and nitrogen concentrations were also assessed. Furthermore, we tested whether: (i) emerging offspring transported propagules of the fungus; (ii) artificially‐contaminated weevils may transmit the disease to healthy trees; and (iii) field collected P. castaneus carry the fungus.
• 3 The fungus enhanced weevil colonization and brood production in both seasons. During winter and spring, adults from trees where the pathogen was inoculated prior to weevil introduction emerged earlier than weevils from trees where they had been introduced simultaneously with the fungus. During winter, weevils from pre‐inoculated trees were also larger. Sex ratio and larval mortality were not affected. Leptographium serpens did not affect phloem nitrogen content but phosphorus content was greater in plants inoculated with the pathogen, which may explain the findings on weevil growth.
• 4 Sixty‐five percent of the weevils that emerged from inoculated trees carried spores of L. serpens, although no successful isolation was made from field collected weevils. The fungus was recovered from 25% of the trees infested with artificially‐contaminated weevils.
• 5 These results suggest that P. castaneus benefits from the presence of L. serpens and may contribute to its spread.

     

Concentrations of Trace Metals in Dominant Aquatic Plants of the Lake Provala (Vojvodina, Yugoslavia)

The trace metal (Fe, Mn, Zn, Cu, Ni, Pb, Cd, Sr, and Cr) contents in the most common submerged and floating aquatic plants Ceratophyllum demersum L., Myriophyllum spicatum L., and Nymphoides flava Hill. of Provala Lake were evaluated. Considerable higher contents of iron, manganese, zinc, nickel, lead and strontium were found in submerged species than in the floating ones. The presence of cadmium and lead in plant tissues points to a certain degree of lake water pollution.     

Variation in hostplant preference and performance by the milfoil weevil, Euhrychiopsis lecontei Dietz, exposed to native and exotic watermilfoils

Exotic plants often displace native plants and thus alter the availability of native hostplants for specialist herbivorous insects. The submersed aquatic weevil Euhrychiopsis lecontei Dietz is endemic to North America, but there are now source populations on the exotic Eurasian watermilfoil (Myriophyllum spicatum L.) as well as on the weevil's ancestral host, northern watermilfoil (Myriophyllum sibiricum Komarov). This provides an opportunity to examine a host range expansion in progress. To further define the host range of the weevil and to determine how population source and rearing plant influence host plant preference and performance, we conducted a series of preference and rearing experiments with weevils from two source populations reared on northern milfoil, on Eurasian milfoil, switched late in larval development from northern to Eurasian milfoil, and vice versa. We also included two rearing treatments with milfoils on which the weevil has not been documented: the native M. verticillatum L. and the exotic M. aquaticum Verd. Preference by weevils in the switched rearing treatments was similar to preferences exhibited by weevils reared solely on the second (later) milfoil species and an increase in preference for Eurasian milfoil was induced by adult exposure to Eurasian milfoil for 2 weeks. In contrast, sizes and development times of weevils in the switched rearing treatments were similar to sizes and development times exhibited by weevils reared solely on the first (early) milfoil species. These results indicate that preference by the milfoil weevil is determined late in larval development or later and Hopkins' host selection principle is not supported. However, size and development time were most affected by hostplant quality during larval development when larvae must acquire the resources needed for pupation. Oviposition preference in the milfoil weevil was a population attribute, not a fixed individual attribute and there was no significant variation in preference among individuals reared on northern milfoil, but significant variation in preference was detected among weevils reared on Eurasian milfoil. Weevils oviposited on all four milfoil species and completed development on three of them, but did not develop beyond the larval stage on M. aquaticum. Weevils reared on Eurasian milfoil developed faster and reached larger adult sizes than weevils in any other rearing treatment. The smallest sizes and longest development times were for weevils reared on the natives, northern milfoil and M. verticillatum. The milfoil weevil oviposits on an array of milfoil species and is unable to distinguish an unsuitable host (M. aquaticum) within this genus. The influence of rearing plant and adult exposure to Eurasian milfoil on hostplant preference suggests that host range expansion to novel congeners may occur more rapidly than predicted by models which assume that genetic variation is required. Significant variation among individuals in hostplant preference suggests the potential for a host shift to a plant for which E. lecontei appears pre-adapted.     

Leaf scarring by the weevils Neochetina eichhorniae and N. bruchi enhances infection by the fungus Cercospora piaropi on waterhyacinth, Eichhornia crassipes

Additive or synergistic effects among introduced and native insect and plant pathogen agents are necessary to achieve biological control of waterhyacinth (Eichhornia crassipes), a globally damaging aquatic weed. In field plots, plants were infested with waterhyacinth weevils (Neoechetina bruchi and N. eichhorniae) and leaves were scarred by weevil feeding. Subsequent infection by the fungal pathogen Cercospora piaropi caused necrotic lesions to form on leaves. Necrosis development was 7.5- and 10.5-fold greater in plots augmented with both weevils and C. piaropi and weevils alone, respectively, than in plots receiving only C. piaropi. Twenty-four days after weevil infestation, the percentage of laminar area covered by lesions on third-youngest and oldest live leaves was elevated 2.3–2.5-fold in plots augmented with weevils. Scar density and necrosis coverage on young leaf laminae were positively correlated, even though antipathogenic soluble peroxidases were elevated 3-fold in plots augmented with weevils alone or weevils and C. piaropi. Combined weevil and fungal augmentation decreased shoot densities and leaves per plant. In a no-choice bioassay, weevil feeding on oldest but not young leaves was reduced 44 two weeks after C. piaropi inoculation. Protein content and peroxidase activities were elevated 2–6-fold in oldest leaves three weeks after inoculation. Augmentation with both waterhyacinth weevils and C. piaropi led to the development of an additive biological control impact, mediated by one or more direct interactions between these agents, and not plant quality effects.     

Epiphytic bacterial community composition on two common submerged macrophytes in brackish water and freshwater

Background  

Plants and their heterotrophic bacterial biofilm communities possibly strongly interact, especially in aquatic systems. We aimed to ascertain whether different macrophytes or their habitats determine bacterial community composition. We compared the composition of epiphytic bacteria on two common aquatic macrophytes, the macroalga Chara aspera Willd. and the angiosperm Myriophyllum spicatum L., in two habitats, freshwater (Lake Constance) and brackish water (Schaproder Bodden), using fluorescence in situ hybridization. The bacterial community composition was analysed based on habitat, plant species, and plant part.     

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

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