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The Emergency Recovery Plan for freshwater biodiversity recognizes that addressing nonnative species is one of six principal actions needed to bend the curve in freshwater biodiversity loss. This is because introduction rates of nonnative species continue to accelerate globally and where these species develop invasive populations, they can have severe impacts on freshwater biodiversity. The most effective management measure to protect freshwater biodiversity is to prevent introductions of nonnative species. Should a nonnative species be introduced, however, then its early detection and the implementation of rapid reaction measures can avoid it establishing and dispersing. If these measures are unsuccessful and the species becomes invasive, then control and containment measures can minimize its further spread and impact. Minimizing further spread and impact includes control methods to reduce invader abundance and containment methods such as screening of invaded sites and strict biosecurity to avoid the invader dispersing to neighbouring basins. These management actions have benefitted from developments in invasion risk assessment that can prioritize species according to their invasion risk and, for species already invasive, ensure that management actions are commensurate with assessed risk. The successful management of freshwater nonnative species still requires the overcoming of some implementation challenges, including nonnative species often being a symptom of degraded habitats rather than the main driver of ecological change, and eradication methods often being nonspecies specific. Given the multiple anthropogenic stressors in freshwaters, nonnative species management must work with other restoration strategies if it is to deliver the Emergency Recovery Plan for freshwater biodiversity.

International audience; Invasive alien aquatic plant species (IAAPs) cause serious ecological and economic impact and are a major driver of changes in aquatic plant communities. Their invasive success is influenced by both abiotic and biotic factors. Here, we summarize the existing knowledge on the biology of 21 IAAPs (four free-floating species, eight sediment-rooted, emerged or floating-leaved species, and nine sediment-rooted, submerged species) to highlight traits that are linked to their invasive success. We focus on those traits which were documented as closely linked to plant invasions, including dispersal and growth patterns, allelopathy and herbivore defence. The traits are generally specific to the different growth forms of IAAPs. In general, the species show effective dispersal and spread mechanisms, even though sexual and vegetative spread differs strongly between species. Moreover, IAAPs show varying strategies to cope with the environment. The presented overview of traits of IAAPs will help to identify potential invasive alien aquatic plants. Further, the information provided is of interest for developing species-specific management strategies and effective prevention measures.

Floating aquatic weed infestations have negative socio-economic and environmental consequences to the ecosystems they invade. Despite the long history of invasion by macrophytes, only a few studies focus on their impacts on biodiversity, while the ecological benefits of biological control programmes against these species have been poorly quantified. We investigated the process of biotic homogenization following invasion by Pistia stratiotes on aquatic biodiversity, and recovery provided by biological control of this weed. Biotic homogenization is the increased similarity of biota as a result of introductions of non-native species. The study quantified the effect of P. stratiotes, and its biological control through the introduction of the weevil, Neohydronomus affinis on recruitment of benthic macroinvertebrates to artificial substrates. Mats of P. stratiotes altered the community composition and reduced diversity of benthic macroinvertebrates in comparison to an uninvaded control. However, reduction in percentage cover of the weed through biological control resulted in a significant increase in dissolved oxygen, and recovery of the benthic macroinvertebrate community that was comparable to the uninvaded state. This highlights the process of homogenization by an invasive macrophyte, providing a justification for sustained ecological and restoration efforts in the biological control of P. stratiotes where this plant is problematic.

Silica (Si) plays a significant role in alleviating the effects of biotic and abiotic stressors in many plants, especially in an agricultural context. With increased use of Si-based fertilisers, understanding plant responses to the addition of Si to their environment, particularly aquatic environments, is important. We investigated how two invasive macrophytes, Eichhornia crassipes and Pistia stratiotes, responded to different nutrient and Si concentrations, in the presence and absence of herbivory. Both species incorporated Si into their foliage, but uptake of Si did not increase under high nutrient availability but rather decreased, especially in P. stratiotes. Plant quality (i.e. C:N) for both weed species was affected more by nutrient concentrations than Si content, and the addition of Si had a negative effect on plant growth. Eichhornia crassipes increased daughter plant production under high Si conditions, while P. stratiotes plants showed no reproductive response to increased Si except in low nutrient conditions where reproduction was reduced. The addition of Si resulted in increased biomass of E. crassipes, while P. stratiotes was unaffected. These results highlight that runoff of Si from fertiliser alter aquatic plant–insect interactions, which has consequences for biological control.

Egeria densa Planchon (Hydrocharitaceae) is a submerged macrophyte native to South America. It forms part of a new suite of invasive aquatic plants that has benefited from open nutrient-rich freshwater systems following the successful biological control of floating aquatic plants in South Africa. The specificity of the leaf-mining fly, Hydrellia egeriae Rodrigues (Diptera: Ephydridae) was tested, using traditional laboratory host-specificity testing (i.e., no-choice and paired choice). Only one non-target species, Lagarosiphon major Deeming (Hydrocharitaceae) supported larval development during pair-choice tests. In order to avoid the rejection of a safe and potentially effective agent, continuation (i.e., multiple generations) tests were conducted to measure the ability of the non-target species to nutritionally support a population indefinitely. None of these species could sustain a viable agent population for more than three generations. Laboratory host-specificity tests are limited as they exempt certain insect-host behaviours. To enhance the interpretation of host-specificity results, a risk assessment was conducted using agent preference (i.e., choice tests) and performance (i.e., choice and continuation tests) results. The feeding and reproductive risk that H. egeriae poses to non-target species is below 2%. Based on these findings, permission for its release in South Africa has been obtained.

Water hyacinth is considered the most damaging aquatic weed in South Africa. The success of biocontrol initiatives against the weed varies nation-wide, but control remains generally unattainable in higher altitude, temperate regions. Eccritotarsus catarinensis (Hemiptera: Miridae) is a biocontrol agent of water hyacinth that was first released in South Africa in 1996. By 2011, it was established at over 30 sites across the country. These include the Kubusi River, a site with a temperate climate where agent establishment and persistence was unexpected. This study compared the critical thermal limits of the Kubusi River insect population with a laboratory-reared culture to determine whether any physiological plasticity was evident that could account for its unexpected establishment. There were no significant differences in critical thermal maxima (CTmax) or minima (CTmin) between sexes, while the effect of rate of temperature change on the thermal parameters in the experiments had a significant impact in some trials. Both CTmax and CTmin differed significantly between the two populations, with the field individuals tolerating significantly lower temperatures (CTmin: −0.3°C ± 0.063 [SE], CTmax: 42.8°C ± 0.155 [SE]) than those maintained in the laboratory (CTmin: 1.1°C ± 0.054 [SE], CTmax: 44.9°C ± 0.196 [SE]). Acclimation of each population to the environmental conditions typical of the other for a five-day period illustrated that short-term acclimation accounted for some, but not all of the variation between their lower thermal limits. This study provides evidence for the first cold-adapted strain of E. catarinensis in the field, with potential value for introduction into other colder regions where water hyacinth control is currently unattainable.

Simple Summary Mismatched distributions between biological control agents and their host plants occur for a variety of reasons but are often linked to climate, specifically differences in their low-temperature tolerances. How to measure and use low-temperature tolerances of control agents to inform agent prioritization, selection for redistribution, or predict efficacy is vitally important, but has not been previously synthesized in a single source. We discuss causes of climate mismatches between agents and target weeds, the traditional and non-traditional approaches that could be used to decrease the degree of mismatch and improve control, and regulatory issues to consider when taking such approaches. We also discuss the variety of cold tolerance metrics, their measurement and ecological value, and the types of modeling that can be carried out to improve predictions about potential distributions of agents. We also briefly touch on molecular bases for cold tolerance and opportunities for improving cold tolerance of agents using modern molecular tools. Abstract 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.

The Brazilian waterweed, Egeria densa Planchon, 1849 (Hydrocharitaceae), is an invasive species in South Africa where it is a host plant for the aquatic leaf-miner Hydrellia egeriae Rodrigues-Júnior, 2015 (Ephydridae, Diptera). Efficacy of the biocontrol agent can potentially be affected by parasitoids. Three species of braconid parasitoid wasps were reared from puparia of Hydrellia egeriae. By comparison with the type specimens, these species have been determined to be Ademon lagarosiphonae van Achterberg, 2012 (Braconidae: Opiinae), Chaenusa anervata van Achterberg, 2012 and Chaenusa seminervata van Achterberg, 2012 (Braconidae: Alysiinae: Dacnusini), all previously recorded as parasitoids of an ephydrid dipterous aquatic leaf-miner, Hydrellia lagarosiphon Deeming, 2012, on Lagarosiphon major (Ridley, 1886) Moss ex Wager (Hydrocharitaceae) in South Africa. The chalcidoid, Janicharis africanus Gumovsky & Delvare, 2006 (Eulophidae), was also reared from Hydrellia egeriae and is possibly a hyperparasitoid of the braconids. South Africa is a new country record for J. africanus. We provide comprehensive images of all species including the braconid types and illustrated identification keys to the Afrotropical species of the two braconid genera are also provided. All images and online keys are available on WaspWeb (http://www.waspweb.org).

Salvinia molesta D.S. Mitchell (Salviniaceae) is a damaging free-floating invasive alien macrophyte native to South America. The biological control programme against S. molesta by the weevil Cyrtobagous salviniae Calder and Sands (Erirhinidae) has been successful in controlling S. molesta infestations in the introduced range, however, there is some debate as to how biological control success is measured. This study measured the response of epilithic algae and aquatic macroinvertebrate communities in a S. molesta-dominated state and subsequently where the weed had been cleared by biological control, as a proxy for ecosystem recovery in a before&ndash, after control&ndash, impact mesocosm experiment. The restored treatment (S. molesta and C. salviniae) demonstrated epilithic algae and aquatic macroinvertebrate recovery during the &ldquo, after&rdquo, biological control phase, defined as similar to the control treatment. Comparatively, the impacted treatment (100% S. molesta) showed a drastic decline in biodiversity and shifts in community assemblages. We conclude that the biological control effort by C. salviniae facilitated biodiversity recovery of the impacted treatment. Furthermore, epilithic algae and aquatic macroinvertebrate communities were reliable biological indicators for measuring ecological impacts of invasion and ecosystem recovery following biological control, and thus represent potential tools for evaluating biological control success and ecological restoration.

International audience; Globally, freshwater ecosystems are under threat. The main threats come from catchment land-use changes, altered water regimes, eutrophication, invasive species, climate change and combinations of these factors. We need scientific research to respond to these challenges by providing solutions to halt the deterioration and improve the condition of our valuable freshwaters. This requires a good understanding of aquatic ecosystems, and the nature and scale of changes occurring. Macrophytes play a fundamental role in aquatic systems. They are sensitive indicators of ecosystem health, as they are affected by run-off from agricultural, industrial or urban areas. On the other hand, alien macrophytes are increasingly invading aquatic systems all over the world. Improving our knowledge on the ecology and management of both native and alien plants is indispensable to address threats to freshwaters in order to protect and restore aquatic habitats. The International Aquatic Plants Group (IAPG) brings together scientists and practitioners based at universities, research and environmental organisations around the world. The main themes of the 15th symposium 2018 in New Zealand were biodiversity and conservation, management, invasive species, and ecosystem response and restoration. This Virtual Special Issue provides a comprehensive review from the symposium, addressing the ecology of native macrophytes, including those of conservation concern, and highly invasive alien macrophytes, and the implications of management interventions. In this editorial paper, we highlight insights and paradigms on the ecology and management of native and alien macrophytes gathered during the meeting.

Invasive alien plant species negatively affect agricultural production, degrade conservation areas, reduce water supplies, and increase the intensity of wild fires. Since 1913, biological control agents i.e. plant-feeding insects, mites, and fungal pathogens, have been deployed in South Africa to supplement other management practices (herbicides and mechanical controls) used against these invasive plant species. We do not describe the biological control agent species that have been used, or what they do, or how they damage the target plant species. We focus instead on evaluations (informed opinions, evidence and quantifications) of what has been achieved in South Africa by using biological control to suppress populations and impacts of invasive plant species. Satisfactory long-term evaluations of outcomes are difficult and expensive, but many have been done, providing ample evidence that biological control is often highly successful. We use case studies from South Africa to support this assertion and to make the point that successes may be largely forgotten in a relatively short time. Biological control of invasive plants in South Africa is demonstrably cost effective and has become generally accepted as a preferred management strategy. However, it is not a panacea, and we discuss several issues which complicate our understanding of its effectiveness and which raise research and implementation challenges. Further studies are needed on the economic and social benefits of biological control of invasive plants to inform and involve the wider South African community.

1. Echthrodesis lamorali Masner, 1968 is the only known parasitoid of the eggs of the intertidal rocky shore spider Desis formidabilis O.P. Cambridge 1890 and is endemic to a small area of South Africa. 2. The abundance of spider nests and parasitoid presence were assessed in relation to their in- and between-shore location at multiple sites within the distribution of E. lamorali along the Cape Peninsula (Western Cape, South Africa). 3. Desis formidabilis nests were more abundant in the mid-shore zone than higher up or lower down the shore. Spider population sizes also differed between collection sites, with higher numbers recorded on the cooler western coast of the peninsula. 4. Evidence of parasitoid activity was recorded in 43.31% of the 127 nests and 13.85% of the 592 egg sacs they contained. 5. Where parasitoids gained entry to a spider egg sac, oviposition took place into all of the eggs present. 6. Incidence of wasp activity was positively correlated with spider nest concentration, not with height up the shore, suggesting that both the host and parasitoid are tolerant of salt-water inundation. 7. These results should assist managers of the Table Mountain National Park, in which the full distribution of E. lamorali falls, to better understand this component of rocky shore community dynamics.

Natural enemies are known to modify competitive hierarchies among terrestrial plants. Here we examine whether the same applies to freshwater systems. Lagarosiphon major (Hydrocharitaceae) is a submerged aquatic macrophyte, indigenous to South Africa. Outside its native range, it outcompetes with indigenous submerged species and degrades aquatic habitats. Hydrellia lagarosiphon (Diptera: Ephydridae) is the most abundant and ubiquitous herbivore associated with L. major in South Africa and is a potential biological control agent elsewhere. Chaenusa anervata (Hymenoptera: Braconidae: Alysiinae) is its main parasitoid. We generated an experimental system involving one, two or three trophic levels to monitor variation in the competitive ability of L. major relative to that of Myriophyllum spicatum (Haloragaceae), a second submerged macrophyte that can also be invasive. Using inverse linear models to monitor competition, we found that herbivory by H. lagarosiphon greatly reduced the competitive ability of L. major. Addition of the wasp at typical field densities halved the impact of herbivory and re-established the competitive advantage of L. major. Our results demonstrate how multitrophic interactions modify relative competitive abilities among aquatic plants, emphasize the significance of higher tropic levels in these systems and illustrate how parasitoids can reduce the effectiveness of insects released as biocontrol agents.

Water hyacinth is invasive in many countries, where it reduces aquatic biodiversity and limits water resource utilisation. Biological control of water hyacinth has been successful in South Africa, but has suffered from a lack of empirical data to prove causation. Insect exclusion trials were conducted to quantify the contribution of Neochetina eichhorniae and N. bruchi to the integrated control of water hyacinth on the Nseleni River, South Africa. Insecticide was not expected to induce phytotoxicity, but would prevent weevil damage in water hyacinth plants; and weevil herbivory was predicted to reduce plant petiole length, and above/below surface biomass. Results showed that insecticide had no phytotoxic effects and excluded weevils for 3 weeks, providing a baseline for field applications. Biological control on the Nseleni River directly affected water hyacinth biomass and petiole length, but did not affect plant cover. Plants subject to weevil herbivory demonstrated reductions in above and below surface biomass and had shorter petioles compared to insect-free plants. Dead biomass was also higher in biological control treatments. Biological control strongly affects plant size, biomass and vigour; however, further integrated control is required to facilitate reduction in mat cover, which is the goalpost for successful control of floating aquatic plants.

Insect biological control agents directly damage target weeds by removal of plant biomass, but herbivorous insects have both direct and indirect impacts on their host plants and can also facilitate pathogen infection. Megamelus scutellaris Berg (Hemiptera: Delphacidae) was recently released into South Africa to help control invasive water hyacinth (Eichhornia crassipes, Pontederiaceae). We compared the impact of fungicide surface-sterilised and unsterilised M. scutellaris individuals and water hyacinth leaves on growth of the weed at two nutrient levels. The survival and reproduction of adult M. scutellaris was not reduced by sterilisation. Under high nutrient conditions, unsterilised M. scutellaris with unsterilised leaves reduced water hyacinth daughter plant production by 32%, length of the second petiole by 15%, chlorophyll content by 27% and wet weight biomass by 48%, while also increasing leaf chlorosis 17-fold, in relation to control plants under the same nutrient regime. Surface sterilisation of the insect and/or plant surfaces led to a general reduction in these impacts on water hyacinth growth and health. This contrast was less evident under low nutrient conditions. Megamelus scutellaris facilitated infection by fungal and other pathogens, thus its biology is compatible with pathogens that could be developed into mycoherbicides. This integrated approach may be ideal for management of infestations of water hyacinth in eutrophic water systems where control has been problematic, both in South Africa and elsewhere.

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.

Biological control of Azolla filiculoides in South Africa with the Azolla specialist Stenopelmus rufinasus has been highly successful. However, field surveys showed that the agent utilized another Azolla species, thought to be the native Azolla pinnata subsp. africana, which contradicted host specificity trials. It is notoriously difficult to determine Azolla species based on morphology so genetic analyses were required to confirm the identity of the Azolla used by the agent. Extensive sampling was conducted and samples were sequenced at the trnL-trnF and trnG-trnR chloroplastic regions and the nuclear ITS1 region. Current literature reported A. filiculoides as the only Section Azolla species in southern Africa but 24 samples were identified as Azolla cristata, an introduced species within Section Azolla that was not used during host specificity trials. A. pinnata subsp. africana was only located at one site in southern Africa, while the alien A. pinnata subsp. asiatica was located at three. What was thought to be A. pinnata subsp. africana was in fact A. cristata, a closer relative of A. filiculoides and a suitable host according to specificity trials. This study confirms that S. rufinasus is a proficient Azolla taxonomist but also supports the use of molecular techniques for resolving taxonomic conundrums.

Variability in aquatic plant morphology is usually driven by phenotypic plasticity and local adaptations to environmental conditions experienced. This study aimed to elucidate which of these drivers is responsible for the morphological variation exhibited by three populations of Myriophyllum spicatum L. (Haloragaceae), a submerged aquatic plant whose status as native or exotic within southern Africa is uncertain. Individuals from three populations on the Vaal River (Northern Cape), Klipplaat River (Eastern Cape) and Lake Sibaya (KwaZulu-Natal) were grown under two nutrient treatments (high: 30 mg N/kg sediment and low: sediment only), while all other variables were kept the same. Morphological characteristics were measured at the start of the experiment to obtain a baseline morphology, and again eight weeks later. By the end of the experiment, the individuals from each population had responded to the different growing conditions. In most cases, the individuals from each population were significantly larger under the high nutrient treatment (Stem diameter: F(5,86) = 18.435, P

There is debate over the native or exotic status of Myriophyllum spicatum L. (Haloragaceae) in South Africa, which has important implications for developing and implementing management strategies. The aim of this study was to determine if M. spicatum was recently introduced from Eurasia by reconstructing the genetic relationships between South African and Eurasian M. spicatum using both a nuclear ribosomal (ITS1-5.8S-ITS2-26S) and a chloroplast intron (trnQ-rps16) sequence from 40 populations. For both these DNA markers, the South African populations were distinct from Eurasian populations, but always stemmed from a European origin. The data suggest that South African and European M. spicatum share a common ancestor, however the divergence of both markers are characteristic of a long period of isolation rather than a recent introduction from Europe. The genetic data from this study suggest that M. spicatum has not been introduced recently, but is most likely a native component of the South African flora.

Nymphaea mexicana Zuccarini (Nymphaeaceae) (Mexican waterlily) is an emergent floating-leaved aquatic plant from the southeastern USA that is invasive in South Africa. In invaded waterbodies this plant restricts water movement, increases siltation, decreases recreational activities, and can deplete water oxygen levels, which in turn negatively impacts aquatic fauna. Currently there are no chemical, mechanical, or biological control programs in place for N. mexicana in South Africa, but the sustainability of biological control makes this the most desirable option. Field surveys for potential biological control agents were conducted in the native range of N. mexicana in Florida, Louisiana, and Texas from Aug to Oct 2018. Leaves, stems, flowers, and roots of N. mexicana were searched for insect herbivores by hand and using Berlese funnels. Insects were prioritized for use as biological control agents by considering the extent and type of feeding damage, field host range, and incidence (percentage of sites in which each species was found). In total, 15 confirmed species were found feeding on N. mexicana, and some taxa were identified only to family level. Incidence coverage estimator mean, MMRuns, Chao 2 mean, and Chao 2 upper 95% CI species accumulation estimators predicted that between 2 and 5 species were missed during the surveys. Based on field observations, Bagous americanus LeConte (Coleoptera: Curculionidae), and Megamelus toddi Beamer (Hemiptera: Delphacidae) were prioritized. Host specificity trials will be conducted to determine whether these insects may be used as biological control agents of N. mexicana.

Specialist herbivores have evolved adaptations to overcome plant defensive chemicals and thus prefer younger leaves with their higher nutritional value, whereas generalist herbivores are unable to overcome these chemical defences, and opt for older leaves which have fewer nutrients, but are less defended (Center & Wright 1990). Plants such as the invasive aquatic weed Eichhornia crassipes (Martius) Solms-Laubach (Pontederiaceae) (water hyacinth) continuously produce young, unfurling leaves that require high concentrations of digestibility-reducing chemicals in their tissues to ensure protection against herbivory (Center & Wright 1990). These defensive chemicals are products of secondary metabolism (Berenbaum 1995) and ensure that young leaves are typically unpalatable to herbivores and resistant to disease (Center & Wright 1990).

Background : Aquatic ecosystems in South Africa are prone to invasion by several invasive alien aquatic weeds, most notably, Eichhornia crassipes (Mart.) Solms-Laub. (Pontederiaceae) (water hyacinth); Pistia stratiotes L. (Araceae) (water lettuce); Salvinia molesta D.S. Mitch. (Salviniaceae) (salvinia); Myriophyllum aquaticum (Vell. Conc.) Verd. (parrot’s feather); and Azolla filiculoides Lam. (Azollaceae) (red water fern). Objective : We review the biological control programme on waterweeds in South Africa. Results : Our review shows significant reductions in the extent of invasions, and a return on biodiversity and socio-economic benefits through the use of this method. These studies provide justification for the control of widespread and emerging freshwater invasive alien aquatic weeds in South Africa. Conclusions : The long-term management of alien aquatic vegetation relies on the correct implementation of biological control for those species already in the country and the prevention of other species entering South Africa.

The southern African coastline plays host to nine spider species. Two of these, namely Desis formidabilis (O. P.-Cambridge, 1890) (Araneae: Desidae) and Amaurobioides africanus Hewitt, 1917 (Araneae: Anyphaenidae), are recorded as hosts for an intertidal spider egg parasitoid, Echthrodesis lamorali Masner, 1968 (Hymenoptera: Platygastridae: Scelioninae). These two spider species occur from Luderitz (Namibia) along the coast to East London (Eastern Cape Province, South Africa), while their parasitoid has been known from only a single locality on the Cape Peninsula. The South African coastline was surveyed from Jacobsbaai (Western Cape Province) to East London in an attempt to determine the full distribution of E. lamorali. The wasp was only reared from host eggs collected on the Cape Peninsula, confirming a high degree of endemism for this species.

The submerged aquatic macrophyte Myriophyllum spicatum L. (Haloragaceae) has been recorded in southern Africa since 1829. It was first considered problematic in 2005 on the Vaal River, which has highlighted the need for an assessment of the invasion status and an appropriate management strategy to be identified. We used a unified invasion framework to categorise M. spicatum in southern Africa and define appropriate management strategies. Historical records and field surveys were used to assess the invasion status. Populations were considered established if collection records from a particular catchment or river system spanned at least 10 years and in some cases several localities. Of the 21 river systems where it has been recorded, it was evaluated as established in 13 and in the D1-E category of invasion. The disjunct populations suggest that it was point source introductions and thus the major barrier that prevents M. spicatum from proceeding along the invasion framework is its inability to naturally disperse the greater distances between catchments. Appropriate management strategies need to be catchment or river system based depending on the stage or state and category of invasion. Where M. spicatum is in the D1-E stage – established and potentially spreading, the appropriate management strategies include containment, and where possible, mitigation.

The leaf-mining fly, Hydrellia lagarosiphon Deeming (Diptera: Ephydridae), was investigated in its native range in South Africa, to determine its potential as a biological control agent for Lagarosiphon major (Ridl.) Moss ex Wager (Hydrocharitaceae), an invasive submerged macrophyte that is weedy in many parts of the world. The fly was found throughout the indigenous range of the plant in South Africa. High larval abundance was recorded at field sites with nearly all L. major shoots sampled ontaining larvae, with densities of up to 10 larvae per shoot. Adults laid batches of up to 15 eggs, usually on the abaxial sides of L. major leaves. The larvae mined internally, leaving the epidermal tissues of the upper and lower leaves intact. The larvae underwent three instars which took an average of 24 days and pupated within the leaf tissue, from which the adults emerged. Impact studies in the laboratory showed that H. lagarosiphon larval feeding significantly restricted the formation of L. major side branches. Based on its biology and damage caused to the plant, Hydrellia lagarosiphon could be considered as a useful biological control candidate for L. major in countries where the plant is invasive.

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.

The integrated control of water hyacinth, Eichhornia crassipes (Martius) Solms-Laubach (Pontederiaceae) has become necessary in South Africa, as biological control alone is perceived to be too slow in controlling the weed. In total, seven insect biological control agents have been released on water hyacinth in South Africa. At the same time, herbicides are applied by the water authorities in areas where the weed continues to be troublesome. This study investigated the assumption that the two control methods are compatible by testing the direct toxicity of a range of herbicide formulations and surfactants on two of the biological control agents released against water hyacinth, the weevil, Neochetina eichhorniae Warner (Coleoptera: Curculionidae) and the water hyacinth mirid, Eccritotarsus catarinensis (Carvalho) (Hemiptera: Miridae). A number of the formulations used resulted in significant mortality of the mirid and the weevil. Products containing 2,4-D amine and diquat as active ingredients caus...

Biological control against water hyacinth, Eichhornia crassipes (C.Mart.) Solms (Pontederiaceae), salvinia, Salvinia molesta D.S.Mitch. (Salviniaceae), water lettuce, Pistia stratiotes L. (Araceae), parrot's feather, Myriophyllum aquaticum (Vell.) Verdc. (Haloragaceae), and red water fern, Azolla filiculoides Lam. (Azollaceae) has been ongoing in South Africa since the release of the first biological control agent on water hyacinth in 1974. This review provides an account of progress for the period from 1999. Post-release evaluations over the last three years have shown that, with the exception of water hyacinth, all of these problematic aquatic plants have been suppressed effectively using classical biological control. In eutrophic water bodies at high elevations that experience cold winters, an integrated approach, that includes herbicide application and augmentive biological control, is required against water hyacinth. The grasshopper Cornops aquaticum (Bruner) (Orthoptera: Acrididae: Leptysminae) has ...

Historically, biological control efforts against aquatic plants in South Africa have focused on floating species, and as a result, there has been a dearth of research into the invasion and control of submerged macrophytes. With numerous submerged invasive species already established in South Africa, thriving horticultural and aquarium industries, nutrient-rich water systems, and a limited knowledge of the drivers of invasions of submerged macrophytes, South Africa is highly vulnerable to a second phase of aquatic plant problems. Experience gained in the U.S.A. on biological control against submerged weeds, such as hydrilla, Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) and spiked/Eurasian watermilfoil, Myriophyllum spicatum L. (Haloragaceae), have provided South African researchers with the necessary foundation to initiate programmes against these weeds. Research in South Africa is currently focused on pre-release studies on the biological control of H. verticillata, using an undescribed fly, Hydr...

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.

Recovery of indigenous species subsequent to the clearing of invasive alien plants (IAPs) is crucial for ecosystem recovery to occur. However, cleared sites are often just left in the hope that revegetation will occur naturally. In riparian areas of Kruger National Park (KNP), the Working for Water (WfW) Programme has cleared IAPs on a regular basis, but little post-clearance monitoring has taken place. Thus investigating short-term effects of IAPs and IAP clearing on plant community diversity and vegetation recovery provided an ideal opportunity to assess feasible targets of natural ecosystem recovery in similar areas. Vegetation was sampled from twelve transects along the Sabie River in and adjacent to the KNP, before (March/April 2006) and after (March 2007) the annual clearing of IAPs by WfW. Rarefied species richness, alpha diversity and evenness of distribution of species all declined with increasing density of IAPs (Pb 0.05). There was a mean reduction in IAP density of 80% (S.E ± 6%) (P = 0.002) through the clearing by WfW. After clearing of IAPs, indigenous vegetation densities increased, with herbaceous growth forms showing the largest increase in transects that were previously heavily invaded. Thus, in this system, which is relatively undisturbed by human activities, initial recovery of indigenous vegetation can occur without further restorative interventions. This process is more than likely aided by the continuous clearing of IAPs by WfW as this acts to deplete alien seed banks and maintain IAPs at acceptable and manageable levels. © 2008 SAAB. Published by Elsevier B.V. All rights reserved.

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.

Submerged aquatic invasive plant species are increasingly being recognised as a major threat to South African water ways. Pet stores, aquarists and the internet-mediated trade were investigated as pathways for submerged invasive macrophyte introductions into South Africa. Online and manually distributed surveys were used to determine the extent of movement of invasive as well as indigenous submerged plant species in South Africa. Sixty-four stores and twenty-three aquarists were surveyed. Four areas of risk were identified in this study. Firstly, and most importantly, a variety of invasive and/or prohibited plants are sold by pet stores. Secondly, there is a lack of knowledge regarding identification as well as regulation of submerged species, which may then result in the unintentional trade of potentially invasive species. It seems that, in many cases, the pet stores are ignorant or misinformed of the potential dangers, rather than intentionally attempting to breach the legislation. Thirdly, aquarists own, trade and move plants in and around the country, which makes it very difficult to monitor which species are being moved around South Africa and to what extent. Finally, the internet is a pathway of potential concern, but it is difficult to quantify its contribution to the trade of invasive species in South Africa.

We investigated water hyacinth seed banks in several aquatic systems in South Africa. Fifteen sites, mainly in the Eastern and Western Cape provinces, were surveyed from August to October 2009. Soil seed density varied between 0 and 2534 seeds/m 2 but did not differ significantly between the type of waterbody (impoundment vs. river) or the history of control carried out at a site. Average germination was 54.17% with very fast velocities (Vigour Index = 36.66) and maximum germination around three days. Although we demonstrated the existence of an important reservoir of seeds, results from this study indicated that a combination of factors such as water fluctuation, eutrophication and seed decomposition might have had a great influence on dispersal and persistence of seeds.