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

Sample size assessments for thermal physiology studies: An R package and R Shiny application

Required sample sizes for a study need to be carefully assessed to account for logistics, cost, ethics and statistical rigour. For example, many studies have shown that methodological variations can impact the critical thermal limits (CTLs) recorded for a species, although studies on the impact of sample size on these measures are lacking. Here, we present ThermalSampleR; an R CRAN package and Shiny application that can assist researchers in determining when adequate sample sizes have been reached for their data. The method is particularly useful because it is not taxon specific. The Shiny application offers a user‐friendly interface equivalent to the package for users not familiar with R programming. ThermalSampleR is accompanied by an in‐built example dataset, which we use to guide the user through the workflow with a fully worked tutorial.Funder: National Research Foundation; doi: http://dx.doi.org/10.13039/501100001321 Funder: South African Research Chairs Initiative of the Department of Science and Technology Funder: Working for Water (WfW) programme of the Department of Environmental Affairs: Natural Resource Management programme (DEA: NRM

The establishment and rapid spread of Sagittaria Platyphylla in South Africa:

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’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

Biological control of South African plants that are invasive elsewhere in the world: A review of earlier and current programmes

South Africa supports a rich floral diversity, with 21 643 native plant taxa that include a high proportion (76.3%) of endemic species, and many of these favoured as ornamentals, both locally and globally. Consequently, South Africa has contributed substantially to global plant invasions, with 1093 native taxa (5% of all species) naturalized in other countries. At least 80 taxa are invasive in natural or semi-natural ecosystems elsewhere, while an additional 132 taxa are potentially invasive. Of the global naturalized flora, 8.2% originate from South Africa and largely comprise species of Poaceae, Asteraceae, Iridaceae and Fabaceae. Australia, in particular, but also Europe and North America are major recipients of South African weeds. However, few countries have targeted South African plants for biological control (biocontrol), with most efforts undertaken by Australia. Previous and current targets have involved only 26 species with 17 agents (15 insects, one mite and one rust fungus) of South African origin released on five target species in Australia and the United States of America. South Africa’s history of weed biocontrol, together with a large cohort of active scientists, is currently facilitating several internationally funded programmes targeting invasive plants of South African origin. In particular, the recently inaugurated Centre for Biological Control at Rhodes University and the University of KwaZulu-Natal have provided the impetus for novel efforts on five new target species and renewed efforts on four previously targeted species. In this contribution, we review the history of earlier biocontrol programmes against weeds of South African origin and the status of projects currently in progress in South Africa

Preventing and controlling nonnative species invasions to bend the curve of global freshwater biodiversity loss

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. Thesemanagement 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 non-species 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 review of the biocontrol programmes against aquatic weeds in South Africa

Biological control (biocontrol) against invasive macrophytes is one of the longest standing programmes in South Africa, initiated in the 1970s against water hyacinth, Pontederia crassipes Mart. (Pontederiaceae). Since then, 15 agent species (13 insects, one mite and one pathogen) have been released against six weeds, most of which are floating macrophytes, with excellent levels of success. The release of the water hyacinth planthopper Megamelus scutellaris Berg (Hemiptera: Delphacidae) in particular, has improved biocontrol prospects for water hyacinth since 2018. In the last decade, however, a new suite of submerged and rooted emergent invasive macrophytes has been targeted. The first release against a submerged macrophyte in South Africa, and the first release against Brazilian waterweed, Egeria densa Planch. (Hydrocharitaceae), anywhere in the world, was achieved with the release of a leafmining fly, Hydrellia egeriae Rodrigues-Júnior, Mathis and Hauser (Diptera: Ephydridae). Yellow flag, Iris pseudacorus L. (Iridaceae) and Mexican waterlily, Nymphaea mexicana Zucc. (Nymphaeaceae), have also been targeted for biocontrol for the first time worldwide, and are in the early stages of agent development. Post-release evaluations, long term monitoring and controlled experiments have highlighted the need for a more holistic approach to managing aquatic invasive plants in South Africa, whose presence is largely driven by eutrophication, resulting in regime shifts between floating and submerged invaded states