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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.

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.

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.

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.

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.

Sagittaria platyphylla Engelm. (Alismataceae) is a freshwater aquatic macrophyte that has become an important invasive weed in freshwater systems in South Africa, New Zealand, Australia, and recently China. In South Africa, due to its rapid increase in distribution and ineffective control options, it is recognised as one of the country&rsquo, s worst invasive aquatic alien plants. In this paper, we investigate the spread of the plant since its first detection in 2008, and the management strategies currently carried out against it. Despite early detection and rapid response programmes, which included chemical and mechanical control measures, the plant was able to spread both within and between sites, increasing from just one site in 2008 to 72 by 2019. Once introduced into a lotic system, the plant was able to spread rapidly, in some cases up to 120 km within 6 years, with an average of 10 km per year. The plant was successfully extirpated at some sites, however, due to the failure of chemical and mechanical control, biological control is currently being considered as a potential control option.

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.

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.

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%.

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.

Classical biological control for the management of floating invasive plants has been highly successful in South Africa. However, restoring ecosystem services has been compromised by a new suite of submerged invasive plants. This study proposes that biological control of floating invasive macrophytes acts as a catalyst in a regime shift between floating and submerged invasive plant dominance. Regime shifts are large and sudden changes in the structure and functioning of ecosystems. The proposed shift is driven by the rapid decomposition of floating plants and subsequent increase in availability of nutrients and light. A mesocosm experiment explored the effect of biological control on floating Pistia stratiotes L. (Araceae) upon the growth of invasive submerged Egeria densa Planch. (Hydrocharitaceae), and native submerged plant species of the same family; Lagarosiphon major (Ridl.) Moss (Hydrocharitaceae). The results revealed a cascade effect of biological control of P. stratiotes on the availability of nitrogen, resulting in increased relative growth rates and invasive capacity for E. densa. In contrast, the native L. major could not compete with healthy or damaged P. stratiotes. These findings highlight the vulnerability of South African freshwater systems to submerged plant invasions and demonstrate the importance of a more holistic approach to invasive plant management.

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.

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.

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.