Julie A. Coetzee

The role of eutrophication in the biological control of water hyacinth, Eichhornia crassipes , in South Africa

South Africa has some of the most eutrophic aquatic systems in the world, as a result of the adoption of an unnecessarily high 1xa0mgxa0l−1 phosphorus (P) standard for all water treatment works in the 1970xa0s. The floating aquatic macrophyte, water hyacinth (Eichhornia crassipes (Mart.) Solms (Pontederiaceae)), has taken advantage of these nutrient rich systems, becoming highly invasive and damaging. Despite the implementation of a biological control programme in South Africa, water hyacinth remains the worst aquatic weed. A meta-analysis of published and unpublished laboratory studies that investigated the combined effect of P and nitrogen (N) water nutrient concentration and control agent herbivory showed that water nutrient status was more important than herbivory in water hyacinth growth. Analysis of long-term field data collected monthly from 14 sites around South Africa between 2004 and 2005 supported these findings. Therefore the first step in any water hyacinth control programme should be to reduce the nutrient status of the water body.

Potential spread of the invasive plant Hydrilla verticillata in South Africa based on anthropogenic spread and climate suitability

Hydrilla verticillata (L.f.) Royle is a submerged aquatic plant native to Asia and Australia that is highly invasive in the USA and was first recorded in South Africa in 2006. It is only known from one locality, Pongolapoort Dam in KwaZulu-Natal Province, but there are fears that it might spread to other sites. The primary vector of spread in the USA is recreational boaters and anglers. A survey at a fishing competition on Pongolapoort Dam showed that anglers travel considerable distances around South Africa (73% of water bodies were >200xa0km, visited by 68% of the respondents). A Threat Index for freshwater bodies throughout South Africa visited by participants of the competition was calculated showing that dams in the vicinity of the infestation were more at risk from invasion. Further, the potential distribution of the weed based on climatic matching with the region of origin showed that most of the country was suitable for establishment, with the exception of the high-lying interior of the country. Recommendations for reducing the potential spread of hydrilla in South Africa are presented.

Prospects for the Biological Control of Submerged Macrophytes in South Africa

Historically, biological control efforts against aquatic plants in South Africa have focused on floating species, and as a result, there has been a dearth of research into the invasion and control of submerged macrophytes. With numerous submerged invasive species already established in South Africa, thriving horticultural and aquarium industries, nutrient-rich water systems, and a limited knowledge of the drivers of invasions of submerged macrophytes, South Africa is highly vulnerable to a second phase of aquatic plant problems. Experience gained in the U.S.A. on biological control against submerged weeds, such as hydrilla, Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) and spiked/Eurasian watermilfoil, Myriophyllum spicatum L. (Haloragaceae), have provided South African researchers with the necessary foundation to initiate programmes against these weeds. Research in South Africa is currently focused on pre-release studies on the biological control of H. verticillata, using an undescribed fly, Hydrellia sp. (Diptera: Ephydridae) and a weevil, Bagous hydrillae OBrien (Coleoptera: Curculionidae); and on M. spicatum using a North American weevil, Euhrychiopsis lecontei Dietz (Coleoptera: Curculionidae). Feasibility studies into biological control of some incipient submerged weeds are also being conducted, including Brazilian water weed, Egeria densa Planch. (Hydrocharitaceae), Canadian water weed, Elodea canadensis Mitch. (Hydrocharitaceae) and cabomba, Cabomba caroliniana A.Gray (Cabombaceae). Progress with, and potential constraints that may limit these programmes, are discussed.

Comparisons of the thermal physiology of water hyacinth biological control agents: predicting establishment and distribution pre- and post-release

Investigations into the thermal physiology of weed biological control agents may elucidate reasons for establishment failure following release. Such studies have shown that the success of water hyacinth biological control in South Africa remains variable in the high‐lying interior Highveld region, because the control agents are restricted to establishment and development due to extreme winter conditions. To determine the importance of thermal physiology studies, both pre‐ and post‐release, this study compared the known thermal requirements of Eccritotarsus catarinensis (Carvalho) (Hemiptera: Miridae) released in 1996, with those of an agent released in 1990, Niphograpta albiguttalis (Warren) (Lepidoptera: Pyralidae) and a candidate agent, Megamelus scutellaris Berg (Hemiptera: Delphacidae), which is currently under consideration for release. The lower developmental threshold (to) and rate of development (K) were determined for N. albiguttalis and M. scutellaris, using a reduced axis regression, and incorporated into a degree‐day model which compared the number of generations that E. catarinensis, N. albiguttalis, and M. scutellaris are capable of producing annually at any given site in South Africa. The degree‐day models predicted that N. albiguttalis (K = 439.43, to = 9.866) can complete 4–11 generations per year, whereas M. scutellaris (K =502.96, to = 11.458) can only complete 0–10 generations per year, compared with E. catarinensis (K = 342, to = 10.3) which is predicted to complete 3–14 generations per year. This suggests that the candidate agent, M. scutellaris, will not fare better in establishment than the other two agents that have been released in the Highveld, and that it may not be worth releasing an agent with higher thermal requirements than the agents that already occur in these high‐lying areas. Thermal physiology studies conducted prior to release are important tools in biological control programmes, particularly those in resource‐limited countries, to prevent wasting efforts in getting an agent established.

Natural enemies from South Africa for biological control of Lagarosiphon major (Ridl.) Moss ex Wager (Hydrocharitaceae) in Europe

The non-native invasive plant, Lagarosiphon major (Hydrocharitaceae) is a submersed aquatic macrophyte that poses a significant threat to water bodies in Europe. Dense infestations prove difficult to manage using traditional methods. In order to initiate a biocontrol programme, a survey for natural enemies of Lagarosiphon was conducted in South Africa. Several phytophagous species were recorded for the first time, with at least three showing notable promise as candidate agents. Amongst these, a leaf-mining fly, Hydrellia sp. (Ephydridae) that occurred over a wide distribution causes significant leaf damage despite high levels of parasitism by braconid wasps. Another yet unidentified fly was recorded mining the stem of L. major. Two leaf-feeding and shoot boring weevils, cf. Bagous sp. (Curculionidae) were recorded damaging the shoot tips and stunting the growth of the stem. Several leaf-feeding lepidopteran species (Nymphulinae) were frequently recorded, but are expected to feed on a wide range of plant species and are not considered for importation before other candidates are assessed. The discovery of several natural enemies in the country of origin improves the biological control prospects of L. major in Europe.

Water hyacinth, Eichhornia crassipes (Pontederiaceae), reduces benthic macroinvertebrate diversity in a protected subtropical lake in South Africa

The socio-economic impacts of the free-floating aquatic plant water hyacinth, Eichhornia crassipes (Pontederiaceae), on aquatic systems are well documented, yet the impacts on aquatic biodiversity, particularly invertebrate biodiversity, are less well understood. This study aimed to determine whether the presence of water hyacinth altered the diversity and assemblage structure of benthic macroinvertebrates in a conservation area. The benthic macroinvertebrate assemblage was sampled over 1xa0year at five sites under water hyacinth mats and at five sites without water hyacinth at Lake Nsezi—Nseleni River in the vicinity of Richards Bay, KwaZulu-Natal, South Africa. Artificial substrates were placed beneath water hyacinth mats or in the open water to allow for colonization by freshwater macroinvertebrates, and left for a period of 6xa0weeks, repeated on seven occasions. Twenty nine families comprising 18,797 individuals were collected, 817 (13 families) individuals were from under water hyacinth mat sites compared to 17,980 (27 families) individuals from open water sites. Ninety-eight percent of individuals collected were, however, the invasive snail, Tarebia granifera. Open water samples were separated from samples beneath the water hyacinth mat by non-metric Multidimensional Scaling, indicating reduced biodiversity associated with the presence of water hyacinth. Exclusion of the dominant Thiaridae from the analysis did not alter the groupings. Family richness(s) and abundance (N) were significantly higher in open water communities(S: H3xa0=xa021.09; Pxa0=xa00.0001; N: H3xa0=xa022.58; Pxa0=xa00.00001), while evenness (J’) was higher under water hyacinth mats (H3xa0=xa020.13; Pxa0=xa00.0002). The presence of water hyacinth had a significantly negative impact on aquatic macroinvertebrate biodiversity in a conservation area, and therefore the control of this invasive aquatic plant must play a major role in catchment management.

Two in one: cryptic species discovered in biological control agent populations using molecular data and crossbreeding experiments

Abstract 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.

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

Abstract 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 (Carvalho) (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.

Unraveling the biogeographic origins of the Eurasian watermilfoil (Myriophyllum spicatum) invasion in North America

PREMISE OF THE STUDYnUsing 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.nnnMETHODSnDNA 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.nnnKEY RESULTSnThe 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.nnnCONCLUSIONSnThe 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.

Distributional Range of the South African Maritime Spider-Egg Parasitoid Wasp, Echthrodesis lamorali (Hymenoptera: Platygastridae: Scelioninae)

ABSTRACT The southern African coastline plays host to nine spider species. Two of these, namely Desis formidabilis (O. P.-Cambridge, 1890) (Araneae: Desidae) and Amaurobioides africanus Hewitt, 1917 (Araneae: Anyphaenidae), are recorded as hosts for an intertidal spider egg parasitoid, Echthrodesis lamorali Masner, 1968 (Hymenoptera: Platygastridae: Scelioninae). These two spider species occur from Lüderitz (Namibia) along the coast to East London (Eastern Cape Province, South Africa), while their parasitoid has been known from only a single locality on the Cape Peninsula. The South African coastline was surveyed from Jacobsbaai (Western Cape Province) to East London in an attempt to determine the full distribution of E. lamorali. The wasp was only reared from host eggs collected on the Cape Peninsula, confirming a high degree of endemism for this species.