Feeding impact of the planthopper Taosa longula (Hemiptera: Dictyopharidae) on water hyacinth,Eichhornia crassipes (Pontederiaceae)

Taosa longula Remes Lenicov (Hemiptera: Dictyopharidae), a planthopper native to South America, is a candidate for the biological control of water hyacinth, Eichhornia crassipes (Mart.) Solms-Laubach (Pontederiaceae), a serious weed worldwide. Biological control requires agents that are not only specific but also effective. Damage caused by sap-sucking insects is difficult to assess. In this work we designed an experimental and analytical procedure to evaluate the potential damage of T. longula on water hyacinth. The damage that T. longula causes to the clonal reproduction, biomass production, and growth of water hyacinth was studied through a paired greenhouse trial with floating cages. The performance of the plant, starting from two plants per treatment, was evaluated at different insect densities (5, 10, 15 and 20 nymphs per cage) until all the nymphs moulted to adults. The tests showed that individual growth and biomass production of water hyacinth was reduced due to the effect of the insect feeding above five nymphs per cage. The number of new plants produced by clonal reproduction was only significantly different above 15 nymphs per cage. These results suggest that this planthopper could be an effective agent for the biological control of water hyacinth.     

Assessing the risk of Eccritotarsus eichhorniae to pickerelweed,Pontederia cordata in North America

The mirid Eccritotarsus eichhorniae is a newly described species with potential as a biological control agent for water hyacinth, Eichhornia crassipes, in North America. A series of quarantine-based studies quantified E. eichhorniae usage of Pontederia cordata with no-choice, adult choice, nymph choice, multi-generation, and impact tests. Although greater numbers of E. eichhorniae adults developed on E. crassipes in two separate no-choice trials, some E. eichhorniae were able to complete development on P. cordata in both trials. Adults showed a clear oviposition preference for E. crassipes and F₁ progeny reduced the mean relative growth rate of E. crassipes but not P. cordata. Naïve nymphs were more likely to leave P. cordata for E. crassipes but not all nymphs left P. cordata when given a choice. Both multi-generation trials showed a rapid increase in the density of adults and nymphs on E. crassipes over four to five generations while population densities on P. cordata did not change, indicating that this insect can maintain populations on this plant. The modest damage inflicted by a sibling species on E. crassipes, coupled with the ability of E. eichhorniae to persist on P. cordata, may pose an interpretation challenge to U.S. regulatory agencies concerned with non-target impacts by biological control agents. Another option for the continuing efforts to increase the biological control of E. crassipes is to prioritise the evaluation of other potential agents like Taosa longula Remes Lenicov (Homoptera: Dictopharidae) or Thrypticus truncatus Bickel and Hernández (Diptera: Dolichopodidae) over E. eichhorniae.     

Feeding behavior and spatial distribution of two planthoppers,Megamelus scutellaris (Delphacidae) and Taosa longula (Dictyopharidae), on water hyacinth

Megamelus scutellaris Berg (Delphacidae) and Taosa (Cuernavaca) longula Remes Lenicov (Dictyopharidae) are specialist planthoppers that feed and reproduce on the invasive aquatic weed, Eichhornia crassipes (Martius) Solms-Laubach (Pontederiaceae). They overlap geographically in several regions of South America and may, therefore, interact and compete for food and microhabitat. Preliminary observations indicated that both species do not feed on the same part of the plant. We hypothesized that they partition the resource; hence, we studied (1) the feeding mechanism at the tissue level and (2) the spatial distribution of both species on the water hyacinth plant. Salivary sheaths were detected through histological sections of plant tissues using light microscopy. The location of either planthopper species on the plant was recorded when in the presence or absence of the other species. Both species produced true salivary sheaths, mostly branched (M. scutellaris: 82%; T. longula: 84%), ending in phloem (M. scutellaris: 56%; T. longula: 52%), and xylem tissues (M. scutellaris: 24%; T. longula: 28%). They resided on different parts of the water hyacinth plant even when they did not coexist; nymphs of T. longula occurred primarily on the back side of the leaf laminas, while nymphs of M. scutellaris occupied the basal zone of the petioles. This study shows that these planthoppers complement each other and could be used in combination as control agents for water hyacinth. Further experimental studies and field observations are necessary to quantify interactions.     

Field efficacy and predicted host range of the pickerelweed borer, Bellura densa, a potential biological control agent of water hyacinth

The North American noctuidmoth Bellura densa offers promise as abiological control agent for use in Africa andother countries invaded by water hyacinth. Anaugmentative release at a pond in Florida, USA,eliminated water hyacinth within a few months. Laboratory studies, though, indicated thatoviposition was indiscriminate and thatdevelopment was completed on taro (Colocasia esculenta [Araceae]) as well as onseveral Pontederiaceae. Acceptability of taroas a larval food plant was confirmed in thefield when larvae were found in isolated standsof taro in Florida. Evidence of use of Peltandra virginica (Linnaeus) (Araceae) wasnoted at another site. The distribution oflarval damage was compared at a site containinga mixture of 97% taro and 3% pickerelweed(Pontederia cordata). Larvae damaged87% of the pickerelweed compared to only about5% of the taro, suggesting spillover ontotaro. In another study, 416 larvae wereliberated into a concrete tank containing waterhyacinth (818 plants) surrounded by taro (96plants). Three months later, taro accountedfor only 4% of the damaged plants, less thanthe 11% expected if host selection had beenrandom. In a similar study, larvae wereliberated onto water hyacinth in a large tankdivided into thirds, with pickerelweed or taroat either end and water hyacinth in the middle. The distributions of F₁ egg masses andincidence of damage 3 months later indicatedthat pickerelweed was preferred over taro, but26% of the taro plants were damaged. Weconclude that while B. densa prefersplants in the Pontederiaceae, it is notrestricted to this plant family. Plants in theAraceae would be at risk if this insect werereleased outside of North America, particularlyin cropping situations near water hyacinthinfestations. Bellura densa could beuseful for water hyacinth management in theU.S. if effective augmentation strategies weredeveloped.     

Life History and Laboratory Host Range ofEccritotarsus catarinensis(Carvalho) (Heteroptera: Miridae), a New Natural Enemy Released on Water Hyacinth (Eichhornia crassipes(Mart.) Solms-Laub.) (Pontederiaceae) in South Africa

A mirid,Eccritotarsus catarinensis(Carvalho), was studied as a potentially damaging natural enemy for water hyacinth, (Eichhornia crassipes(Mart.) Solms-Laub.), in South Africa. In the laboratory, eggs were inserted into the leaf tissue parallel to the leaf surface. The four nymphal instars fed gregariously with the adults mainly on the undersurface of the leaves, causing severe chlorosis at high population levels. The duration of immature stages (egg and nymphs) was approximately 23 days, while the adults survived for approximately 50 days. Favorable biological characteristics ofE. catarinensisincluded a high rate of increase, gregarious habits, long-lived and mobile adults, and several generations per year. Laboratory host range of the mirid was determined by adult choice trials on 67 plant species in 36 families and adult no-choice trials on five species in the Pontederiaceae. Feeding was recorded on all Pontederiaceae tested and oviposition on four of the five species. However, these plant species proved to be inferior hosts forE. catarinensisin comparison to water hyacinth, suggesting thatE. catarinensiswould be an acceptable natural enemy for water hyacinth in South Africa.     

Megamelus scutellaris (Hemiptera: Delphacidae), a biocontrol agent of water hyacinth,is not sufficiently specific for release in Australia

Water hyacinth Eichhornia crassipes (Pontederiaceae) is one of the world's worst invasive species, responsible for damaging aquatic systems in many warmer parts of the globe including north America, Africa, Asia and Australia. The planthopper Megamelus scutellaris Berg (Delphacidae) has been released in USA and approved for release in South Africa for biocontrol of water hyacinth. We assessed this agent for suitability for release in Australia and found that a related native aquatic plant, Monochoria cyanea (Pontederiaceae) is within the fundamental host range of this insect. Adult survival, oviposition and development of nymphs to adult was equally high on M. cyanea as on the target species, although the quality of these next generation adults was lower than those reared on the target species. This demonstrates that M. scutellaris is not sufficiently specific for release in Australia. Nymphal development to adults occurred only in very low numbers on the three other Australian species of Monochoria. M. cyanea only occurs in Australia so M. scutellaris is still a possible water hyacinth biocontrol candidate for other regions depending on the results of assessment of the risk to local species of Monochoria. This study demonstrates the effectiveness of modern biocontrol agent assessment and reinforces the importance of testing of local non-target species.     

Should the mirid,Eccritotarsus catarinensis (Heteroptera: Miridae), be considered for release against water hyacinth in the United States of America?

Between one and seven biological control agents have been released against water hyacinth (Eichhornia crassipes (Mart.) Solms) in at least 30 countries, with varied success. A mirid, Eccritotarsus catarinensis (Carvalho) (Heteroptera: Miridae), the most recent agent released, is damaging to the plant on the African continent. It could be useful in the USA where water hyacinth remains a problem, but its introduction remains in doubt because during host specificity trials, it developed on Pontederia cordata L. (pickerelweed), indigenous to the USA. However, it did not establish on pickerelweed monocultures during South African field trials, and only light spillover feeding occurred where the two plants coexisted suggesting that the use of P. cordata as a host is a laboratory artefact and it may be suitable for use in the USA, if its thermal physiology allows establishment. We reran models developed for South Africa using CLIMEX to predict whether the mirid will establish where water hyacinth and pickerelweed co-occur, but not where pickerelweed occurs in the absence of water hyacinth. The models suggest that the mirid's distribution will be limited by cold winter temperatures and insufficient thermal accumulation to the southern states of the USA, within the main distribution of water hyacinth. Even though some spillover feeding on pickerelweed might result where the two plants co-occur, the risk of population level effects seems minimal and the risk to more northern pickerelweed negligible. The benefits, including improved habitat for pickerelweed, associated with further suppression of water hyacinth, outweigh the minimal risk of collateral damage to pickerelweed.     

Meteorological weather station data can be used in climate matching studies of biological control agents

Meteorological weather station data are often used in climate matching studies to predict potential distributions of biological control agents, yet, this does not take into account the effects of microclimates experienced by the agents. Comparisons of the number of generations that the mirid, Eccritotarsus catarinensis, a biological control agent of water hyacinth, Eichhornia crassipes (Mart.) Solms-Laub (Pontederiaceae), was predicted to complete using meteorological weather station data, on site air temperature and water hyacinth canopy microclimate temperatures recorded over two years showed that there were no significant differences between the temperature data sources. Therefore, meteorological weather station data used in degree-day models of biological control agents are useful in explaining broad establishment patterns.     

Variation in expression of trimorphic incompatibility in Pontederia cordata L. (Pontederiaceae)

Summary Pontederia cordata L. (Pontederiaceae), a perennial diploid, possesses the rare genetic polymorphism tristyly. A controlled pollination programme was conducted over a three year period, under glasshouse conditions, on 36 clones of P. cordata var. cordata to examine the nature of the self-incompatibility system. The three major findings of the pollination study were: (1) the three floral morphs display different levels of self-incompatibility, (2) pollen from the two anther levels within a flower exhibits different compatibility behaviour in self-pollinations, (3) considerable individual genetic variation in the expression of self-incompatibility is evident among clones within floral morphs. Similar results were also obtained from a smaller study on 15 clones of P. cordata var. lancifolia conducted over a 6 month period. In common with other Pontederia species the mid-styled morph (M) of P. cordata produces large amounts of seed when self-pollinated with pollen from long-level anthers. A developmental model is proposed to explain the high level of self-compatibility of the M morph in Pontederia species. Self-pollination of segregating progenies from M and S morphs of known incompatibility status demonstrated that the expression of incompatibility is closely associated with style length. It is suggested that overall differences in incompatibility behaviour among the floral morphs may be due to the pleiotropic effects of major genes controlling sub-characters of the tristylous syndrome, rather than linked modifier genes. However, the variable expression of trimorphic incompatibility within floral morphs suggests that this variation may be polygenic in origin.     

The role of eutrophication in the biological control of water hyacinth, <Emphasis Type="Italic">Eichhornia crassipes</Emphasis>, 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 1 mg l⁻¹ phosphorus (P) standard for all water treatment works in the 1970 s. 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.     

Cryptic species of a water hyacinth biological control agent revealed in South Africa: host specificity,impact, and thermal tolerance

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 Host Range ofEccritotarsus catarinensis(Heteroptera: Miridae), a Potential Agent for the Biological Control of Waterhyacinth (Eichhornia crassipes)

The host range ofEccritotarsus catarinensiswas determined using 33 plant species to assess the risk of using this insect, a native of South America, for the classical biological control of waterhyacinth (Eichhornia crassipes) in Australia and Papua New Guinea. The results, in conjunction with the results of Hillet al.(1999) who tested 67 species (mostly South African), strongly suggest thatE. catarinensisis restricted to the Pontederiaceae, a family of aquatic plants. All five species of Pontederiaceae in the Australian testing,E. crassipes, Pontederia cordata, Monochoria vaginalis, M. cyanea,andM. australasica,were suitable for insect development. Colonies persisted for at least four generations onE. crassipes, P. cordata,andM. vaginalis.Two-way choice and multiple-choice preference trials were conducted and discussed.E. catarinensisdid not exhibit a clear preference for waterhyacinth over other Pontederiaceae in these trials. Most oviposition occurred into the upper surface of the lamina during laboratory testing despite observations that the underside was preferred in the field. Although not considered suitable for release in Australia, this insect may be useful in other countries where more serious waterhyacinth problems occur and whereM. vaginalisis a serious weed, such as in Southeast Asia.     

Fungi associated with Eichhornia crassipes in South Africa and their pathogenicity under controlled conditions

Eichhornia crassipes Mart. Solms-Laubach (Pontederiaceae), water hyacinth, continues to be the world's worst aquatic weed. In South Africa, considerable research has been conducted on biological control agents associated with water hyacinth, with the release of six arthropods and one fungus, but little is known about the occurrence and impacts of native phytopathogenic fungi. Nation-wide surveys were conducted in 2010 and 2011 on various aquatic bodies of South Africa to identify the fungal pathogens associated with water hyacinth. Diseased plant parts were collected and fungi were isolated and identified. Some 250 isolates belonging to more than 25 genera were collected. Some of these represent new host records, as well as undescribed taxa. Isolates of Acremonium zonatum (Sawada) Gams, Alternaria eichhorniae Nag Raj and Ponnappa, Bipolaris hawaiiensis (M.B. Ellis) Uchida and Aragaki, Fusarium Link, Myrothecium roridum Tode ex Fr. and Ulocladium sp., showed the highest pathogenicity and have the potential to be useful in complementing the ongoing biocontrol programme on water hyacinth in South Africa.     

Predicting the distribution of Eccritotarsus catarinensis, a natural enemy released on water hyacinth in South Africa

Water hyacinth [Eichhornia crassipes (Mart.) Solms (Pontederiaceae)] is the most damaging aquatic weed in South Africa, where five arthropod biological control agents have been released against it. The most recent introduction of Eccritotarsus catarinensis (Carvalho) (Heteroptera: Miridae) has failed to establish permanent populations at a number of sites in South Africa where water hyacinth is a problem. Cold winter temperatures at these sites are assumed to be the reason for these establishment failures. This assumption was tested by investigating the thermal physiology of the mirid, then incorporating these data into various predictive distribution models. Degree‐day models predict 3–14 generations per year at different localities in South Africa, and five generations at a Johannesburg site where the mirid failed to overwinter. The inability to develop sufficiently rapidly during winter months may hinder overwintering of this insect, which was predicted to develop through only one generation during the winter months of April to August in Johannesburg. A CLIMEX model also showed that cold stress limits the mirid's ability to overwinter in the interior of the country, while determination of the lower lethal limit (–3.5 °C) and critical thermal minimum (1.2 ± 1.17 °C) also indicated that extreme temperatures will limit establishment at certain sites. It is concluded that E. catarinensis is limited in its distribution in South Africa by low winter temperatures.     

A Leaf Spot of Water Hyacinth Caused by Drechslera spicifera

In the present study, corn meal agar was found as the best medium for growth of Drechslera spicifera, a pathogen of water hyacinth. Small water hyacinth plants exhibited a smaller number of lesions/leaf when inoculated with D. spicifera compared with inoculated medium and large plants. However, percentage total diseased area/leaf was almost the same among the different plant sizes. The pathogen did not interfere with the vegetative plant growth, since, both infected and healthy water hyacinth plants of the same age showed similar percentages of new leaf growth. Leaf diffusates of water hyacinth plants significantly reduced conidiospore germination in D. spicifera.     

Macrolophus caliginosus: Functional response to whiteflies and preference and switching capacity between whiteflies and spider mites

Laboratory experiments were performed with adult female Macrolophus caliginosus Wagner (Heteroptera: Miridae) at 22ºC on bean plants to determine the functional response towards whiteflies, as well as the preference and switching capacity between the two prey species: whiteflies and spider mites. Predation of females presented with first instars of Trialeurodes vaporariorum Westwood (Homoptera: Aleyrodidae) was of a Type III functional response. The observed maximum predation was approximately 75 first instars at high prey densities within a 24-h period. The preference of M. caliginosus females between eggs of T. vaporariorum and Tetranychus urticaeKoch (Acarina: Tetranychidae) changed with the ratio of offered prey. The preference for T. vaporariorum eggs increased non-linearly with increasing proportions of this prey type. The average maximum predation of whitefly and spider mite eggs were approximately 166 and 111 eggs per day, respectively, at the highest ratio of the two preys. The proportion of M. caliginosus females found on the test plants at the end of the experiment increased with prey density suggesting that this mirid spends more time in areas with high prey density. Macrolophus caliginosus females are voracious predators of eggs and first instars of T. vaporariorum as well as of spider mite eggs and may thus be a valuable addition to existing methods of biological control of T. vaporariorum and T. urticae.     

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 (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.     

Predation of the apefly,Spalgis epius (Lepidoptera: Lycaenidae) on citrus mealybug,Planococcus citri (Hemiptera: Pseudococcidae)

The citrus mealybug, Planococcus citri (Risso) (Hemiptera: Pseudococcidae) is a serious pest of economically important crops worldwide. The apefly, Spalgis epius (Westwood) (Lepidoptera: Lycaenidae) is a potential predator of various species of mealybugs. Earlier investigation on its daily preying capacity and preference for prey stages on P. citri is incomplete. Hence, a study was conducted to find out the daily prey consumption ability and preference for prey stages by different larval instars of S. epius reared on P. citri in the laboratory. Through the 8-day developmental period with four larval instars, the daily prey consumption of S. epius increased from the first to the seventh day and decreased on the eighth day prior to the prepupal stage. Generally, there was a significant difference in the prey consumption on different days. When the prey stages were offered separately, the first to fourth instar larva of S. epius consumed, respectively, a mean of 199.6, 722.6, 1908.8, and 4625.6 eggs or 21.5, 77.0, 168.5, and 670.5 nymphs or 3.2, 7.2, 16.0, and 35.1 adults of P. citri. When an S. epius larva was fed on P. citri eggs, nymphs and adults separately, it consumed a mean of 7456.7 eggs, 937.6 nymphs, or 62.3 adults during its entire development. When the prey stages were offered all together, a single S. epius larva consumed 2618.4 eggs, 170.4 nymphs, and 39.7 adults of P. citri throughout its entire development. The study revealed that S. epius is a voracious predator of P. citri and thus could be utilized as a major biological control agent.     

Host specificity assessment and potential impact of Megamelus scutellaris (Hemiptera: Delphacidae) on waterhyacinth Eichhornia crassipes (Pontederiales: Pontederiaceae)

The delphacid Megamelus scutellaris Berg was evaluated for host specificity and potential impact as part of a biological control program targeting Eichhornia. crassipes. Survival and development of adults and nymphs were used as metrics with no-choice, two-choice, nymph transfer, and sustainability tests conducted under quarantine conditions. A total of 69 plant species were tested including 12 from the Pontederiaceae (including E. crassipes). Additionally, 27 native and 5 exotic associated wetland species and 11 economic species were tested. Megamelus scutellaris exhibited a high level of oviposition and developmental fidelity to E. crassipes by failing to sustain populations on any non-target test plant past the F₁ generation. Nymph transfer tests which simulated potential spill-over events found that survival was virtually non-existent on associated wetland plants, regardless of taxonomic relatedness, including on Pontederia cordata, an important and widespread native species. Eichhornia crassipes plants exposed to two consecutive generations of feeding produced 66.9% less biomass and 73.4% fewer leaves than those in the controls. We conclude that Megamelus scutellaris is safe to release on E. crassipes in the United States.     

Observations on the effect of the weevils Neochetina eichhorniae Warner and Neochetina bruchi Hustache on the growth of water hyacinth

Water hyacinth Eichhornia crassipes (Mart.) Solms. is an aquatic weed that infests most of the White Nile system in the Sudan. Serious economical and ecological problems are caused by this weed. The two weevils Neochetina eichhorniae and Neochetina bruchi were imported and released in an attempted biological control against the weed. The adults of these weevils attack the plant and feed by removing tissues from the leaf pseudolamina and petioles. The larvae tunnel inside the petioles and the crown. The optimum temperature for feeding and development of both species is 25° C. Results obtained from stocking hyacinth plants with adults and larvae of both species separately revealed that N. bruchi is more efficient in checking the growth of the plant. The progeny of a pair of N. bruchi and N. eichhorniae reared separately on 41 hyacinth plants for a period of 61 days (one generation period) reduced their population growth by 25.4% and 12.7% respectively. The progeny of both species in a mixed culture reduced the growth of the plants by 22.5% in the same period, while in the control the population of the plants increased 136.6%.