Vision is Important for Plant Location by the Phytophagous Aquatic Specialist Euhrychiopsis lecontei Dietz (Coleoptera: Curculionidae)

The aquatic milfoil weevil Euhrychiopsis lecontei Dietz (Coleoptera: Curculionidae) is a specialist on Myriophyllum spp. and is used as a biological control agent for Eurasian watermilfoil (Myriophyllum spicatum L.), an invasive aquatic macrophyte. We show evidence that visual cues are important for plant detection by these weevils. Weevils had difficulty locating plants in dark conditions and were highly attracted to plant stems in the light, even when the plant sample was sealed in a vial. However, weevils were equally attracted to both M. spicatum and another aquatic macrophyte, coontail (Ceratophyllum demersum L.) in vials. Turbidity (0–100 NTU) did not significantly influence visual plant detection by the weevils. This work fills a void in the literature regarding visual plant location by aquatic specialists and may help lead to a better understanding of when and where these weevils will find, accept, and damage their target host-plants.     

Modeling the interaction between the exotic invasive aquatic macrophyte <Emphasis Type="Italic">Myriophyllum spicatum</Emphasis> and the native biocontrol agent <Emphasis Type="Italic">Euhrychiopsis lecontei</Emphasis> to improve augmented management programs

Mathematical modeling is used to investigate control of the invasive aquatic plant Myriophyllum spicatum L. by the native watermilfoil weevil Euhrychiopsis lecontei Dietz. An age-structured population model with discrete development class is used to separate the destructive activity of Euhrychiopsis lecontei Dietz larvae from the relatively benign activity of the other developmental stages. Empirical model behavior is evaluated and agrees with expected system characteristics. Simulation results demonstrate diminishing returns in the control impact of E. lecontei Dietz as a function of population stocking density. These results indicate that predictive modeling of this type may be used to optimize the utilization of Euhrychiopsis lecontei Dietz as a biological control agent.     

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.     

Induced responses to grazing by an insect herbivore (Acentria ephemerella) in an immature macrophyte (Myriophyllum spicatum): an isotopic study

While the mechanisms by which adult terrestrial plants deploy constitutive and induced responses to grazing pressure are well known, the means by which young aquatic plants defend themselves from herbivory are little studied. This study addresses nitrogen transport in the aquatic angiosperm Myriophyllum spicatum in response to herbivore exposure. Nitrogen tracers were used to monitor nitrogen uptake and reallocation in young plants in response to grazing by the generalist insect herbivore Acentria ephemerella. Total nitrogen content (N%) and patterns of nitrogen uptake and allocation (δ¹⁵N) were assessed in various plant tissues after 24 and 48 h. Following 24 h exposure to herbivore damage (Experiment 1), nitrogen content of plant apices was significantly elevated. This rapid early reaction may be an adaptation allowing the grazer to be sated as fast as possible, or indicate the accumulation of nitrogenous defense chemicals. After 48 h (Experiment 2), plants' tips showed depletion in nitrogen levels of ca. 60‰ in stem sections vulnerable to grazing. In addition, nitrogen uptake by grazed and grazing‐prone upper plant parts was reduced and nutrient allocation into the relatively secure lower parts increased. The results point to three conclusions: (1) exposure to an insect herbivore induces a similar response in immature M. spicatum as previously observed in mature terrestrial species, namely a rapid (within 48 h) reduction in the nutritional value (N%) of vulnerable tissues, (2) high grazing intensity (100% of growing tips affected) did not limit the ability of young plants to induce resistance; and (3) young plants exposed to herbivory exhibit different patterns of nutrient allocation in vulnerable and secure tissues. These results provide evidence of induced defense and resource reallocation in immature aquatic macrophytes which is in line with the responses shown for mature aquatic macrophytes and terrestrial plants.     

Population genetic structure in six sympatric and widespread aquatic plants inhabiting diverse lake environments in China

Many aquatic plant species are distributed over large areas and diverse environments with populations interconnected by abiotic and biotic mediators. Here, we examined differences and similarities in the population genetic structure of six sympatric and widespread aquatic plant species. We sampled the aquatic species from six Chinese lakes found on plateaus, plains, and different river systems and analyzed them using inter‐simple sequence repeat (ISSR) markers. Samples originating from each lake tended to cluster together. Of the six species, only Nymphoides peltata and Myriophyllum spicatum could be divided into plateau and plain groups, once Taihu Lake individuals were excluded. Genetic similarities between populations connected by the Yangtze River were not consistently higher than unconnected populations. Populations from Taihu Lake and/or Weishanhu Lake were distant from other lake populations for all species except Potamogeton lucens. The Taihu and Weishanhu populations clustered for Ceratophyllum demersum and Typha latifolia. Hydrophilous C. demersum had the lowest gene flow (Nm = 0.913), whereas the entomophilous Hydrocharis dubia (Nm = 2.084) and N. peltata (Nm = 2.204) had the highest gene flow. The genetic relationships among distant populations of aquatic plants reflect the comprehensive effects of environmental selection pressure and biotic and abiotic connectivity. Differences in environmental factors between plateau and plain lakes and long distance hydrochory have limited importance on aquatic plant genetic structures. Among multiple evolutionary forces, gene flow mediated by birds may play the most important role in the formation of genetic patterns in the six species examined. For example, the close genetic relationship between Taihu Lake and Weishanhu Lake populations, each in different river systems and with different climates, may be related to the migration routes of birds. Differences in gene flow among the six aquatic plants may be attributable to different bird‐transport and the fruit traits of each species.     

Evaluation of the predatory mite <Emphasis Type="Italic">Phytoseiulus macropilis</Emphasis> (Acari: Phytoseiidae) as a biological control agent of the two-spotted spider mite on strawberry plants under greenhouse conditions

The predatory mite Phytoseiulus macropilis is a potential biological control agent of the two-spotted spider mite (TSSM) Tetranychus urticae on strawberry plants. Its ability to control TSSM was recently assessed under laboratory conditions, but its ability to locate and control TSSM under greenhouse conditions has not been tested so far. We evaluated whether P. macropilis is able to control TSSM on strawberry plants and to locate strawberry plants infested with TSSM under greenhouse conditions. Additionally, we tested, in an olfactometer, whether odours play a role in prey-finding by P. macropilis. The predatory mite P. macropilis required about 20 days to achive reduction of the TSSM population on strawberry plants initially infested with 100 TSSM females per plant. TSSM-infested plants attract an average of 27.5 ± 1.0% of the predators recaptured per plant and uninfested plants attracted only 5.8 ± 1.0% per plant. The predatory mites were able to suppress TSSM populations on a single strawberry plant and were able to use odours from TSSM-infested strawberry plants to locate prey in both olfactometer and arena experiments. Hence, it is concluded that P. macropilis can locate and reduce TSSM population on strawberry plants under greenhouse conditions.     

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.     

Effects of an artificial blend of host‐infested plant volatiles on plant attractiveness to parasitic wasps

For the biological control of diamondback moth (DBM) larvae in commercial greenhouses, we have previously identified a blend of volatiles that attracted Cotesia vestalis, a parasitoid of DBM larvae. Here, we tested the effects of an artificial volatile blend on the attractiveness of komatsuna plants (Japanese mustard spinach; Brassica rapa var. perviridis) to C. vestalis under greenhouse conditions. First, we showed that female C. vestalis preferred infested komatsuna plants to uninfested plants in the greenhouse. Under the same conditions, placing the artificial attractants near both infested and uninfested plants did not affect the wasps’ preference. However, when comparing infested komatsuna plants coupled with the artificial attractants with infested plants without them, significantly more female C. vestalis were attracted to the former. The possible use of artificial C. vestalis attractants for the biological control of DBM is discussed.     

Orientation behavior of Propylaea japonica toward visual and olfactory cues from its prey–host plant combination

The lady beetle Propylaea japonica (Thunberg) (Coleoptera: Coccinellidae) is an important predator of aphids in agroecosystems. The inundative release of coccinellid beetles can be an effective biological control strategy. An understanding of how biological control agents perceive and use stimuli from host plants is the key to successfully implement commercially produced predators. Here, we studied the relative role of visual and volatile cues. Dual‐choice assays using foraging‐naïve and foraging‐experienced P. japonica adults were conducted using cotton plants [Gossypium hirsutum L. (Malvaceae)] with or without infestation by the cotton aphid, Aphis gossypii (Glover) (Hemiptera: Aphididae). Overall, experienced beetles were more attracted than naïve beetles toward cues associated with aphid‐infested plants. Experienced beetles were also more responsive to olfactory cues compared with naïve beetles. Both foraging‐naïve and ‐experienced lady beetles integrate olfactory and visual cues from plants infested with aphids, with an apparently greater reliance on olfactory cues. The results suggest that foraging experience may increase prey location in P. japonica.     

Comparison of the Adsorption Capabilities of Myriophyllum spicatum and Ceratophyllum demersum for Zinc,Copper and Lead

Industrial wastewaters contain various heavy metal components and therefore threaten aquatic bodies. Heavy metals can be adsorbed by living or non‐living biomass. Submerged aquatic plants can be used for the removal of heavy metals. This paper exhibits the comparison of the adsorption properties of two aquatic plants Myriophyllum spicatum and Ceratophyllum demersum for lead, zinc, and copper. The data obtained from batch studies conformed well to the Langmuir Model. Maximum adsorption capacities (q_max) were obtained for both plant species and each metal. The maximum adsorption capacities (q_max) achieved with M. spicatum were 10.37 mg/g for Cu²⁺, and 15.59 mg/g for Zn²⁺ as well as 46.49 mg/g for Pb²⁺ and with C. demersum they were 6.17 mg/g for Cu²⁺, 13.98 mg/g for Zn²⁺ and 44.8 mg/g for Pb²⁺. It was found that M. spicatum has a better adsorption capacity than C. demersum for each metal tested. Gibbs free energy and the specific surface area based on the q_max values were also determined for each metal.     

Sorption of Co, Cu, Ni and Zn from industrial effluents by the submerged aquatic macrophyte Myriophyllum spicatum L.

The submerged aquatic plant Myriophyllum spicatum L. (Eurasian water milfoil) has been suggested as an efficient plant species for the treatment of metal-contaminated industrial wastewater. The process of metal removal by plants involves a combination of rapid sorption on the surface and slow accumulation and translocation in the biomass. This study focussed on the sorption/desorption characteristics of the surface of M. spicatum for Co, Cu, Ni and Zn. Batch sorption tests with mixed metal solutions covering a range of 0, 1, 5, 10, 50 and 100 mg l⁻¹ of each metal, were performed. For Co, Ni and Zn, the sorption process was well described by the Langmuir model, whereas sorption of Cu was better described by the Freundlich model. The biomass showed the highest affinity for Cu and Zn. Langmuir sorption maxima of Co, Ni and Zn were 2.3, 3.0 and 6.8 mg g⁻¹ DM, respectively. At the highest initial concentration of 100 mg l⁻¹, a maximum of 29 mg g⁻¹ DM of Cu was sorbed onto the surface of the biomass. Desorption by 0.1 M HCl did not fully recover the metals sorbed onto the surface and there was evidence of leaching from within the biomass. Recovery of heavy metals and regeneration of the biomass by washing with 0.1 M HCl was therefore not suggested as a viable strategy.     

Effects of invasive species on plant communities: an example using submersed aquatic plants at the regional scale

Submersed aquatic plants have a key role in maintaining functioning aquatic ecosystems through their effects on the hydrological regime, sedimentation, nutrient cycling and habitat of associated fauna. Modifications of aquatic plant communities, for example through the introduction of invasive species, can alter these functions. In the Sacramento-San Joaquin River Delta, California, a major invasive submersed plant, Brazilian waterweed Egeria densa, has become widespread and greatly affected the functionality of the submersed aquatic plant community. Rapid assessments of the distribution and abundance of this species are therefore crucial to direct management actions early in the season. Given the E. densa bimodal growth pattern (late spring and fall growth peaks), summer assessments of this species may indicate which and where other submersed species may occur and fall assessments may indicate where this and other species may occur in the following spring, primarily because the Delta’s winter water temperatures are usually insufficient to kill submersed aquatic plant species. We assessed community composition and distribution in the fall of 2007 and summer of 2008 using geostatistical analysis; and measured summer biomass, temperature, pH, salinity, and turbidity. In the fall of 2007, submersed aquatic plants covered a much higher proportion of the waterways (60.7%) than in the summer of 2008 (37.4%), with a significant overlap between the seasonal distribution of native and non-native species. Most patches were monospecific, and multispecies patches had significantly higher dominance by E. densa, co-occurring especially with Ceratophyllum demersum. As species richness of non-natives increased there was a significant decrease in richness of natives, and of native biomass. Sustained E. densa summer biomass negatively affected the likelihood of presence of Myriophyllum spicatum, Potamogeton crispus, and Elodea canadensis but not their biomass within patches. Depth, temperature and salinity were associated with biomass; however, the direction of the effect was species specific. Our results suggest that despite native and invasive non-native submersed plant species sharing available niches in the Delta, E. densa affects aquatic plant community structure and composition by facilitating persistence of some species and reducing the likelihood of establishment of other species. Successful management of this species may therefore facilitate shifts in existing non-native or native plant species.     

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.     

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)     

Effects of Light,Temperature, and Water Depth on Growth of a Rare Aquatic Plant, <Emphasis Type="Italic">Ranunculus kadzusensis</Emphasis>

Ranunculus kadzusensis is an endangered aquatic plant species that commonly reproduces in the rice paddies of Korea and Japan during winter and early spring. Here, we investigated the effects of main aquatic environmental factors—light, temperature, and water depth—on its growth, with the goal of seeking information that will contribute to its in situ conservation. As the amount of shading increased, biomass, maximum shoot length, number of branches, flowers, and fruits, main stem diameter, and maximum leaf length decreased. Although seed germination occurred under a 12-h photoperiod and at either 30/20°C (day/night) or 20/15°C, most plants died at the higher temperature. Survival was 0% for surface-grown plant sets when tested in a wintertime pond experiment. The rate of maximum shoot extension was greatest for plants grown at depths of 50 and 100 cm versus those at 20 cm. Thus, we demonstrated that R. kadzusensis is intolerant of high temperatures and shade, which may explain why its growth is limited to paddies with no shading and where temperatures are low early in the year, before rice cultivation begins.     

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.     

<Emphasis Type="Italic">Mayaca fluviatilis</Emphasis> Aubl.: an ornamental aquatic with invasive potential in Sri Lanka

The ornamental aquatic plant industry in Sri Lanka has grown substantially during the last decade and there is a renewed demand for several exotic species. Mayaca fluviatilis, a submerged plant used in aquaria, is not recorded as a naturally occurring plant in Sri Lanka. However, recent studies recorded it in the wet zone of Sri Lanka and certain countries have recognized it as a problematic plant. The knowledge of the mode of propagation of invasive species is vital in the management of biological invasions. Therefore, the present study was conducted to investigate the propagation ability of vegetative parts of M. fluviatilis. A field survey was conducted in the wet zone of Sri Lanka, and populations of M. fluviatilis were monitored between 2006 and 2009. The propagation ability of different maturity stages of stem cuttings (terminal, middle and basal) were investigated, and terminal stem cuttings with five different lengths (2, 4, 6, 8 and 10 cm) were used to investigate the propagation ability of fragmented vegetative parts. Terminal shoots showed the highest survival rates and significantly high (P < 0.05) shoot and root lengths compared to middle and basal parts of the stem cuttings. In the other experiment with terminal stem cuttings with different lengths, all the stem fragments were capable of developing into a new plant. Increasing lengths of fragments was significantly related to the number of roots. Nevertheless, as even the 2-cm fragment was capable of developing into a new plant, removal of even the smallest fragments is necessary to control this plant. Submerged plants can spread easily through fragmentation and manual removal will likely increase its spread. Nevertheless, it is considered as effective for small infestations. Therefore, repeated removal of plant parts can be suggested as an ecologically sound management approach to prevent further spread of M. fluviatilis since it is in the early stage of establishment.     

Chlorophyll fluorometry sheds light on the role of desiccation resistance for vegetative overland dispersal of aquatic plants

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Go with the flow: Fragment retention patterns shape the vegetative dispersal of aquatic plants in lowland streams

1. The dispersal of aquatic plant propagules is highly facilitated in streams due to flow. As many aquatic plants predominantly spread through vegetative propagules, the specific retention and thus drift distance of dispersed plant fragments largely contribute to the rapid spread along the course of a stream.
2. We determined fragment retention for four aquatic plant species (Elodea canadensis, Myriophyllum spicatum, Ceratophyllum demersum, Salvinia natans; representing four different common morpho-structural groups) in sections of small to medium-sized German streams with different levels of stream sinuosity.
3. The number of fragments showed a logistic decline over drift distance. In two small streams, 90% of drifting fragments were retained at distances (D₉₀) of only 5–9 m and 19–70 m, while higher D₉₀ values of 116–903 m and 153–2,367 m were determined for sections of a medium-sized stream. The likelihood of retention thereby decreased significantly with increasing stream size and was reduced in straightened stream sections.
4. Differences in retention were more strongly related to fragment buoyancy rather than fragment size and morphology. Increasing buoyancy significantly lowered the likelihood of fragment retention over drift distance by a factor of 3–8, whereas contrasting effects were documented for size and morphology of fragments.
5. The relevance of different obstacles was highly stream section-specific and depended on obstacle abundance, distribution, and the degree of submergence/emergence.
6. Our findings elucidate the dynamic retention patterns of plant fragments and highlight the strong interplay between extrinsic (stream) and intrinsic (fragment) properties. We conclude that straightened lowland streams of intermediate size promote the rapid dispersal of invasive aquatic plants and are particularly prone to invaders producing large amounts of small and highly buoyant plant fragments. Information on the species-specific fragment colonisation dynamics in the field is further required to improve our understanding of the vegetative dispersal capacity of invasive aquatic plants in stream ecosystems.