Causes of macrophyte mass development and management recommendations.

Aquatic plants (macrophytes) are important for ecosystem structure and function. Macrophyte mass developments are, however, often perceived as a nuisance and are commonly managed by mechanical removal. This is costly and often ineffective due to macrophyte regrowth. There is insufficient understanding about what causes macrophyte mass development, what people who use water bodies consider to be a nuisance, or the potential negative effects of macrophyte removal on the structure and function of ecosystems. To address these gaps, we performed a standardized set of in situ experiments and questionnaires at six sites (lakes, reservoirs, and rivers) on three continents where macrophyte mass developments occur. We then derived monetary values of ecosystem services for different scenarios of macrophyte management ("do nothing", "current practice", "maximum removal"), and developed a decision support system for the management of water courses experiencing macrophyte mass developments. We found that (a) macrophyte mass developments often occur in ecosystems which (unintentionally) became perfect habitats for aquatic plants, that (b) reduced ecosystem disturbance can cause macrophyte mass developments even if nutrient concentrations are low, that (c) macrophyte mass developments are indeed perceived negatively, but visitors tend to regard them as less of a nuisance than residents do, that (d) macrophyte removal lowers the water level of streams and adjacent groundwater, but this may have positive or negative overall societal effects, and that (e) the effects of macrophyte removal on water quality, greenhouse gas emissions, and biodiversity vary, and likely depend on ecosystem characteristics and macrophyte life form. Overall, we found that aquatic plant management often does not greatly affect the overall societal value of the ecosystem, and we suggest that the "do nothing" option should not be easily discarded in the management of perceived nuisance mass developments of aquatic plants.

Impact of Mechanical Removal on the Regeneration and Colonization Abilities of the Alien Aquatic Macrophyte Egeria densa.

The development of aquatic plant beds can obstruct boat traffic, hinder the practice of water activities, and impact the functioning of freshwaters. In order to mitigate their effects, mechanical removal is often the preferred management solution. The objective of this study was to test, in mesocosms, the effect of frequency (none, one, and two cuts) and cutting dates (May and/or July) on the regeneration and colonization capabilities of the aquatic plant Egeria densa, an invasive alien species in France. The cutting date had no effect on the capabilities of E. densa, but the two cuts significantly reduced the plant's biomass. Removal produced numerous fragments, which exhibited very high survival and anchoring rates. However, summer removal produced fragments with lower regeneration and colonization abilities compared to fragments from spring cutting. Mechanical removal only temporarily reduced the biomass of the aquatic plant beds and could promote the formation of new beds from the fragments generated by management and dispersed by water flow.

Responses of three invasive alien aquatic plant species to climate warming and plant density.

Climate warming may impact plant invasion success directly, as well as indirectly through changes among interactions within plant communities. However, the responses of invasive alien aquatic species to plant density and rising temperatures remain largely unknown. We tested the effects of plant density and neighbour plant identity at different temperatures to better understand the performance of a community of invasive species exposed to climate warming. A microcosm experiment was conducted with three invasive aquatic plants species-Elodea canadensis, Egeria densa and Lagarosiphon major-, at mono and polycultures with low and high plant density, at 16 °C, 19 °C and 23 °C. The results clearly demonstrated that rising temperature influenced, either as a single parameter or as a combined factor, at least one of the measured traits of the three invasive species. Leaf area of E. densa, root number of L. major and growth of E. densa and L. major were influenced by temperature, plant density and neighbour identity. Plant density influenced all traits with the exception of leaf area of E. canadensis and lateral branch production of E. densa. Neighbour identity had no effect on growth rate and leaf area of E. canadensis, on lateral branch and roots production of E. densa and on leaf area of L. major. These findings establish that rising temperature could enhance competition or facilitation among E. canadensis, L. major and E. densa and could cancel the beneficial effects of the presence of a neighbour species; however, the magnitude of this effect was strongly dependent on plant density. Rising temperature due to climate change will likely play a crucial role in interactions between invasive species within plant communities and in the further spread of these invasive aquatic plants.

Drivers of Perceived Nuisance Growth by Aquatic Plants.

Mass developments of macrophytes occur frequently worldwide and are often considered a nuisance when interfering with human activities. It is crucial to understand the drivers of this perception if we are to develop effective management strategies for ecosystems with macrophyte mass developments. Using a comprehensive survey spanning five sites with different macrophyte species in four countries (Norway, France, Germany and South Africa), we quantified the perception of macrophyte growth as a nuisance among residents and visitors, and for different recreational activities (swimming, boating, angling, appreciation of biodiversity, appreciation of landscape and birdwatching). We then used a Bayesian network approach to integrate the perception of nuisance with the consequences of plant removal. From the 1234 responses collected from the five sites, a range of 73-93% of the respondents across the sites considered macrophyte growth a nuisance at each site. Residents perceived macrophytes up to 23% more problematic than visitors. Environmental mindedness of respondents did not influence the perception of nuisance. Perceived nuisance of macrophytes was relatively similar for different recreational activities that were possible in each case study site, although we found some site-specific variation. Finally, we illustrate how Bayesian networks can be used to choose the best management option by balancing people's perception of macrophyte growth with the potential consequences of macrophyte removal.

Short-term effects of macrophyte removal on aquatic biodiversity in rivers and lakes.

Mass development of macrophytes is an increasing problem in many aquatic systems worldwide. Dense mats of macrophytes can negatively affect activities like boating, fishing or hydropower production and one of the management measures often applied is mechanical removal. In this study, we analyzed the effect of mechanical macrophyte removal on phytoplankton, zooplankton, and macroinvertebrate (pelagic and benthic samples) assemblages. Our study covered five sites in four countries in Europe and Africa with highly variable characteristics. In all sites, dense mats of different macrophyte species (Juncus bulbosus in a river in Norway; a mix of native macrophytes in a German river, Elodea nuttallii in a lake in Germany, Ludwigia spp. In a French lake and Pontederia crassipes in a South African lake) are problematic and mechanical removal was applied. In every country, we repeated the same BACI (Before-After-Control-Impact) design, including "before", "one week after", and "six weeks after" sampling in a control and an impact section. Repeating the same experimental design at all sites allowed us to disentangle common effects across all sites from site-specific effects. For each taxonomic group, we analyzed three structural and three functional parameters, which we combined in a scoring system. Overall, the removal of macrophytes negatively affected biodiversity, in particular of zooplankton and macroinvertebrate assemblages. In contrast, plant removal had positive effects on the phytoplankton assemblages. Effects were more pronounced one week after removal than six weeks after. Consequently, we suggest a stronger consideration of the effect of plant removal on biodiversity to arrive at more sustainable management practices in the future.

Advancing biological invasion hypothesis testing using functional diversity indices.

Pioneering investigations on the effects of introduced populations on community structure, ecosystem functioning and services have focused on the effects of invaders on taxonomic diversity. However, taxonomic-based diversity metrics overlook the heterogeneity of species roles within and among communities. As the homogenizing effects of biological invasions on community and ecosystem processes can be subtle, they may require the use of functional diversity indices to be properly evidenced. Starting from the listing of major functional diversity indices, alongside the presentation of their strengths and limitations, we focus on studies pertaining to the effects of invasive species on native communities and recipient ecosystems using functional diversity indices. By doing so, we reveal that functional diversity of the recipient community may strongly vary at the onset of the invasion process, while it stabilizes at intermediate and high levels of invasion. As functional changes occurring during the lag phase of an invasion have been poorly investigated, we show that it is still unknown whether there are consistent changes in functional diversity metrics that could indicate the end of the lag phase. Thus, we recommend providing information on the invasion stage under consideration when computing functional diversity metrics. For the existing literature, it is also surprising that very few studies explored the functional difference between organisms from the recipient communities and invaders of the same trophic levels, or assessed the effects of non-native organism establishment into a non-analogue versus an analogue community. By providing valuable tools for obtaining in-depth diagnostics of community structure and functioning, functional diversity indices can be applied for timely implementation of restoration plans and improved conservation strategies. To conclude, our work provides a first synthetic guide for their use in hypothesis testing in invasion biology.

Heterogenous Impact of Water Warming on Exotic and Native Submerged and Emergent Plants in Outdoor Mesocosms.

Some aquatic plants present high biomass production with serious consequences on ecosystem functioning. Such mass development can be favored by environmental factors. Temperature increases are expected to modify individual species responses that could shape future communities. We explored the impact of rising water temperature on the growth, phenology, and metabolism of six macrophytes belonging to two biogeographic origins (exotic, native) and two growth forms (submerged, emergent). From June to October, they were exposed to ambient temperatures and a 3 °C warming in outdoor mesocosms. Percent cover and canopy height were favored by warmer water for the exotic emergent Ludwigia hexapetala. Warming did not modify total final biomass for any of the species but led to a decrease in total soluble sugars for all, possibly indicating changes in carbon allocation. Three emergent species presented lower flavonol and anthocyanin contents under increased temperatures, suggesting lower investment in defense mechanisms and mitigation of the stress generated by autumn temperatures. Finally, the 3 °C warming extended and shortened flowering period for L. hexapetala and Myosotis scorpioides, respectively. The changes generated by increased temperature in outdoor conditions were heterogenous and varied depending on species but not on species biogeographic origin or growth form. Results suggest that climate warming could favor the invasiveness of L. hexapetala and impact the structure and composition of aquatic plants communities.

Seed source regions drive fitness differences in invasive macrophytes.

PREMISE: Worldwide, ecosystems are threatened by global changes, including biological invasions. Invasive species arriving in novel environments experience new climatic conditions that can affect their successful establishment. Determining the response of functional traits and fitness components of invasive populations from contrasting environments can provide a useful framework to assess species responses to climate change and the variability of these responses among source populations. Much research on macrophytes has focused on establishment from clonal fragments; however, colonization from sexual propagules has rarely been studied. Our objective was to compare trait responses of plants generated from sexual propagules sourced from three climatic regions but grown under common environmental conditions, using L. peploides subsp. montevidensis as a model taxon. METHODS: We grew seedlings to reproductive stage in experimental mesocosms under a mediterranean California (MCA) climate from seeds collected in oceanic France (OFR), mediterranean France (MFR), and MCA. RESULTS: Seed source region was a major factor influencing differences among invasive plants recruiting from sexual propagules of L. peploides subsp. montevidensis. Trait responses of young individual recruits from MCA and OFR, sourced from geographically distant and climatically distinct source regions, were the most different. The MCA individuals accumulated more biomass, flowered earlier, and had higher leaf N concentrations than the OFR plants. Those from MFR had intermediate profiles. CONCLUSIONS: By showing that the closer a seedling is from its parental climate, the better it performs, this study provides new insights to the understanding of colonization of invasive plant species and informs its management under novel and changing environmental conditions.

Allelopathic Effects of Native Versus Invasive Plants on One Major Invader.

Allelopathy is defined as the effects (stimulatory and inhibitory) of a plant on the development of neighboring plants through the release of secondary compounds. Autoallelophaty is the beneficial or harmful effect of a plant species on itself. The allelopathic potential belonging to a native species could induce a biotic resistance against invasive plants, whereas allelochemicals released by exotic species could favor the establishment of invasive species (invasional meltdown). The aim of our study was to examine the potential allelopathic effect of four plant species on the target species Ludwigia hexapetala using two experiments. In the first experiment, we tested the allelopathic effect of root and leaf leachates of the two congeneric exotic species Ludwigia hexapetala and Ludwigia peploides on L. hexapetala, while in the second experiment, we studied the allelopathic effect of root and leaf leachates of a sympatric exotic species Myriophyllum aquaticum and of one native species Mentha aquatica on L. hexapetala. We measured the stem length to calculate the relative growth rate and four physiological traits (nitrogen balance index and flavonol, chorophyll, anthocyanin indices) of the target plants on a weekly basis. At the end of the experiment, we determined the aboveground and belowground biomass. We also counted the number of lateral branches and measured their lengths. We found that the root leachates of L. peploides and of Myriophyllum aquaticum had stimulated the synthesis of flavonols of L. hexapetala. Leaf leachate of L. hexapetala also stimulated its own flavonol synthesis. Also, the root leachate of L. peploides had stimulated the total biomass and length of lateral branches of L. hexapetala, whereas the production of lateral branches had been stimulated by root leachates of both Ludwigia species and by leaf leachate of Myriophyllum aquaticum. The autoallelopathy of L. hexapetala could explain its invasiveness. Both leachates produced by Mentha aquatica had no effect on the physiological and morphological traits of the invasive L. hexapetala and indicated no biotic resistance in the recipient community. The two invasive plant species Myriophyllum aquaticum and L. peploides could favor the establishment of L. hexapetala. These results suggested an "invasional meltdown."

Allelopathic Effect of the Invasive Ludwigia hexapetala on Growth of Three Macrophyte Species.

The release of allelochemicals by plants can affect the performance of other organisms positively or negatively. We tested the effects of aqueous extracts and leachates derived from the leaves and roots of the invasive water primrose (Ludwigia hexapetala) on one submerged native species - Ceratophyllum demersum, and two exotic species - the submerged Egeria densa and the emergent growth form of Myriophyllum aquaticum. The effect of the aqueous extracts and leachates of L. hexapetala on photosynthetic yield, growth (i.e., relative growth rate, leaf area), root length, and length of the lateral shoots of each species were analyzed in spring and in autumn. In autumn, an allelopathic effect was established on the traits of the three macrophytes species. The root extracts stimulated leaf area and the photosynthetic yield of C. demersum and of E. densa, whereas leaf treatments (leachates and extracts) and root leachate reduced the leaf area of M. aquaticum. The autumnal root leachate of L. hexapetala decreased the relative growth rate of C. demersum, whereas it had no effect on the two others plants. The root extract increased the length of lateral branches of M. aquaticum in autumn, suggesting a positive effect of L. hexapetala on the lateral growth of M. aquaticum. Three main allelochemicals were identified in leaves: quercitrin, prunin, myricitrin. The concentrations of these allelochemicals were greater in the leaf extract taken from L. hexapetala in autumn than in spring, and those found in the leaf leachates for both seasons. This assessment of autumnal allelopathy could help to explain the patterns of plant community succession in invaded areas.

Response of Macrophyte Traits to Herbivory and Neighboring Species: Integration of the Functional Trait Framework in the Context of Ecological Invasions.

With the increase in the number of introduced species each year, biological invasions are considered as one of the most important environmental problems for native biodiversity. In invaded habitats, the establishment of exotic plant species depends on the abiotic and biotic environment. Herbivores and neighboring plants (native or exotic) comprise an important part of the latter. Herbivores cause trophic and non-trophic damage to focal plants, which respond to herbivory by varying their different traits quantitatively (e.g., growth rate and biomass changes) and qualitatively (e.g., variation in morphological and chemical defenses strategies affecting plant palatability). Neighboring plant species also affect functional traits and the fitness of focal plant species, thus herbivore effects on a focal plant could also depend indirectly on the palatability and defensive traits of the neighboring species inside the community. Here, in a first step toward the integration of associational susceptibility/resistance theories in the field of ecological invasion, we performed a microcosm experiment to consider the effects of an exotic crayfish on the growth rate, morphological traits and damage level of three macrophytes (two exotic, one native) growing in pairwise combinations. We found that (i) the response to herbivore presence and to neighboring species identity seemed to be species specific, and (ii) crayfish enhance the fragmentation rate of the two exotic macrophytes Ludwigia grandiflora and Egeria densa in the presence of the native macrophyte Myriophyllum spicatum, which could indirectly facilitate their invasion success. Indeed, fragmentation can increase dispersal abilities of the exotic macrophytes considered in this study as they are able to generate new plants from their fragments. However, our results showed that the interaction herbivore-neighbor species was hardly significant. Our paper presents some first results on associational resistance/susceptibility and lays the foundation for developing a general framework that combines plant community ecology and biological invasion ecology to explain invasive species success.

Germination and Seedling Growth of Water Primroses: A Cross Experiment between Two Invaded Ranges with Contrasting Climates.

Aquatic ecosystems are vulnerable to biological invasions, and will also be strongly impacted by climate change, including temperature increase. Understanding the colonization dynamics of aquatic invasive plant species is of high importance for preservation of native biodiversity. Many aquatic invasive plants rely on clonal reproduction to spread, but mixed reproductive modes are common. Under future climate changes, these species may favor a sexual reproductive mode. The aim of this study was to test the germination capacity and the seedling growth of two water primrose species, Ludwigia hexapetala and Ludwigia peploides, both invasive in Europe and in the United States. We performed a reciprocal transplant of seeds of L. hexapetala and L. peploides from two invasive ranges into experimental gardens characterized by Oceanic and Mediterranean-type climates. Our results showed that higher temperatures increased or maintained germination percentages and velocity, decreased survivorship of germinants, but increased their production of biomass. The origin of the seeds had low impact on L. hexapetala responses to temperature, but greatly influenced those of L. peploides. The invasiveness of water primroses in ranges with Oceanic climates might increase with temperature. The recruitment from seed banks by these species should be considered by managers to improve the conservation of native aquatic and wetland plant species.

Experimental assessment of the water quality influence on the phosphorus uptake of an invasive aquatic plant: biological responses throughout its phenological stage.

Understanding how an invasive plant can colonize a large range of environments is still a great challenge in freshwater ecology. For the first time, we assessed the relative importance of four factors on the phosphorus uptake and growth of an invasive macrophyte Elodea nuttallii (Planch.) St. John. This study provided data on its phenotypic plasticity, which is frequently suggested as an important mechanism but remains poorly investigated. The phosphorus uptake of two Elodea nuttallii subpopulations was experimentally studied under contrasting environmental conditions. Plants were sampled in the Rhine floodplain and in the Northern Vosges mountains, and then maintained in aquaria in hard (Rhine) or soft (Vosges) water. Under these conditions, we tested the influence of two trophic states (eutrophic state, 100 µg x l(-1) P-PO4(3-) and hypertrophic state, 300 µg x l(-1) P-PO4(3-)) on the P metabolism of plant subpopulations collected at three seasons (winter, spring and summer). Elodea nuttallii was able to absorb high levels of phosphorus through its shoots and enhance its phosphorus uptake, continually, after an increase of the resource availability (hypertrophic > eutrophic). The lowest efficiency in nutrient use was observed in winter, whereas the highest was recorded in spring, what revealed thus a storage strategy which can be beneficial to new shoots. This experiment provided evidence that generally, the water trophic state is the main factor governing P uptake, and the mineral status (softwater > hardwater) of the stream water is the second main factor. The phenological stage appeared to be a confounding factor to P level in water. Nonetheless, phenology played a role in P turnover in the plant. Finally, phenotypic plasticity allows both subpopulations to adapt to a changing environment.

Characterization of the salt stress vulnerability of three invasive freshwater plant species using a metabolic profiling approach.

The effects of salt stress on freshwater plants has been little studied up to now, despite the fact that they are expected to present different levels of salt sensitivity or salt resistance depending on the species. The aim of this work was to assess the effect of NaCl at two concentrations on three invasive freshwater species, Elodea canadensis, Myriophyllum aquaticum and Ludwigia grandiflora, by examining morphological and physiological parameters and using metabolic profiling. The growth rate (biomass and stem length) was reduced for all species, whatever the salt treatment, but the response to salt differed between the three species, depending on the NaCl concentration. For E. canadensis, the physiological traits and metabolic profiles were only slightly modified in response to salt, whereas M. aquaticum and L. grandiflora showed great changes. In both of these species, root number, photosynthetic pigment content, amino acids and carbohydrate metabolism were affected by the salt treatments. Moreover, we are the first to report the salt-induced accumulation of compatible solutes in both species. Indeed, in response to NaCl, L. grandiflora mainly accumulated sucrose. The response of M. aquaticum was more complex, because it accumulated not only sucrose and myo-inositol whatever the level of salt stress, but also amino acids such as proline and GABA, but only at high NaCl concentrations. These responses are the metabolic responses typically found in terrestrial plants.

Does competition for phosphate supply explain the invasion pattern of Elodea species?

Two invasive aquatic plants, Elodea canadensis and Elodea nuttallii, occurred in north-eastern France. In this study, we examine the influence of phosphorus availability in soft water streams to explain the invasion pattern of exotic species (E. nuttallii and E. canadensis) compared to native plants (Callitriche platycarpa, Ranunculus peltatus). Total phosphorus was measured in these four aquatic macrophytes. Sediment total phosphorus and water-soluble reactive phosphorus were also analysed each season in 2001. Phosphorus content in the two invasive species and in R. peltatus was higher than in C. platycarpa. Elodea species are adapted to the seasonal phosphorus fluctuations as well as R. peltatus and exhibited high phosphorus storage ability. The high fluctuation availability of resources in space or/and time favoured the spread of the invasive plants and confirms the theory of invasibility of Davis et al. [2000. Fluctuating resources in plant communities: a general theory of invasibility. J. Ecol. 88, 528-534]. The eutrophication process increases the invasibility of E. nuttallii's, while inducing competition between E. nuttallii and native macrophyte species.

Are trophic and diversity indices based on macrophyte communities pertinent tools to monitor water quality?

Diversity and trophic indices based on macrophyte communities were calculated to test their pertinence to monitoring water quality in the Northern Vosges (NE of France). Highly significant correlations were found between the four tested chemical variables (bicarbonate, calcium, phosphorus and ammonium nitrogen) and trophic indices and between them and abundance and richness. Trophic indices and McIntosh's index appeared to be more effective in predicting water quality than diversity indices. Diversity indices do not necessarily provide any direct information on the quality or degree of degradation of the environment from which the sample was taken, whereas trophic indices do.