Field assessment of a frond-feeding weevil, a successful biological control agent of red waterfern, Azolla filiculoides, in southern Africa

Azolla filiculoides (red waterfern) is a small, floating fern native to South America, that has invaded aquatic habitats, predominantly water resevoirs in southern Africa. A frond-feeding weevil, Stenopelmus rufinasus Gyllenhal (Coleoptera: Curculionidae), was imported from Florida, USA, and released as a biological control agent against this weed in South Africa at the end of 1997. To date, 24,700 weevils have been released, which has resulted in local extinction of red waterfern at 81% of the 112 release sites. The weevil has not failed to control a single site. Several sites were, however, lost due to flooding or drainage of dams. The surface area of weed controlled totalled 203.5 ha. On average, A. filiculoides was controlled in infested sites in 6.9 (±4.3) months. The weed recolonized at 22 of the sites (through either spore germination or dispersal by waterfowl), but the weevils subsequently spread to all of these sites and successfully caused local extinction of the weed at 18 of the sites. Five years after the release of the weevil, the weed no longer poses a threat to aquatic systems in southern Africa. In comparison to other biological control programs of aquatic weeds, the program against A. filiculoides in southern Africa ranks among the most successful cases anywhere in the world.     

Integration of biological control agents with other weed management technologies: Successes from the leafy spurge (Euphorbia esula) IPM program

An invasive weed can occupy a variety of environments and ecological niches and generally no single control method can be used across all areas the weed is found. Biological control agents integrated with other methods can increase and/or improve site-specific weed control, but such combinatorial approaches have not been widely utilized. The successful leafy spurge (Euphorbia esula L.) control program provides examples for future integrated weed programs that utilize biological control agents with traditional methods. Weed control methods can be used separately, such as when the leafy spurge gall midge (Spurgia esulae Gagné) reduced seed production in wooded areas while herbicides prevented further spread outside the tree line. Traditional methods also can be used directly with biological control agents. Incorporation of Aphthona spp. with herbicides has resulted in more rapid and complete leafy spurge control than either method used alone. Also, the insect population often increased rapidly following herbicide treatment, especially in areas where Aphthona spp. were established for several years but had been ineffective. Incorporation of Aphthona spp. with sheep or goat grazing has resulted in a larger decline in leafy spurge production than insects alone and in weed density than grazing alone. Controlled burns can aid establishment of biological control agents in marginally suitable environments, but timing of the fire must be coordinated to the insect’s life-cycle to ensure survival. Integration of biological control agents with revegetation programs required the agent to be the last method introduced because the cultivation and herbicide treatments necessary to establish desirable grasses and forbs were destructive to the insect. In a practical application, herbicides were combined with Aphthona spp. to help the insect establish and control leafy spurge in the habitat of the western prairie fringed orchid (Platanthera praeclara Sheviak and Bowles), an endangered species. Several experimental designs can be used to evaluate biological control agents with cultural, mechanical, and chemical control methods or with additional biological agents.     

The fixed plot survey method for determining the host range of the flowerbud-feeding weevil Dicomada rufa, a candidate for the biological control of Hakea sericea in South Africa

The hakea bud weevil, Dicomada rufa (Curculionidae), is a promising candidate for the biological control of the weed Hakea sericea (Proteaceae) in South Africa. Because D. rufa could not be successfully cultured on potted plants in quarantine, most of theconventional methods for host range determination were not suitable. A type of open-field testing method, the fixed plot survey method, was developed to show that D. rufa is host specific to H. sericea. The trial was conducted in three 1–2 ha plots at three localities in New SouthWales, Australia, involving 41 test plantspecies. This result was combined with otherconventional considerations to apply for therelease of D. rufa in South Africa. Theseincluded a multiple choice feeding trial inquarantine in South Africa, during which 10test species from seven genera of South AfricanProteaceae were not accepted for feeding. AllSouth African Proteaceae, except Brabejumstellatifolium, are phylogenetically distinctfrom H. sericea. Also, there are norecords of D. rufa interacting withcommercially important plants in Australia,including commercially cultivated South AfricanProteaceae. Dicomada rufa adult andlarval feeding destroys buds, flowers, smallfruits and succulent shoots. It is expectedthat this damage will supplement that of thetwo seed-feeding biological control agentsalready established in South Africa and furtherreduce the reproductive potential of the weed,particularly that of young plants regeneratingafter fires. Without compromising safety, thefixed plot survey method may also contribute toreducing the time and cost normally associatedwith conventional host specificity testing.     

Biological control of three floating water weeds,<Emphasis Type="Italic">Eichhornia crassipes,Pistia stratiotes</Emphasis>, and <Emphasis Type="Italic">Salviniamolesta</Emphasis> in the Republic of Congo

Since 1999, four specific weevils (Coleoptera, Curculionidae) were released in the Republic of Congo against three exotic floating water weeds: Neochetina eichhorniae Warner and N. bruchi Hustache against water hyacinth, Neohydronomus affinis Hustache against water lettuce, and Cyrtobagous salviniae Calder and Sands against water fern. Recoveries of exotic weevils were made from all 24 release sites except one, and all four species have established and spread (up to 800 km for water hyacinth weevils). Within a few years of releases, control of water fern and water lettuce was such that fishing and navigation could be resumed, while reductions of water hyacinth populations were only beginning.     

Biology and host range of Ophiomyia camarae, a biological control agent for Lantana camara in South Africa

The herringbone leaf-mining fly, Ophiomyiacamarae Spencer, is a promising candidateagent for the biological control of the alieninvasive weed Lantana camara L. in SouthAfrica. During extensive host-specificity testsinvolving 39 test plant species from 12families, survival to adulthood was restrictedto L. camara, L. trifolia, and fourspecies of the closely-related genus Lippia (Verbenaceae) in no-choice tests. However, survival of the immature stages wassignificantly lower on L. trifolia andthe four Lippia species than on L.camara. In addition, O. camaraedisplayed very strong oviposition preferencefor L. camara during paired-choice tests.This narrow laboratory host range suggests thatO. camarae will not pose any risks tonon-target verbenaceous plants if released inSouth Africa. Permission to release O. camaraeinto South Africa was approved by the regulatoryauthorities in 2001.     

Alcidodes sedi (Col.: Curculionidae), a natural enemy of Bryophyllum delagoense (Crassulaceae) in South Africa and a possible candidate agent for the biological control of this weed in Australia

The Madagascan endemic, Bryophyllum delagoense (Crassulaceae), is a major weed in Queensland, Australia. Despite having first been recorded in Australia in the 1940s, it is far more invasive there than on the African mainland where it was introduced more than 170 years ago. This may be due to a number of factors, one of which could be the occurrence of new natural enemy associations in southern Africa. Among the insects of crassulaceous plants that have extended their host ranges, a stem-boring weevil, Alcidodes sedi, was studied to elucidate its status as a natural enemy of B. delagoense in southern Africa and as a candidate biological control agent for introduction to Australia. Laboratory studies indicated that damage inflicted by adult and larval feeding caused significant reductions in stem length and number of leaves. Preliminary host-range trials revealed that A. sedi can complete its development on other species in the Crassulaceae, including most of the introduced Bryophyllum species and some Kalanchoe species native to South Africa. Despite the oligophagous nature of A. sedi and the fact that it can complete its development on a number of ornamental species in the Crassulaceae, it should be considered a potential biological control agent in Australia. All of the native Crassulaceae in Australia are in the genus Crassula, most of which are very small and therefore unlikely to support the development of a large weevil like A. sedi. However, additional host-range trials will have to be undertaken in Australia to determine whether the weevil can be considered safe for release.     

Molecular evidence that fireweed (Senecio madagascariensis, Asteraceae) is of South African origin

ITS1 sequence data was obtained for fireweed (Senecio madagascariensis) andS. lautus from Australia,S. madagascariensis andS. inaequidens from South Africa andS. madagascariensis from Madagascar. Despite the low level of variation (0.0–3.4%), these data further resolve the controversy concerning the identity and origin of fireweed. They confirm that fireweed is part of the South AfricanS. madagascariensis/S. inaequidens complex, and indicate that the infestation in Australia originated from South Africa as opposed to Madagascar. This will facilitate a resumption of biological control efforts in Australia and will direct surveys for control agents to South Africa.     

The effect of two biological control agents, the weevil Neochetina eichhorniae and the mirid Eccritotarsus catarinensis on water hyacinth, Eichhornia crassipes , grown in culture with water lettuce, Pistia stratiotes

We assessed the effect of two biological control agents, the mirid Eccritotarsus catarinensis (Carvalho) and the weevil Neochetina eichhorniae (Warner), singly or in combination, on the competitive ability of their host plant, water hyacinth, Eichhornia crassipes (Mart.) Solms-Laub., grown in a screen house, in competition with another aquatic plant (Pistia stratiotes L.). Water hyacinth plant growth characteristics measured included fresh weight, leaf and petiole lengths, number of inflorescences produced, and new shoots. Without herbivory, water hyacinth was 18 times more competitive than water lettuce (across all experimental combinations of initial plant densities), as estimated from fresh weights. Both insect species, singly or in combination, reduced water hyacinth plant growth characteristics. E. catarinensis alone was less damaging than the weevil and under normal conditions, i.e., floating water hyacinth, is not expected to increase control of water hyacinth beyond that of the weevil. When combined with the weevil, half the inoculum of weevils and half the inoculum of mirids produced the same growth reduction as the full inoculum of the weevil. Under conditions where the weevils are not effective because water hyacinths are seasonally rooted in mud, the mirid, which lives entirely on leaves, should become a useful additional biological control agent. Handling Editor: John Scott.     

The system management approach of biological weed control: Some theoretical considerations and aspects of application

Theoretical considerations behind the system management approach of biological weed control are presented. These include, a part describing and explaining the effects of parasitic fungi on crop – weed competition, a part describing and explaining the epidemic spread of parasitic fungi on weeds, and a part relating crop – weed competition at the population level to epidemics. The theoretical framework developed may also provide a basis for the use of other natural enemies, like insects, for biological weed control following the system management approach. Aspects of application are discussed using data of the interaction between the annual weed Senecio vulgaris and the rust fungusPuccinia lagenophorae.     

Direct and Indirect Effects of a Shoot-Base Boring Weevil and Plant Competition on the Performance of Creeping Thistle, Cirsium arvense

Creeping thistle or Canada thistle, Cirsium arvense (L.) Scop., is considered one of the world's worst weeds and the third most important weed in Europe. Biological control of this indigenous weed in Europe by use of native agents may provide a low-cost alternative to use of chemical or mechanical control measures and contribute to a more sustainable weed management. We investigated the potential of a shoot-base boring weevil, Apion onopordi Kirby (Coleoptera: Apionidae), for biological weed control, in the presence or absence of plant competition by three grass species. Infestation of thistle shoots by A. onopordi at natural infestation levels reduced above- and belowground plant performance after 2 years. Plant competition at natural levels had an overall greater effect than that of herbivory, significantly reducing both above- and belowground thistle performance in both years, thereby slowing the propagation of the weed. Weevil infestation and grass competition had a synergistic effect on C. arvense growth; the combined effects of the two factors was greater than the sum of both single-factor effects. The experiment revealed that A. onopordi promotes systemic infections of the rust fungus Puccinia punctiformis (Str.) Röhl in the year following weevil infestation. Systemically infected thistle shoots died before the end of the growing season. Although the direct effect of A. onopordi may not be sufficient to control creeping thistle, the synergistic interaction with plant competition and the indirect effect via promotion of systemic rust infections makes A. onopordi a promising agent for the biological control of this weed.     

Biology, host range, and risk assessment supporting release in Africa of Falconia intermedia (Heteroptera: Miridae), a new biocontrol agent for Lantana camara

The ornamental hybrid shrub, Lantana camara L. (lantana), is a serious environmental weed and has been targeted for biological control in South Africa since 1961. The established biocontrol agents cause insufficient levels of damage and additional natural enemies are required to reduce the invasiveness of this weed. The lantana mirid, Falconia intermedia (Distant), is a promising new agent that was imported from the Caribbean for life history and host-range studies. The nymphs and adults are leaf-suckers that cause chlorotic speckling, which reduces the photosynthetic capacity of the plant. Biological studies indicate that F. intermedia has considerable biocontrol potential, in that it has a high intrinsic rate of increase, the potential for multiple generations a year, highly mobile adults, and a high level of damage per individual. Host-specificity trials indicated that the lantana mirid has a narrow host range, with L. camara being the most suitable host, but several indigenous African species in the closely related genus Lippia are suitable alternative host plants. Under multiple-choice conditions, adults showed a significant and strong oviposition preference for L. camara over the Lippia species. A risk assessment of potential nontarget effects indicated that three Lippia species could sustain damage levels in the field. The relatively low probability of damage to indigenous species was considered a justified trade-off for the potentially marked impact on L. camara. The regulatory authorities accepted the results of this study and F. intermedia was released against L. camara in South Africa in April 1999.     

Food Preference and Reproductive Plasticity in an Invasive Freshwater Snail

The freshwater apple snail Pomacea canaliculata has become a major crop pest in southeast Asia and Hawai'i and threatens natural wetland habitats in these regions and elsewhere. Deliberately introduced as a potential human food resource, it has also been proposed as a possible biocontrol agent against aquatic weeds. Various factors may facilitate its rapid invasion of new areas; we focus on two: growth rate and food preference. Our field observations and laboratory experiments suggest that in Hawai'i P. canaliculata reaches reproductive maturity in 10 months or more, less time than in its native temperate and seasonal Argentina, where it takes 2 years, but longer than in parts of southeast Asia, where it may take as little as 2 months. This increased growth rate, and thence reproductive rate, probably facilitate rapid population growth. Although P. canaliculata is usually considered an indiscriminate generalist macrophytophagous feeder, laboratory experiments indicated preferences among the dominant plants at our field site and growth rate differences when constrained to feed only on one of these plants. Water hyacinth (Eichhornia crassipes), a major invasive weed, was not preferred in food choice experiments, and snails offered only water hyacinth on which to feed did not differ in growth rate from unfed snails. Another important invasive weed, water lettuce (Pistia stratiotes), was also not preferred, but snails fed on it did grow, though not as quickly as those fed on green-leaf lettuce. Among the food plants offered in the experiments the native Vigna marina was the most preferred. Therefore, although a generalist, P. canaliculata exhibits some discrimination among food plants. We recommend that it not be introduced for use as a biological control agent for aquatic weeds.     

油菜类型和品种对外来入侵杂草小子虉草的替代控制作用

外来入侵植物小子虉草（Phalaris minor Retz.）是世界公认的冬季农田恶性杂草,掌握农作物对其替代控制作用具有重要的研究价值。前期研究表明,油菜是替代控制小子虉草的优良农作物,然而,目前尚不清楚油菜类型与品种对其控制能力的影响。为此选取与小子虉草同域发生的不同类型（白菜型油菜、芥菜型油菜和甘蓝型油菜）油菜品种各3种,通过田间小区实验和室内化感作用测定,对比研究其对小子虉草的生长、繁殖、表型以及化感作用的影响。田间实验显示：竞争方式（种内或种间竞争）和油菜类型对小子虉草的地上生物量、种子数、株高、分枝数、叶面积和比叶面积存在极显著（P=0.0001）影响;而油菜品种对小子虉草的地上生物量（P=0.6064）、种子数（P=0.3577）、株高（P=0.4279）、分枝数（P=0.6357）、叶面积（P=0.8839）和比叶面积（P=0.3424）均无显著影响。3种类型油菜对小子虉草生长、繁殖以及表型的影响存在明显差异,其中芥菜型油菜对小子虉草的上述指标的影响最强,而白菜型油菜的影响最弱。室内生物测定显示,油菜对小子虉草具有化感抑制作用,当供试油菜叶片水提液浓度为0.1 g/mL时,小子虉草种子的萌发和幼苗的株高、根长、生物量均被显著抑制;研究也表明不同类型油菜对小子虉草的化感作用显著不同,同等条件下,芥菜型油菜对小子虉草的化感抑制作用最强。综上所述,油菜类型对外来入侵小子虉草的控制作用存在显著差异,其中芥菜型油菜对植物小子虉草的替代控制作用明显优于白菜型油菜和甘蓝型油菜,而其强的化感抑草特性或许是其强控草能力的原因之一。另外,本研究也为进一步利用油菜替代控制入侵植物小子虉草提供了参考。     

Ecological niche modelling of an invasive alien plant and its potential biological control agents

Invasive alien plants are of concern in South Africa. Pompom weed (Campuloclinium macrocephalum) is currently invading the Grassland and Savannah biomes of South Africa and is likely to continue spreading in the southern African sub- region. Two possible biological control agents (Liothrips tractabilis and Cochylis campuloclinium) have been identified for control of pompom weed. We used ecological niche modelling to predict which areas in southern Africa are likely to be suitable for pompom weed and the two potential biological control agents. The overlap between areas predicted to be highly suitable for pompom weed and areas suitable for the biological control agents was assessed. Methods of reducing sampling bias in a data set used for calibrating models were also compared. Finally, the performance of models calibrated using only native range data, only invaded range data and both were also compared. Models indicate that pompom weed is likely to spread across a greater region of southern Africa than it currently occupies, with the Savannah and Grassland biomes being at greatest risk of invasion. Poor overlap was found between the areas predicted to be highly suitable for pompom weed and those areas predicted to be suitable for the biological control agents. However, models of the potential distribution of the biological control agents are interpreted with caution due to the very small sample size of the data set used to calibrate the models. Models calibrated using both native range and invaded range data were found to perform best whilst models calibrated using only native range data performed the worst. There was little difference found between models that were calibrated using spatially reduced (selecting only one record per 30 min grid cell) and randomly reduced (randomly selecting 50% of available records) biased data sets.     

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.     

The trimorphic flea-beetle, <Emphasis Type="Italic">Alagoasa extrema</Emphasis>, not suitable for biocontrol of <Emphasis Type="Italic">Lantana camara</Emphasis> in Africa

The biology and host range of the flea-beetle, Alagoasa extrema (Harold) (Coleoptera: Chrysomelidae: Galerucinae: Alticini), was studied under quarantine laboratory conditions to evaluate the insect’s suitability for release as a biological control agent for the noxious weed, Lantana camara L. (Verbenaceae), in South Africa. Identification of this species proved noteworthy. The adults of A. extrema display three freely interbreeding and very distinct colour morphs, which can be confused with no less than 11 species of Alagoasa found in Mexico and the southwestern U.S. Initially, specimens were identified as two species of the genus Alagoasa Bechyné, i.e. A. quadrilineata (Harold) and A. extrema. Thirteen additional Alagoasa species and Kushelina petaurista (F). that can be confused with A. extremabased on external morphology of adults or larvae are discussed. Favourable biological characteristics included long-lived adults, several overlapping generations per year, and high adult and larval feeding rates. Observations from the insect’s native Mexican range and studies in South Africa suggest that A. extrema would probably be more suited to subtropical, rather than temperate areas in Africa. Host-specificity studies showed A. extrema to be an oligophagous species, capable of feeding and developing on several non-target species, especially two indigenous, African Lippia species (Verbenaceae). The host suitability of these indigenous species was only marginally lower than that of L. camara, and the potential risk to them was deemed to be too high to warrant release. It was therefore recommended that A. extrema be rejected as biocontrol agent for lantana in Africa.     

Biological studies on <Emphasis Type="Italic">Ceutorhynchus portulacae</Emphasis>, a potential natural enemy of the purslane weed <Emphasis Type="Italic">Portulaca oleracea</Emphasis>

Ceutorhynchus portulacae (Coleoptera Curculionidae) is identified as a potential natural enemy of Portulaca oleracea, which is a serious crop weed in many countries of the world. Detailed biological studies on its life cycle, fecundity, longevity and mortality factors revealed that the weevil possess positive attributes of successful biological control agents like short life cycle, prolonged longevity, high fecundity, small size, high rate of increase, phenology of attack, multivoltinism and versatile feeding behaviour. Based on the present study, it could be concluded that C. portulacae could contribute considerably to reduce purslane weed in crop fields.     

Host range of Ceutorhynchus assimilis (Coleoptera: Curculionidae), a candidate for biological control of Lepidium draba (Brassicaceae) in the USA

Ceutorhynchus assimilis has been selected as a potential biological control agent of Lepidium draba, which is a Eurasian invasive weed in North America. Preliminary studies indicated specificity of this weevil collected in southern France on L. draba. This result was in discord with the pest status of C. assimilis found in the literature. Host-specificity tests based both on field and laboratory experiments showed heterogeneity in the host spectrum of the weevils reared from different host-plants as determined by larval development. However, no distinguishable morphological differences could be visually detected between the populations feeding on different host-plants. All sampled populations of weevils were polyphagous as adults. Weevils reared from L. draba were specific to this plant for their complete larval development. Conversely, populations living on other wild and cultivated Brassicaceae species were not able to use L. draba as a host plant. Such differentiation is further highlighted by other biological aspects such as plant infestation rates, sex-ratio, duration of larval development, and differences in the timing of their life cycles. These results demonstrate that C. assimilis, an insect species formerly considered as a pest of Brassicaceae, is characterized by its host-range variability, with one population being potentially useful in the biological control of L. draba. Moreover, this example points to the need to test multiple populations of biological control agents in assessing risk.     

The effect of temperature,darkness, starvation and various food types on growth,survival and reproduction of Helisoma duryi,Biomphalaria alexandrina and Bulinus truncatus (Gastropoda: Planorbidae)

Helisoma duryi has been proposed as a biological control agent in schistosomiasis due to its superiority in laboratory competition experiments with various species of the intermediate host snails. Therefore it was considered important to evaluate the response of this snail species and the intermediate host species, Biomphalaria alexandrina and Bulinus truncatus, to various physical, chemical and biological factors under laboratory conditions in order to obtain information on the similarities in the ecological niches of these species. The factors considered in the present paper are: temperature, darkness, starvation and food. All three species had optimal growth and egg laying at 26–28 °C. Only H. duryi survived for a longer period at 33°C and it was capable of starting egg laying at this temperature although the onset was delayed. However, low temperature (18°C) caused a relatively larger decrease in egg laying of H. duryi than in the other two species. Growth and egg laying was reduced for H. duryi and B. truncatus kept under darkness and B. alexandrina could not tolerate maintenance under darkness. A few days of starvation of juvenile snails had no effect on later growth and egg laying capacity of the survivors, although mortality in B. truncatus was increased. B. alexandrina had a lower tolerance to starvation than the other two species. Egg laying of snails fed only one of the three laboratory food types decreased for all three species in the order: Vov-vov (dog food in dry pellets), Tetramin (fish food) and lettuce. Combinations of lettuce and one or more proteinaceous food types gave optimal growth and egg laying for all three species.     

Biology and host plant range of the soft scale,Steatococcus new species [Hem.: Margarodidae] for the biological control of the weedCryptostegia grandiflora [Asclepiadaceae]

The soft scale,Steatococcus new species, occurs onCryptostegia grandiflora R. Br. in Madagascar. Studies on its biology and host plant range showed colonies could survive for up to 6 months on some genera in theApocynaceae and indefinitely on many genera in the familyAsclepiadaceae. The species was therefore rejected as a biological control agent forC. grandiflora, a serious weed in northern Queensland, Australia.