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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.

A B S T R A C T Iris pseudacorus L. (Iridaceae) is an emergent macrophyte native to Europe, North Africa and western Asia. Considered invasive in wetland habitats around the world, this species is now the target of a biocontrol pro- gramme in the Southern Hemisphere. Native range surveys of the weed led to the selection of the flea beetle, Aphthona nonstriata Goeze (Coleoptera: Chrysomelidae), as a candidate biocontrol agent. An important aspect to consider in weed biocontrol is the ability of an agent to establish and thrive in the environment where it is released. Climatic incompatibility between source and intended release sites can in fact limit the success of a biocontrol programme. In the current study, the potential climatic niche of I. pseudacorus and A. nonstriata in the Southern Hemisphere was analysed. The ecological niche modelling software MaxEnt was used to map the cli- matic suitability of both organisms across invaded regions in South America, southern Africa and Australasia. Furthermore, occurrence records from each invaded range were used independently to model the climatic compatibility of I. pseudacorus in Europe, in order to prioritize areas of the native range to explore during future surveys for potential biocontrol agents. The models identified areas at high risk of invasion by I. pseudacorus in northern Argentina, Uruguay, southern Brazil and central Chile, as well as numerous provinces of eastern South Africa, Lesotho, southern Australia and New Zealand. Accordingly, the highest climatic suitability for A. nonstriata was predicted across the humid temperate climates of north-east Argentina, Uruguay, southern Brazil, southern South Africa, south-east Australia and New Zealand. These results can eventually be used in future release plans to prioritize areas where establishment and survival of the agent is expected to be highest. At the same time, it may be useful to search the native range of the weed for biological control agents showing high climatic adaptation towardsthe intended release sites of each invaded range. In this regards, our climatic compatibility models identified high-priority areas across the Mediterranean regions of Italy and southern France, as well as the temperate regions of central and western Europe. Altogether, the current study provides useful new information to tackle the invasion and advance the biocontrol programme of I. pseudacorus in the Southern Hemisphere

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.

The yellow flag, Iris pseudacorus L. (Iridaceae) is an emergent perennial wetland plant native to the Palearctic region and has been dispersed worldwide as an ornamental plant. In the introduced range, it is highly invasive in natural and artificial waterbodies. Currently, this species is difficult to manage and continues to spread, causing significant environmental impacts. The aims of this study were to assess the geographical range of I. pseudacorus in Argentina by compiling data from literature, collections, citizen science and field surveys. An exhaustive review of herbarium collections and literature was conducted and a database with reports obtained through citizen science was obtained and used to design routes for field surveys. 114 records of I. pseudacorus were obtained, mainly in humid and disturbed areas. The oldest records date from 1931 and 1933, and the number of reported localities showed a gradual increase with time. Fifteen records of the weed were from protected areas, four of which have international conservation status. This study shows that I. pseudacorus is increasing in distribution in Argentina and that citizen science can be used to document the distribution of invasive alien plant species. Taking into account that the number of records obtained from herbarium collections was greater than the number of records from literature, this study shows the value of these as database that allows historical reconstructions. This study is part of a collaborative “Global South” alliance between Argentina, South Africa and New Zealand, determining the extent of I. pseudacorus invasion in the southern hemisphere and generating management strategies for its control.

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.

Water hyacinth, Pontederia crassipes (Martius) [≡Eichhornia crassip es (Martius) Solms-Laubach] (Pontederiaceae), is native to South America, but has expanded its range to many other regions of the...

The impact of the planthopper Megamelus scutellaris, a biocontrol agent of water hyacinth in South Africa, was assessed using chlorophyll fluorometry in a greenhouse study under two differe...

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.

Current global change scenarios predict an increase in atmospheric CO2 from the current 380 ppm to a value ranging from 540 ppm to 960 ppm by the year 2100. The effects of three air CO2 levels (400, 600 and 800 ppm) on five submerged aquatic plants that utilize HCO3− were studied, using the elevated CO2 Open Top Chamber facility at Rhodes University (Grahamstown, South Africa). Plants grew in water with two different initial dissolved inorganic carbon (DIC) concentrations of 1.5 and 3.0 mM. Overall, the growth rates and biomass allocation to roots were not affected by the initial DIC and air CO2, even though differences between the species were found. Furthermore, no overall effects were found on net photosynthesis, chlorophyll and starch content, even though significant effects of CO2 and DIC were observed in some species. In contrast, with increasing DIC and air CO2 a significant global decline in leaf nitrogen content linked with an increased C:N molar ratio was observed. The results indicate that submerged aquatic HCO3− users will be less affected by atmospheric CO2 increases when growing in DIC rich waters, in comparison to obligate CO2 users growing under CO2 limiting conditions as documented in previous studies. However, the changes found in plant nitrogen illustrate that atmospheric CO2 increases will affect nitrogen absorption by submerged plants, with subsequent ecosystem level effects.

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).

Native to Europe, North Africa and western Asia, Iris pseudacorus L. (Iridaceae) has invaded natural and human-modified wetlands worldwide. This species is considered a noxious weed in several countries including Argentina, South Africa and New Zealand. Its broad ecological tolerance, high resilience and reproductive potential make current mechanical and chemical control measures cost-ineffective, and biological control is considered a suitable alternative. In order to prioritise candidate biocontrol agents, a list of organisms reported to attack the plant within its native range has been assembled, and information about their host-range and damaging potential gathered from the literature. Furthermore, surveys for natural enemies of the plant were conducted in Belgium and northern Italy. The insect fauna associated with I. pseudacorus at the sites surveyed comprised mostly incidental visitors and polyphagous feeders, with the exception of the sawfly Rhadinoceraea micans Klug (Hymenoptera: Tenthredinidae), theseed weevil Mononychus punctumalbum Herbst (Coleoptera: Curculionidae), and the flea beetle Aphthona nonstriata Goeze (Coleoptera: Chrysomelidae). The potential of these species for biocontrol was evaluated, and A. nonstriata was given highest priority. A population of this species was imported to quarantine in South Africa, where it is currently undergoing host-specificity testing. Importation of the two remaining candidates is expected shortly. In conclusion, the prospects for the biological control of I. pseudacorus appear promising.

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.

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.

Salvinia molesta D.S. Mitchell (Salviniaceae) (giant salvinia), a floating aquatic fern of Brazilian origin, has been dispersed to much of the tropical and subtropical parts of the world since the mid-1900s, where it is invasive and damaging. Herbicide application and mechanical control of this weed are labor intensive and expensive, but biological control using the host-specific weevil, Cyrtobagous salviniae (Calder and Sands), provides an effective and sustainable solution. The weevil was first released as a biological control agent onto S. molesta in Australia in 1980 and has subsequently been released in further 22 countries affected by the weed. The biological control program against S. molesta has been an extraordinary success story where a single weevil species has resulted in the weed no longer being considered invasive in most countries in a relatively short time of under 3 years. Biological control in temperate regions of the world might require more time than in tropical and subtropical regions, and this difference is presumed to be due to greater winter mortality. Nevertheless, inundative releases of the weevil early in the growing season have resulted in more effective control and less dependence on herbicide applications. However, there are new records of infestations of S. molesta in particular in Africa, and these sites must be prioritized for biological control.

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.

Classical biological control has been used as a management tool for invasive non-native plant species globally for over 200 years. There have been some very successful programmes, most notably on waterweeds, cacti and seed reduction in perennial trees. Seventy per cent of agents released have established in at least one instance, and 66% of the targeted invasive species have showed some level of control. However, some programmes have failed to meet expectations, for example on Lantana camara. The most commonly cited reasons for the failure of establishment or limited efficacy of biological control agents are unsuitable climatic conditions and genotype incompatibility. We propose that antagonistic biotic interactions play a significant role in the outcomes of weed biological control programmes. Induced plant defences (physical and chemical) that can be mounted rapidly by the invasive non-native plants can result in the reduction in agent populations after initial attack. Rapid induction of plant defences have been implicated in the lack of long-term establishment of the agent Falconia intermedia that showed great initial promise against the widespread invasive shrub L. camara. Host range expansion by native natural enemies onto biological control agents have also been shown to reduce population growth of agents. Finally, competition from indigenous plant species aids invasive alien plant population reduction in the presence of herbivory. All three factors have been poorly studied and further work is needed to better explain the outcomes of weed biological control programmes.

Water lettuce, Pistia stratiotes, and red water fern, Azolla filiculoides, are floating aquatic macrophytes that have become problematic in South Africa. Two weevils, Neohydronomus affinis and Sten...

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.

Sagittaria platyphylla (Engelmann) J.G. Smith (Alismataceae) was first recorded in South Africa in 2008 and is considered to be an emerging weed with naturalised populations occurring throughout th...

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.

Biological control offers a cost effective and ecologically sustainable tool for the management of invasive alien plants. Its implementation, however, has historically been slow and poorly co-ordinated. In South Africa, as in many other countries, most aspects of biological control programmes were done by researchers, but from 1995 onwards, with the advent of the Working for Water Programme, a more inclusive approach to biological control has been adopted. In this paper, we report on the development of community-based biological control implementation programmes in South Africa, after 1995, and highlight a number of initiatives, including employing persons with disabilities at mass-rearing facilities and in particular, we outline a suite of educational and outreach programmes for the general public and for schools, which have increased capacity, education and employment in the field of weed biological control.

Eichhornia crassipes is notorious as the world’s worst aquatic weed, and here we present all aspects of its biology, ecology and invasion behaviour within the framework of the new series of Botany ...

Sagittaria platyphylla (Engelm.) J.G. Sm. (Alismataceae) is an emergent aquatic plant native to southern USA. Imported into Australia and South Africa as an ornamental and aquarium plant, the species is now a serious invader of shallow freshwater wetlands, slow-flowing rivers, irrigation channels, drains and along the margins of lakes and reservoirs. As a first step towards initiating a classical biological control program, a population genetic study was conducted to determine the prospects of finding compatible biological control agents and to refine the search for natural enemies to source populations with closest genetic match to Australian and South African genotypes. Using AFLP markers we surveyed genetic diversity and population genetic structure in 26 populations from the USA, 19 from Australia and 7 from South Africa. Interestingly, we have established that populations introduced into South Africa and to a lesser extent Australia have maintained substantial molecular genetic diversity comparable with that in the native range. Results from principal coordinates analysis, population graph theory and Bayesian-based clustering analysis all support the notion that introduced populations in Australia and South Africa were founded by multiple sources from the USA. Furthermore, the divergence of some Australian populations from the USA suggests that intraspecific hybridization between genetically distinct lineages from the native range may have occurred. The implications of these findings in relation to biological control are discussed.

As a result of a variety of chemical, environmental, mechanical and physiological difficulties, insects that spend their entire life spans in the marine or intertidal region are relatively rare. The present study assesses whether morphological and physiological adaptations have evolved in a maritime parasitoid wasp species Echthrodesis lamorali Masner, 1968 (Hymenoptera: Platygastridae, Scelioninae), in response to environmental pressures on its respiratory functioning. Scanning electron and light microscopy of whole and sectioned specimens show the presence of structure-retaining taenidia in the tracheal tubes, although there is an absence of other major adaptations associated with the trachea or spiracles. Histological sectioning reveals the presence of unusual sacs in the female metasoma whose role is unknown, although they are hypothesized to most likely be linked to ovipositor control. Respirometry experiments illustrate the formation of a plastron when submerged, with the longevity of the wasps being increased by quiescence. The critical thermal range of E. lamorali is shown to be large: from −1.1 °C ± 0.16 to 45.7 °C ± 0.26 (mean ± SE). Behavioural and physiological adaptations in E. lamorali appear to have evolved in response to exposure to the heterogeneous environmental conditions experienced within the intertidal zone.

The invasive freshwater snail Tarebia granifera (Lamarck, 1822) was first reported in South Africa in 1999 and it has become widespread across the country, with some evidence to suggest that it reduces benthic macroinvertebrate biodiversity. The current study aimed to identify the primary abiotic drivers behind abundance patterns of T. granifera, by comparing the current abundance of the snail in three different regions, and at three depths, of the highly modified Nseleni River in KwaZulu-Natal, South Africa. Tarebia granifera was well established throughout the Nseleni River system, with an overall preference for shallow waters and seasonal temporal patterns of abundance. Although it is uncertain what the ecological impacts of the snail in this system are, its high abundances suggest that it should be controlled where possible and prevented from invading other systems in the region.

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 discovery that cryptic species are more abundant than previously thought has implications for weed biological control, as there is a risk that cryptic species may be inadvertently released with consequences for the safety of the practice. A cryptic species of a biological control agent released for the control of the invasive alien macrophyte, water hyacinth, Eichhornia crassipes (C. Mart.) Solms. (Pontederiaceae), was recently discovered in South Africa. The two species were considered a single species prior to genetic analysis and interbreeding experiments. The original biological control agent retains the name Eccritotarsus catarinensis (Carvalho) (Heteroptera: Miridae) whereas the new species has been described as Eccritotarsus eichhorniae Henry. In this study, we compared the host specificity, efficacy, and thermal physiologies of the two species. The host specificity of the two species within the Pontederiaceae was very similar and both are safe for release in South Africa. Comparison of the per capita impact of the two species indicated that E. eichhorniae was the more damaging species but this is likely to be influenced by temperature, with E. catarinensis being more effective under lower temperatures and E. eichhorniae being more effective under higher temperatures. Releasing the correct species for the thermal environment of each release site will improve the level of control of water hyacinth in South Africa. This example highlights the need to keep populations of biological control agents from different native range collection localities separate, and to screen for host specificity and efficacy.

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.

Freshwater ecosystems are particularly susceptible to invasions by invasive non-native species (INNS) across a range of taxa, largely as a consequence of anthropogenic influences on these systems, with a number of ecological and socio-economic impacts. This chapter reviews freshwater invasive non-native species across the globe, focusing on fishes, invertebrates, floating macrophytes, and submerged macrophytes emphasising the knowledge gaps in particular that have resulted in biases inherent in assessments of freshwater invasions. These include an ecological bias because the majority of studies focus on terrestrial invasions; a geographical bias as most studies are focused on temperate northern hemisphere systems; and a taxon bias where fish invasions, populate the literature. This chapter highlights some of the approaches needed to survey, monitor, and manage INNS.

Nymphaea mexicana Zuccarini (Nympheaceae) (Mexican waterlily) is a rooted floating-leaved aquatic plant native to southern USA and Mexico that has become a problematic invasive alien plant in South Africa. Biological control is considered a desirable management strategy for the plant in South Africa. A good understanding of the genetic structure of invasive populations has been useful in other biological control programmes because taxonomic uncertainty about the target plant can result in natural enemies that are not adapted to the invasive populations being considered as potential agents. For N. mexicana, hybrids exist in the wild and horticultural trade, but identification is difficult, so understanding the genetic structure of populations is required to ensure that potential agents are collected off plants similar to invasive populations in South Africa. ISSR (inter-simple sequence repeats) analysis was used to determine whether invasive N. mexicana populations from South Africa were genetically similar to native range populations from USA or whether they were hybrids. Results from these analyses were matched with the morphotypes of each population based on petal colour, shape, and size. The genotypes suggested by the ISSR analyses corroborated the presence of both hybrid and pure forms of N. mexicana in South Africa. Populations of N. mexicana in the invaded range that are genetically similar to native range populations are more likely to be suitable for biological control, while other populations are likely to be hybrids formed by crossing of parents from the native range or within the horticultural trade, which may present difficulties for management using biocontrol.

There has been a rapid increase in atmospheric CO2 concentration, from pre-industrial values of 280 ppm to more than 400 ppm currently, and this is expected to double by the end of the 21st century. Studies have shown that plants grown at elevated CO2 concentrations have increased growth rates and invest more in carbon-based defences. This has important implications for the management of invasive alien plants, especially using biological control which is mostly dependent on herbivorous insects. The aim of this study was to investigate the effects of elevated atmospheric CO2 on the biological control of four invasive aquatic weeds (Azolla filiculoides, Salvinia molesta, Pistia stratiotes and Myriophyllum aquaticum). These species are currently under successful control by their respective biological control agents (Stenopelmus rufinasus, Cyrtobagous salviniae, Neohydronomus affinis, and Lysathia sp.) in South Africa. The plant species were grown in a two factorial design experiment, where atmospheric CO2 concentrations were set at ambient (400 ppm) or elevated (800 ppm), and plants were either subjected to or not subjected to herbivory by their target biological control agents. There was an overall increase in biomass production and C:N across all species at elevated CO2, both in the absence and presence of biological control, although C:N of M. aquaticum and biomass of A. filiculoides with herbivory were not constant with this trend. Insect feeding damage was reduced by elevated CO2, except for S. molesta. Thus, we can expect that plants will respond differently to CO2 increase, but the general trend suggests that these species will become more challenging to manage through biological control in future.

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.

Some of South Africa's citrus export markets require mandatory postharvest cold treatment of citrus fruit as a phytosanitary risk mitigation treatment for Thaumatotibia leucotreta (Meyrick) (Lepidoptera: Tortricidae). An alternative to this may be partial cold treatment as one of the final steps in a systems approach to mitigate phytosanitary risk. Consequently, the efficacy of such partial cold treatments was evaluated. It was first determined that a 2°C cold treatment was significantly more effective against fourth and fifth instars (the most cold-tolerant instars) than treatments at 3°C and 4°C for a duration of 18 d. Secondly, it was determined that 2°C for 18 d and 1°C for 16 d were similarly effective, but both treatments were significantly more effective than 1°C for 14 d. Mean mortality of fourth and fifth instars treated with 2°C for 18 d in seven replicates from four trials was 99.94%. Finally, it was determined that the inability of the majority of surviving larvae to develop to adulthood would further increase the efficacy of a 2°C for 18 d treatment to 99.96%. Inclusion of reproductive nonviability of survivors increased mortality to 99.99%.