Predicting the response of Cabomba caroliniana populations to biological control agent damage

Abstract  Cabomba caroliniana is a submerged aquatic plant from South America that is becoming a serious weed worldwide. It spreads by seed and by fragmentation and has an extremely wide climatic range, invading lakes and ponds from tropical (Darwin, Australia: latitude 12°) to cold temperate regions (Peterborough, Canada: latitude 45°). There are currently no effective methods of managing cabomba infestations and funding has been allocated to research biological methods. Surveys have examined cabomba in its native range and have identified several potential biological control agents. The most promising are a stem boring weevil ( Hydrotimetes natans ) and an aquatic moth ( Paracles spp.). Here we predict the change in cabomba populations after the introduction of the biological control agents. Our predictions are based on quantitative surveys of cabomba populations at three lakes in south-east Queensland, qualitative observations of cabomba in its native range, and conceptual knowledge of how the realised niche of cabomba might be affected by herbivore damage.     

Opportunities for biological weed control in Europe

The development and application of biological weed control offer greatopportunities not only for farmers, nature conservationists and othervegetation managers but also for institutions and companies that wish tosell plant protection services and products, and for the general publicthat demands safe food and a visually attractive and diverseenvironment. Despite the obvious opportunities for biological weedcontrol, few control agents are actually being used in Europe. Potentialagent organisms have features that make them particularly strong anduseful for biological control, but they also have weaknesses. Weaknessesinclude a too narrow or too wide host specificity, lack of virulence, orsensitivity to unfavourable environmental conditions.Developing specific knowledge on the interaction between weeds andpotential biological control agents, as well as expertise to increasethe effect of control agents and so achieve sufficient weed control in acost-effective manner, should have the highest priority in researchprogrammes. From 1994 to 2000 most ongoing research on biological weedcontrol in Europe was combined in a cooperative programme. This COSTAction concentrated on the interactions between five target crop weedsand their antagonists (pathogens and insects), on furthercharacterisation of the specific blems and potential control agents andon the most suitable biological control approach.The next major challenge will be to apply the findings provided byCOST-816 to the development of practical control solutions. The leadingobjective of a new concerted research programme with European dimensionswill be to stabilise or even promote biodiversity in the most importantEuropean ecosystems by integrating biological weed control in themanagement of these systems.     

Genetic diversity of alligator weed ecotypes is not the reason for their different responses to biological control

Biological control of alligator weed Alternanthera philoxeroides (Mart.) Griseb. using Agasicles hygrophila, a Chrysomelid beetle, has been successful in limiting growth in water, but not on land. In order to understand a possible genetic basis of this difference, technique using inter-simple sequence repeats (ISSR) markers was applied to analyse genetic diversity of this invasive weed. No genetic variation was detected not only within or between populations growing in the same habitats, but also between land- and water-grown populations. Thus we consider that the genetic variation is not the baseline factor resulting in the biological control difference in China. The differential success of pupation by the beetle may be related to the phenotypic plasticity of the plant stem diameter, rather than to genotypic factors.     

The potential for biological control of invasive alien aquatic weeds in Europe: a review

A retrospective analysis shows that invasive, alien, free-floating and emergent aquatic weeds in Europe are good targets for classical biological control, and that genus-specific chrysomelid and curculionid beetles offer the most potential. Ludwigia spp., Azolla filiculoides, Lemna minuta, Crassula helmsii and Hydrocotyle ranunculoides should be prioritised as targets. Fungal pathogens have been under-utilised as classical agents but, whilst they may have some potential against free-floating weeds, they appear to be poor candidates against submerged species, although the suitability of arthropod agents against these difficult targets still merits investigation. The use of indigenous pathogens as inundative agents (mycoherbicides) shows some promise.     

外来入侵杂草传统生物防治的生态风险及其防范对策

长期大量实践说明,引进天敌防治外来入侵杂草的传统生物防治方法是治理外来入侵杂草的一条切实可行的有效途径,但对其潜在的生态风险——对本土生物的直接或间接不良影响不容忽视。利用传统评价方法预测候选天敌的生态风险存在缺陷,主要表现在：（1）寄主专一性测定过分依赖室内进行的生理寄主范围测定结果,对生态寄主范围（实际寄主范围）问题重视不够,后者指在新环境中的一系列物理和生物条件下的寄主利用预测;（2）在生理寄主范围测定中,过分依赖完成生长发育的可能性,对行为、遗传性状以及系统发育关系重视不够;（3）在风险评估中,过多强调对经济作物的风险,而对自然生态系统的风险重视不够。对此,建议：（1）鼓励对已释放的天敌进行回顾性跟踪研究,从而为杂草生物防治实践提供生态学理论支撑;（2）在运用生物防治手段对付外来入侵杂草实践中,建议采用“有害推论”的预防性原则,以避免在面临入侵生物重大威胁时草率做出释放天敌的决策;（3）在评估候选天敌风险中重视生态效应的风险评估。     

Avoiding conflicts between insect and weed biological control: selection of non-target species to assess host specificity of cabbage seedpod weevil parasitoids

Abstract:  Classical biological control of insect pests and weeds may lead to potential conflicts, where insect pests are closely related to weed biological control agents. Such a conflict may occur in the classical biological control of the cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham) in North America, which belongs to the same subfamily, Ceutorhynchinae, as a number of agents introduced or proposed for introduction against non-indigenous invasive weed species. We propose a step-by-step procedure to select non-target species and thereby to develop a non-target species test list for screening candidate entomophagous biological control agents of a herbivore pest insect in a way that would simultaneously evaluate non-target potential on weed biological control agents and other non-target species. Using these recommendations, we developed a non-target test list for host specificity evaluations in the area of origin (Europe) and the area of introduction (North America) for cabbage seedpod weevil parasitoids. Scientifically based predictions on expected host–parasitoid interactions and ecological information about the ecological host range in the area of origin can help avoid conflicts, while still allowing the introduction of safe and effective agents against both insect pests and weeds.     

Biological control of alligator weed: Unsuccessful attempts to control terrestrial growth using the flea beetleDisonycha argentinensis [Col.: Chrysomelidae]

Biological control of alligator weed growing in aquatic habitats in Australia is successful but the agents, a flea beetleAgasicles hygrophila and a mothVogtia malloi, do not control terrestrial growth. Consequently another flea beetleDisonycha argentinensis was introduced into Australia specifically to control the terrestrial growth. Progeny of adults collected in Brazil from areas similar in climate and habitat to areas infested with alligator weed in Australia, were released but failed to become established. Eggs were laid by females released into a large field cage and some completed development, but the new adults failed to reproduce. Tentative conclusions are that microclimate or predation may have prevented establishment ofD. argentinensis but the results should not preclude attempts to establish this insect in North America, China or elsewhere.      

Strategies for the biological control of invasive willows (Salix spp.) in Australia

Abstract  Willows ( Salix spp.) are Weeds of National Significance in Australia where a large number of taxa are naturalised in temperate regions and can cause serious environmental degradation of riparian and wetland habitats. Several species are of economic or ornamental value and conflicts of interest could arise with planning their suppression. Biological control of six willow species ( S. alba L., S. cinerea L . , S. fragilis L., S. nigra Marshall, S. viminalis L., S . x  rubens Schrank) is under consideration in Australia using three approaches based on targeting different morphological structures. A rich phytophagous biota associated with Salix in their regions of origin indicate a range of organisms are available for development as biological control agents. Implementation strategies will depend on the host species targeted for biological control, their reproductive strategies, the level of control required to reduce environmental impacts and the level of conflict of interest associated with a biological control program.     

Optimizing nutritional conditions for the liquid culture production of effective fungal biological control agents

Spores of fungal pathogens of weeds and insects are unique in their ability to actively infect and kill their pest host. While these capabilities are advantageous in terms of their use as a contact biological control agent, or biopesticide, they also require special consideration during spore production. Directed approaches to medium optimization must consider not only spore yield but also spore qualities such as desiccation tolerance, stability as a dry preparation, and biocontrol efficacy. Nutritional conditions during culture growth and sporulation should direct the accumulation of appropriate endogenous reserves so that newly formed spores possess these advantageous qualities. Studies with the bioherbicidal fungus Colletotrichum truncatum and with the bioinsecticidal fungus Paecilomyces fumosoroseus have demonstrated the impact of nutrition on spore ‘fitness’ for use as a biological control agent. The optimization strategy used in these nutritional studies as well as a comparison of the results are presented. Received 06 February 1997/ Accepted in revised form 29 May 1997     

Biological control of waterhyacinth in Sinaloa Mexico with the weevils Neochetina eichhorniae and N. bruchi

Water hyacinth (Eichhornia crassipes (Mart.) Solms.)creates severe problems in the irrigationdistricts of Mexico, particularly in westernSinaloa. Therefore water hyacinth weevils(Neochetina eichhorniae Warner and N. bruchi Hustache), imported from the USA in1993, were used to initiate a biologicalcontrol program. Precautionary screeningrevealed that some were infected with amicrosporidian so disease-free colonies wereproduced by eliminating infected breedinglines. To demonstrate effectiveness prior toopen field releases, weevils were firstreleased in cages at field sites. Weevilintensity increased to 6.3 weevils/plant after320 days when the plants were all dead ordying. More than 8,600 N. bruchi and14,500 N. eichhorniae were then releasedat various sites during January 1995 to August1996. Waterhyacinth coverage declined atBatamote reservoir (134 ha) from 95% to <3%by 1997; at the 12-ha Hilda reservoir from100% in May 1995 to 1% by March 1998; at the42.3-ha Arroyo Prieto reservoir from 100% to1% during the same interval; and at theMariquita reservoir (492 ha), the largestreservoir in the Humaya system, from 394 ha(80%)to 98.4 ha (20%).     

Long-term population studies and the development of an integrated management programme for control of Opuntia stricta in Kruger National Park, South Africa

1. A cactus, Opuntia stricta , has invaded almost 16 000 ha of conserved, natural habitatand has become a major weed problem in Kruger National Park (KNP), South Africa.
2. The main objectives in the control of O. stricta are to reduce the density of the weed and to curb long-range dispersal of seeds by preventing young plants from reaching the size (28 cladodes) at which they start to produce fruits.
3. Herbicides have failed to provide satisfactory control of O. stricta because the weed infestations are replenished from seeds in the soil and from small plants that are overlooked during spraying.
4. A phycitid moth, Cactoblastis cactorum , was released in KNP during 1988 in an attempt to control O. stricta biologically.
5. Population counts of the biological control agent and of the weed over a 5-year period showed that, even though C. cactorum has not provided complete control of O. stricta in KNP, the moderate levels of larval damage have stunted the growth of O. stricta and have considerably extended the time that the young plants take to reach sexual maturity.
6. Comparisons of modelled (i.e. with no C. cactorum ) and actual populations of O. stricta showed that C. cactorum is making a substantial contribution to the control of O. stricta in residual infestations of the weed that have been treated with herbicides.
7. The need for long-term evaluation studies in biological weed control is demon strated by the development of an integrated management programme for effective control of O. stricta .     

The value of simulating herbivory in selecting effective weed biological control agents

Invasive species have a significant economic and ecological cost and biological control can be a powerful tool in their management. Classical biological control practice involves the re-establishment of trophic links between specialist insect and fungal agents to regulate populations of invasive species. However, the permanent nature of biological control agent introductions raises concerns about the unintended consequences of such introductions on non-target organisms, particularly when such agents are ineffective on their target organisms. In this paper, we explore the current debate in the selection of agents for weed classical biological control. We then propose an alternate approach, based on studying plant response to simulated herbivory, that could minimize the chances of release of ineffective agents. Simulating herbivory could yield insights into the vulnerability of plants to certain types of damage. Selection of agents within guilds that are likely to have a significant influence on the plant can improve the chances of achieving biological control sooner and reduce the likelihood of releasing ineffective agents that may have non-target impacts. We propose a method by which simulated herbivory could be integrated into the agent selection process and discuss its strengths and shortcomings. We present case studies of two Neotropical weeds illustrating how this method could be applied.     

Biological control of the wetlands weed purple loosestrife (Lythrum salicaria) in the Pacific northwestern United States

Purple loosestrife (Lythrum salicaria) is an Eurasian perennial hydrophyte that has become naturalized in wetlands and in and along waterways throughout temperate North America. The ecological integrity of such areas is threatened by rapidly forming monotypic infestations that displace valued flora and diminish critical fish and wildlife habitat. The inability of physical, cultural, and chemical methods to provide adequate control of the weed has led to the development of an insect-based biological control program. The first field releases of the bud and leaf feeding beetles, Galerucella calmariensis and G. pusilla, and a root-mining weevil, Hylobius transversovittatus, were made in the United States and Canada in 1992. A total of 4740 Galerucella spp. adults were released in central Washington during 1992 and 1993 at eight sites and 471 H. transversovittatus egg inoculations were made in 1993 at three locations. Establishment of both Galerucella spp. was confirmed and Hylobius colonization was achieved.     

Analysis of importations for biological control of insect pests and weeds in New Zealand

Importations of biological control agents for insect pests and weeds in New Zealand are summarized and factors contributing to the relative success of the programmes are examined. The establishment rate of 30.9% is similar to that achieved worldwide, but is significantly lower than the rate achieved in the island habitat of Hawaii. The pioneering role of New Zealand in biological control is shown by the high proportion of programmes first attempted in this country. Although this novelty has not reduced the establishment rate, introductions against endemic species have not succeeded. Size of release was not a dominant feature in the establishment of agents. Complete or substantial success is recorded for 17 of the 70 target pests, with a relatively high success rate in forestry programmes. Examples of the influence of climate matching and competitive exclusion are also discussed. Changing practices and attitudes to the introduction of biological control agents are documented to show the increasing emphasis on specialists. No adverse effects of introductions are reported. The challenge to practitioners and regulators is to develop systems to evaluate conflicts of interest and develop workable mechanisms to determine which biological control agents are suitable for release.     

Effects of weed control on species composition of aquatic plants and bank plants and macrofauna in ditches

For several years, the effect of ditch management measures on the species composition of aquatic and bank flora and macrofauna were studied in a polder with a peat soil. The mowing basket increased the presence of floating plants whereas the ditch-scoop increased filamentous algae and submerged plant species. Monospecific stands of aquatic plants had fewer macrofaunal species than stands with many plant species. Direct and indirect effects of management measures on the macrofauna composition were shown.     

Risk analysis and management decisions for weed biological control agents: Ecological theory and modeling results

Biologically based control methods offer many advantages for the control of invasive plant species; however, these methods are not without risks to native species. Thus, there is a need for more effective and efficient methods of risk analysis for biological control agents. We show how the process of ecological risk assessment established by the United States’ Environmental Protection Agency may be adapted to improve assessment of the risks of proposed biological control agents. We discuss the risks posed by weed biological control agents, and present a simple individual-based model of herbivorous insect movement and oviposition on two species of host plant, a target invasive plant species and a non-target native species, in simulated landscapes. The model shows that risks of non-target impacts may be influenced by the details of the movement behavior of biological control agents in heterogeneous landscapes. The specific details of insect movement that appear to be relevant are readily measured in field trials and the general modeling approach is readily adapted to real landscapes. Current biological control risk assessments typically emphasize effects analysis at the expense of exposure analysis; the modeling approach presented here provides a simple and feasible way to incorporate exposure analyses. We conclude that models such as ours should be given serious consideration as part of a comprehensive strategy of risk assessment for proposed weed biological control agents.     

Perspectives on the development of biological control agents in Africa

Concern over the impact of chemical pesticides on human health and the environment continues to stimulate the study of alternative, biological pest control agents (BCA). In Africa, use of BCA products is just emerging, primarily in the export sector where growers are using a mixture of products from new African companies and products imported from outside Africa. In contrast, although interest in BCA for use in domestic markets has been widely supported by development funding partners for many years, few products have become available to these growers. The limited uptake of BCA in Africa suggests there may be constraints and issues working against their wider implementation. Here, we focus on non-technical constraints and issues facing microbial BCA applied inundatively, but arguments extend to a greater or lesser degree to all BCA. The challenges to wider BCA availability could be overcome by: a more coherent approach to their regulation across the continent; support for BCA development that is based on evidence of need and applicability; and political advocacy to lobby for necessary resources being made available to all stakeholders.