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

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

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

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

Chlorophyll fluorometry as a method of determining the effectiveness of a biological control agent in post-release evaluations

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 different eutrophic nutrient treatments and agent densities (high and low). The results indicated that plants grown in low nutrients with high densities of M. scutellaris showed the greatest reduction in the fluorescence parameters Fv/Fm and PIabs. The successful use of chlorophyll fluorometry for the detection of subtle insect damage to water hyacinth leaves could have future application in post-release studies to measure the impact of M. scutellaris in the field

First record of an indigenous South African parasitoid wasp on an imported biological control agent, the water hyacinth hopper

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 world including South Africa. Megamelus scutellaris Berg (Hemiptera: Delphacidae) was released as a biological control agent and has established in several regions. Recently, the indigenous species Echthrodelphax migratorius Benoit, (Hymenoptera: Dryinidae) was discovered in South Africa parasitising M. scutellaris. This newly discovered relationship might have repercussions for the efficacy of biological control of water hyacinth by the delphacid. The wasp may negatively impact M. scutellaris populations making it difficult for the agent to successfully manage the invasive weed. Contrarily, the parasitoid may be beneficial by keeping the M. scutellaris populations stable, serving as a natural enemy

Testing the thermal limits of Eccritotarsus catarinensis: a case of thermal plasticity

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

Biological control of Salvinia molesta in South Africa revisited

The aquatic weed Salvinia molesta D.S. Mitch. (Salviniaceae) was first recorded in South Africa in the early 1900s, and by the 1960s was regarded as one of South Africa’s worst aquatic weeds. Following the release of the weevil, Cyrtobagous salviniae Calder and Sands (Coleoptera: Curculionidae) in 1985, the weed is now considered under successful biological control. However, the post-release evaluation of this biological control programme has been ad hoc, therefore, to assess the efficacy of the agent, annual quantitative surveys of South African freshwater systems have been undertaken since 2008. Over the last ten years, of the 57 S. molesta sites visited annually in South Africa, the weevil has established at all of them. Eighteen sites are under successful biological control, where the weed no longer poses a threat to the system and 19 are under substantial biological control, where biological control has reduced the impact of the weed. Since 2008, the average percentage weed cover at sites has declined significantly from 51–100% cover to 0–5% cover in 2017 (R2 = 0.78; P < 0.05). Observations of site-specific characteristics suggest that biological control is most effective at small sites and more difficult at larger and shaded sites. Our findings show that S. molesta remains under good biological control in South Africa, however, some sites require intermittent strategic management, such as augmentative releases of C. salviniae