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.

Citizen Science Improves the Known and Potential Distribution of a Strong Wetland Invader: Implications for Niche Modeling and Invasion Management.

Invasive alien species are one of the main causes of biodiversity loss and ecosystem alteration. Obtaining up-to-date occurrence records and accurate invasion risk maps has become crucial to develop timely and effective management strategies. Unfortunately, gathering and validating distribution data can be labor-intensive and time-consuming, with different data sources unavoidably leading to biases in the results. In this study, we evaluated the performance of a tailored citizen science project compared with other data sources, in mapping the current and potential distribution of Iris pseudacorus, a strong invasive alien plant in Argentina. To do so, we used geographic information systems and ecological niche modeling with Maxent, and compared data from: i) a citizen science tailored project; ii) the Global Biodiversity Information Facility (GBIF); and iii) an exhaustive professional data collection (i.e. field samplings across Argentina, literature and collections review). Results suggest that the citizen science tailored project provided a larger and more diversified amount of data compared to the other sources. All data-sources showed good performance in the ecological niche models, however, data from the tailored citizen science project predicted a greater suitable area, including regions not yet reported. This allowed us to better identify critical and vulnerable areas, where management and prevention strategies are necessary. Professional data provided more reports in non-urban areas, whereas citizen science based data sources (i.e. GBIF and the citizen science project conducted in this study) reported more sites in urban areas, which indicates that different data-sources are complementary and there is a big potential in combining methods. We encourage the use of tailored citizen science campaigns to gather a more diverse amount of data, generating better knowledge about aquatic invasive species and helping decision-making in ecosystem management.

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.

Where are the seeds? Lack of floral morphs prevent seed production by the tristylous Pontederia cordata in South Africa.

Reproduction is a crucial part of the successful establishment and spread of an invasive species. Invasive plants often produce seeds prolifically to spread into new ranges, yet the invasive macrophyte, Pontederia cordata L., does not appear to produce seeds in South Africa, limiting its invasive potential. Here, we aimed to determine what limits seed production of the tristylous P. cordata in South Africa, where it is widespread with impacts on the ecology of wetlands it invades, South Africa. We measured floral traits and pollen grain size from populations throughout the invasive range in South Africa to determine the relative proportion of tristylous morphs. We speculated that the absence of specialist native pollinators in the invasive range may be responsible for the absence of sexual reproduction and thus conducted a pollination study to determine whether flowers were visited. Thereafter, we hand pollinated 8865 flowers to conclude whether P. cordata exhibited an incompatibility system, which prevented seed production. The floral traits and pollen grain measurements were similar to those reported for short-morphed flowers from the native range. The pollination study confirmed the absence of specialist insect visitors, while the hand-pollination experiments resulted in no seed production. Only short-morphed plants are present in South Africa, and the illegitimate pollination of short-morphed plants prevents seed production. Vegetative spread through rhizome production is thus responsible for the invasion of P. cordata throughout South Africa. These findings suggest that control programs should target the plants' rhizomes to prevent and reduce spread. More importantly, preventing the introduction of medium- and long-morphed plants into South Africa is crucial to preclude P. cordata from producing seeds and enhancing invasion.

Climate Mismatch between Introduced Biological Control Agents and Their Invasive Host Plants: Improving Biological Control of Tropical Weeds in Temperate Regions.

Many weed biological control programs suffer from large-scale spatial variation in success due to restricted distributions or abundances of agents in temperate climates. For some of the world's worst aquatic weeds, agents are established but overwintering conditions limit their survival in higher latitudes or elevations. The resulting need is for new or improved site- or region-specific biological control tools. Here, we review this challenge with a focus on low-temperature limitations of agents and propose a roadmap for improving success. Investigations across spatial scales, from global (e.g., foreign exploration), to local (selective breeding), to individual organisms (molecular modification), are discussed. A combination of traditional (foreign) and non-traditional (introduced range) exploration may lead to the discovery and development of better-adapted agent genotypes. A multivariate approach using ecologically relevant metrics to quantify and compare cold tolerance among agent populations is likely required. These data can be used to inform environmental niche modeling combined with mechanistic modeling of species' fundamental climate niches and life histories to predict where, when, and at what abundance agents will occur. Finally, synthetic and systems biology approaches in conjunction with advanced modern genomics, gene silencing and gene editing technologies may be used to identify and alter the expression of genes enhancing cold tolerance, but this technology in the context of weed biological control has not been fully explored.

Mind the gap: the delayed recovery of a population of the biological control agent Megamelus scutellaris Berg. (Hemiptera: Delphacidae) on water hyacinth after winter.

Cold winter temperatures significantly affect the biological control effort against water hyacinth, Pontederia ( = Eichhornia) crassipes Mart. (Pontederiaceae), in more temperate regions around the world. The population dynamics of the planthopper Megamelus scutellaris Berg. (Hemiptera: Delphacidae), a newly released biological control agent of water hyacinth, were recorded on the Kubusi River in the Eastern Cape Province (South Africa) over 15 months to determine the population recovery post-winter. Megamelus scutellaris incurred a severe population decline at the onset of winter when the water hyacinth plants became frost damaged. The combined effect of a population bottleneck and low minimum winter temperatures (6.12°C) below the agent's lower developmental threshold (11.46°C) caused a post-winter lag in agent density increase. Subsequently, the maximum agent population density was only reached at the end of the following summer growing season which allowed the water hyacinth population to recover in the absence of any significant biological control immediately post-winter. Supplementary releases of agents from mass-reared cultures at the beginning of the growing season (spring) is suggested as a potential method of reducing the lag-period in field populations in colder areas where natural population recovery of agents is slower.

Modeling Top-Down and Bottom-Up Drivers of a Regime Shift in Invasive Aquatic Plant Stable States.

The evidence for alternate stable states characterized by dominance of either floating or submerged plant dominance is well established. Inspired by an existing model and controlled experiments, we conceptually describe a dynamic that we have observed in the field using a simple model, the aim of which was to investigate key interactions of the shift between invasive floating and invasive submerged plant dominance, driven by the rapid decomposition of floating plants as a consequence of herbivory by biological control agents. This study showed that the rate of switch between floating and submerged invasive plant dominance, and the point in time at which the switch occurs, is dependent on the nutrient status of the water and the density of biological control agents on floating plant populations. Therefore, top-down invasive plant biological control efforts using natural enemies can affect systems on a wider scale than the intended agent - plant level, and can be significantly altered by bottom-up changes to the system, i.e., nutrient loading. The implications of this are essential for understanding the multiple roles invasive plants and their control have upon ecosystem dynamics. The results emphasize the importance of multi-trophic considerations for future invasive plant management and offer evidence for new pathways of invasion. The model outputs support the conclusion that, after the shift and in the absence of effective intervention, a submerged invasive stable state will persist.

Two in one: cryptic species discovered in biological control agent populations using molecular data and crossbreeding experiments.

There are many examples of cryptic species that have been identified through DNA-barcoding or other genetic techniques. There are, however, very few confirmations of cryptic species being reproductively isolated. This study presents one of the few cases of cryptic species that has been confirmed to be reproductively isolated and therefore true species according to the biological species concept. The cryptic species are of special interest because they were discovered within biological control agent populations. Two geographically isolated populations of Eccritotarsus catarinensis (Carvalho) [Hemiptera: Miridae], a biological control agent for the invasive aquatic macrophyte, water hyacinth, Eichhornia crassipes (Mart.) Solms [Pontederiaceae], in South Africa, were sampled from the native range of the species in South America. Morphological characteristics indicated that both populations were the same species according to the current taxonomy, but subsequent DNA analysis and breeding experiments revealed that the two populations are reproductively isolated. Crossbreeding experiments resulted in very few hybrid offspring when individuals were forced to interbreed with individuals of the other population, and no hybrid offspring were recorded when a choice of mate from either population was offered. The data indicate that the two populations are cryptic species that are reproductively incompatible. Subtle but reliable diagnostic characteristics were then identified to distinguish between the two species which would have been considered intraspecific variation without the data from the genetics and interbreeding experiments. These findings suggest that all consignments of biological control agents from allopatric populations should be screened for cryptic species using genetic techniques and that the importation of multiple consignments of the same species for biological control should be conducted with caution.

Unraveling the biogeographic origins of the Eurasian watermilfoil (Myriophyllum spicatum) invasion in North America.

PREMISE OF THE STUDY: Using phylogeographic analyses to determine the geographic origins of biological invaders is important for identifying environmental adaptations and genetic composition in their native range as well as biocontrol agents among indigenous herbivores. Eurasian watermilfoil (Myriophyllum spicatum) and its hybrid with northern watermilfoil (M. sibiricum) are found throughout the contiguous United States and southern Canada, forming one of the most economically costly aquatic plant invasions in North America, yet the geographic origin of the invasion remains unknown. The objectives of our study included determining the geographic origin of Eurasian watermilfoil in North America as well as the maternal lineage of the hybrids. METHODS: DNA sequence data from a cpDNA intron and the nrDNA ITS region were compiled for accessions from 110 populations of Eurasian watermilfoil and hybrids from North America and the native range (including Europe, Asia, and Africa). Datasets were analyzed using statistical parsimony and Bayesian phylogenetics to assess the geographic origin of the invasion. KEY RESULTS: The two Eurasian watermilfoil cpDNA haplotypes in North America are also found from China and Korea, but not elsewhere in the native range. These haplotypes did not overlap and were limited in native geographic range. The ovule parent for hybrids can come from either parental lineage, and multiple haplotypes from both parental species were found. CONCLUSIONS: The geographic origin of this prolific aquatic plant invasion of North America is in Asia. This provides critical information to better understand the invasion pathway and inform management into the future.