Pet stores, aquarists and the internet trade as modes of introduction and spread of invasive macrophytes in South Africa

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

Plant–herbivore–parasitoid interactions in an experimental freshwater tritrophic system: higher trophic levels modify competitive interactions between invasive macrophytes

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

Impact of nutrients and herbivory by Eccritotarsus catarinensis on the biological control of water hyacinth, Eichhornia crassipes

Many water hyacinth infestations in South Africa are the symptom of eutrophication, and as a result, biological control of this weed is variable. This study examined the effects of herbivory by the mirid, Eccritotarsus catarinensis, on water hyacinth grown at high, medium and low nitrogen (N) and phosphorus (P) nutrient concentrations. Water nutrient concentration appears to be the overriding factor affecting plant growth parameters of water hyacinth plants—at high nutrient concentrations, leaf and daughter plant production were more than double than at low nutrient concentrations, while stem length was twice as great at high nutrient concentrations compared to low concentrations. Chlorophyll content was also twice as high at high nutrient concentrations than low concentrations. Conversely, flower production at high nutrient concentrations was less than half that at low concentrations. Herbivory by E. catarinensis did not have as great an effect on water hyacinth vigour as nutrient concentration did, although it significantly reduced the production of daughter plants by 23 ± 9%, the length of the second petiole by 13 ± 5%, and chlorophyll content of water hyacinth leaves by 15 ± 6%. In terms of insect numbers, mirids performed better on plants grown under medium nutrient conditions (99 ± 28 S.E.), compared to high nutrient concentrations (52 ± 27 S.E.), and low nutrient concentrations (25 ± 30 S.E.). Thus, these results suggest that the fastest and most significant reduction in water hyacinth proliferation would be reached by lowering the water nutrient concentrations, and herbivory by E. catarinensis alone is not sufficient to reduce all aspects of water hyacinth vigour, especially at very high nutrient concentrations

Risk assessment to interpret the physiological host range of Hydrellia egeriae, a biocontrol agent for Egeria densa

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

Love at first bite? Pre-release surveys reveal a novel association between a native weevil and the invasive Nymphaea mexicana Zuccarini (Nymphaeaceae) in South Africa

Classical biological control aims to suppress alien invasive plant populations by introducing host-specific natural enemies from the native range. This relies on the assumption that invasive plant populations in the invaded range benefit from the release of natural enemies. Pre-release surveys in the invaded range are a useful way to determine if enemy release applies to a particular invasive alien plant, and to determine what other factors may contribute to the invasion. Similarly, pre-release surveys gather information that can be used to compare invaded sites before and after the release of biological control agents and may also identify whether natural enemies have been accidentally introduced into the country. Pre-release surveys were conducted in South Africa on the invasive Nymphaea mexicana Zuccarini (Nymphaeaceae) to gather such information about this species, for which a biological control programme is being developed. There was lower diversity and abundance of herbivores in the native range compared to South Africa, suggesting that N. mexicana does experience enemy release at most sites in South Africa. This support for the enemy release hypothesis justifies the investment in biological control for its management. However, a native weevil, Bagous longulus Gyllenhal (Coleoptera: Curculionidae), was found feeding and reproducing on N. mexicana at three sites, resulting in damage to the leaves and suggesting that a novel association has formed between these species. Bagous longulus may have potential to be distributed to sites of N. mexicana where it is not present, though further investigation is necessary to confirm if its host range is suitable for this to be a safe endeavour. With the exception of sites where B. longulus was present, leaf sizes were large and damage was low, and there is no evidence that any natural enemies have been accidentally introduced from the native range. Findings such as these emphasise the importance of conducting thorough surveys during the development of biological control programmes

Contrasting effects of climate change on the invasion risk and biocontrol potential of the invasive Iris pseudacorus L. between Northern and Southern Hemisphere

Iris pseudacorus is both a prized ornamental and an invasive aquatic plant that tends to grow dense monospecific stands, displacing the local vegetation and altering the hydrology of freshwater ecosystems. Originally from Europe, this species has historically invaded North America, China and Japan, and more recently spread through Argentina, South Africa and Australasia, where it is now a target for biological control. Field surveys within its native range have led to the selection of three candidate biocontrol agents. Prioritizing the best candidates for different regions constitutes a critical step, which could save significant time and resources before further cost-intensive experimental studies are conducted. Climate change is seldom taken into consideration in the prioritization process. In this regard, climatic suitability can be used to model the potential distributions of weeds and their candidate agents, both in space and time, thus allowing to identify areas at risk of invasion and predict where agents will be able to establish long-term. Accordingly, the objectives of this work were (i) to predict I. pseudacorus invasions and range shifts in the context of climate change; (ii) to identify wetland areas most at risk of invasion under present and future climatic conditions; and (iii) to prioritize the best suite of candidate biocontrol agents for different invaded ranges, worldwide. To do so, we modelled the present and future (2040–2060) climatic suitability of I. pseudacorus and its candidate agents using the software MaxEnt. Our results highlight a clear distinction between predictions for the Northern and Southern Hemispheres. In North America and eastern Asia, the area climatically suitable for I. pseudacorus is expected to increase and shift northwards. As for its biocontrol agents, very low suitability is predicted across these regions, further decreasing under future climatic conditions. On the other hand, climatically suitable areas for the plant in South America, southern Africa and Australasia are predicted, on average, to reduce in response to climate change. A decrease in climatic suitability is also expected for its candidate biocontrol agents which, however, would still maintain a significant range overlap with their host. These results can be used to prioritize areas most at risk of invasion and identify which combination of candidates could potentially provide the best level of control across different invaded ranges

Climatic suitability and compatibility of the invasive Iris pseudacorus L.(Iridaceae) in the Southern Hemisphere: Considerations for biocontrol

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 programme 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 climatic 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 towards the 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

Risk assessment to interpret the physiological host range of Hydrellia egeriae, a biocontrol agent for Egeria densa

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

Morphological variations in southern African populations of Myriophyllum spicatum: Phenotypic plasticity or local adaptation?

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 is less than 0.001, Internode length: F(5,86) = 5.0747, P is less than 0.001, Leaf length: F(5,86) = 19.692, P is less than 0.001). Despite these differences in nutrient treatments, the growth pattern of each population remained true to the original starting point indicated by the lack of overlap between populations in the PCA groupings. This suggests that local adaptations are responsible for the differences in morphology between populations of M. spicatum, but shows that phenotypic plasticity does play a role as evidenced by individual responses to the different nutrient conditions. The development of these local adaptations within southern Africa suggests that the populations have had a long evolutionary history in the region and are relatively isolated with little reproductive mixing

Toxic effect of herbicides used for water hyacinth control on two insects released for its biological control in South Africa

The integrated control of water hyacinth, Eichhornia crassipes (Martius) Solms- Laubach (Pontederiaceae) has become necessary in South Africa, as biological control alone is perceived to be too slow in controlling the weed. In total, seven insect biological control agents have been released on water hyacinth in South Africa. At the same time, herbicides are applied by the water authorities in areas where the weed continues to be troublesome. This study investigated the assumption that the two control methods are compatible by testing the direct toxicity of a range of herbicide formulations and surfactants on two of the biological control agents released against water hyacinth, the weevil, Neochetina eichhorniae Warner (Coleoptera: Curculionidae) and the water hyacinth mirid, Eccritotarsus catarinensis (Carvahlo) (Hemiptera: Miridae). A number of the formulations used resulted in significant mortality of the mirid and the weevil. Products containing 2,4-D amine and diquat as active ingredients caused higher mortality of both agents (up to 80% for the mirid) than formulations containing glyphosate. Furthermore, when surfactants were added to enhance herbicide efficiency, it resulted in increased toxicity to the insects. We recommend that glyphosate formulations should be used in integrated control programmes, and that surfactants be avoided in order to reduce the toxic nature of spray formulations to the insect biological control agents released against water hyacinth.Water Research Commission of South Africa (Project 915/1/01).http://www.tandfonline.com/loi/cbst20hb201