Environmental weeds in Australia and New Zealand: issues and approaches to management

Environmental weeds are plants that invade natural ecosystems and are considered to be a serious threat to nature conservation. Australia and New Zealand, where biota with a high degree of endemism have evolved, are particularly susceptible to environmental weeds. Environmental weeds have been implicated in the extinction of several indigenous plant species, and they also threaten ecosystem stability and functional complexity. Historically, emphasis has been placed on the chemical or manual ‘control’ of weed infestations, often with little consideration of the long‐term effectiveness or the ecological consequences of such an approach. As the threat from environmental weeds is becoming more fully recognized, an integrated, strategic and ecological approach to weed management is being recommended. In both countries, systems for screening new plants before allowing entry for cultivation have been developed. For already established plants, management is conducted within a legislative and policy framework such as the Regional Pest Management Strategies that operate through the Biosecurity Act 1993 in New Zealand. Noxious weed legislation in Australia has historically focused on agricultural weeds, but some Acts are (or have recently been) undergoing revision to give greater emphasis to environmental weeds and the involvement of the community in weed management. Quarantine, legislation, research and on‐ground management are complemented by education programmes about the impact and control of environmental weeds. This paper provides an overview of the ‘tool‐kit’ needed to manage environmental weeds in Australia and New Zealand, comparing and contrasting the approaches taken in the two countries. It also provides a broad framework for the case studies that make up this special issue on the ecology and management of environmental weeds in both countries.     

Factors affecting the cost of weed biocontrol programs in New Zealand

Many national schemes for setting priorities for invasive weed management have emphasized the current or future impacts of the weed more than the cost or feasibility of control, perhaps because the latter may be difficult to estimate. As part of a project to improve prioritization of weed biocontrol targets in New Zealand, we investigated factors that were hypothesized to influence the cost of conducting weed biological control, using data from New Zealand programs. Taxonomic isolation of the target weed, relative to commercially important plants and native flora was not a significant influence on program cost, although we present evidence that disease, which to date has only affected agents released against taxonomically isolated weed targets, has masked the importance of taxonomic isolation in New Zealand. Opposition to biocontrol has caused delays, but has not had a major influence on the cost of biocontrol in New Zealand, probably because weed species with the greatest potential for opposition were identified during feasibility studies and avoided, or because conflicts were resolved by conducting cost-benefit analyses that were minor components of the total program costs. Only two factors explained virtually all the variance in program cost: program type (repeat programs were cheaper than novel/pioneering programs); and the number of agent species released. The predicted cost of future weed biocontrol programs can now be incorporated into decision-making tools ranking New Zealand weed biocontrol targets. Efficiencies in future programs are most likely to be gained by better agent selection so that fewer agents are released. For repeat programs this could be achieved by waiting until monitoring has been conducted overseas, so that the best agents or combination of agents can be selected for any particular weed. This reiterates the need for better post-release evaluation of weed biocontrol agent effectiveness worldwide.     

Establishment and impact of insect agents deployed for the biological control of invasive Asteraceae: prospects for the control of <Emphasis Type="Italic">Senecio madagascariensis</Emphasis>

Several invasive Asteraceae have been targeted for biological control worldwide, with variable success. Senecio madagascariensis Poiret, which invades agricultural lands in Australia and Hawaii, is a recent target. Since several potential insect agents were recorded in the plant’s native range in South Africa, we assessed biocontrol efforts against asteraceous weeds to determine those most likely to deliver success. Some 108 insect species, from five orders and 23 families, were deployed against 38 weed taxa, mostly in the mainland USA, Canada, Australia and New Zealand. Coleoptera (mainly Curculionidae and Chrysomelidae), Diptera (Tephritidae) and Lepidoptera (Tortricidae) featured the most. Despite high establishment success (73% of releases across countries), only 37% of successful releases achieved meaningful impact. Although root-feeding and stem-feeding insects appeared to be the best candidates, neither insect family nor feeding guild significantly influenced the probability of success. This synthesis of the global contribution of different guilds of specialist herbivores to the management of invasive Asteraceae is guiding the selection of candidate agents for the biocontrol of S. madagascariensis in Australia.     

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.     

The current status of biological control of weeds in southern China and future options

China has become one of the countries most seriously affected by invasive alien weeds in the world. Weeds impact agriculture, the environment and human health, and conventional control methods such as herbicides are expensive, damaging to human health and unsustainable. As the impacts and costs of weed control in China increase, there is an urgent need to manage some of the more important weeds through more sustainable methods. Classical biological control of invasive alien weeds is environmentally-friendly and sustainable. Biological control in China began in the 1930s with the introduction of two agents into Hong Kong for the control of Lantana camara. Since then, a further seven biological control agents have been introduced into China to control four weed species. In addition, 11 biological control agents targeting seven weed species have naturally spread into China. Together, these biological control agents are helping to control some of China's worst weeds. However, these efforts are only a small portion of the weeds that could be targeted for weed biological control. This paper reviews the current status of weed biological control efforts against introduced weeds in ten provinces and regions in southern China and provides a platform to identify the most effective and appropriate weed biological control opportunities and programmes to pursue in the future. Introducing additional safe and effective biological control agents into China to help manage some of the worst weeds in the region should reduce the use of herbicides and impacts on human health and the environment, while increasing productivity and food security.     

A review of <Emphasis Type="Italic">Solanum mauritianum</Emphasis> biocontrol: prospects,promise and problems: a way forward for South Africa and globally

The invasive tree Solanum mauritianum Scopoli remains one of the world’s most widespread environmental weeds. Despite biocontrol providing one of the few viable long-term solutions to tackling S. mauritianum invasions globally, only South Africa and, more recently, New Zealand, have programmes in place. Ongoing biocontrol efforts against S. mauritianum are reviewed here with particular reference to South Africa. The South African programme has suffered a troubled history, with considerable research efforts culminating in the eventual release and establishment of only two insect agents, Gargaphia decoris Drake and Anthonomus santacruzi Hustache. The difficulties experienced have hindered research into new agents, causing apprehension in using biocontrol internationally. However, recent studies have demonstrated that biocontrol may be deserving of renewed investment, particularly within an integrated management context. In this review, we advocate for the revival of the S. mauritianum biocontrol programme in South Africa, and discuss possible avenues for future research internationally.     

Biological control of weeds: an analysis of introductions,rates of establishment and estimates of success,worldwide

The foremost document that comprehensively reports on biological control introductions against weeds—‘Biological control of weeds: a world catalogue of agents and their target weeds’—has been updated and now includes all deliberate releases made through 2012. It includes data on 1555 intentional releases of 468 biological control agent species used against 175 species of target weeds in 48 plant families, in 90 countries. For 55 (31.4%) of the target weed species, only one biocontrol agent was introduced. The largest number of agent species (44) was introduced for the biological control of Lantana camara (Verbenaceae). Three insect orders (Coleoptera, Lepidoptera and Diptera) comprised about 80% of all biocontrol agent species released and releases made. Of the 468 biocontrol agent species introduced, 332 (70.9%) established in at least one instance. Of the 313 species, for which impact could be categorized, 172 (55.0%) caused medium, variable or heavy levels of damage (impacts). Of all releases made through 2012, 982 (63.2%) led to establishment. Forty-two releases were judged too early post-release to categorize impact, leaving 940 releases for which impact analyses were conducted. Similar to agent species, approximately half of the established releases (503 or 53.5%) caused medium, variable or heavy levels of damage on the target weeds, and almost a quarter of releases (225 or 23.9%) caused heavy impact. Across all countries and regions, 65.7% of the weeds targeted for biological control experienced some level of control. These data indicate the value of this practice, on its own, or as a supplement to other methods, in the management of invasive plants.     

利用传统生物防治控制外来杂草的入侵

随着国际贸易的日益频繁,外来有害植物入侵,严重威胁我国的自然环境和生物多样性。利用从原产地引入食性较专一的天敌来控制外来杂草是杂草生物防治的主要方式之一,有保护环境一劳永逸的效果。简要介绍了国际生物防治概况,统计表明全世界至少有133种目标杂草进行生物防治,主要分布在菊科、仙人掌科和含羞草科,63科369种无脊椎动物和真菌作为杂草生物防治的天敌,利用最多的天敌是鞘翅目象甲科和叶甲科昆虫,其中大多数项目是治理外来杂草的。杂草生物防治最活跃的国家依次为美国、澳大利亚、南非、加拿大和新西兰。重点论述了利用传统生物防治方法防治外来杂草的经典项目、国内外研究概况,以及目前面临的问题和应用前景。我国杂草生物防治起步晚,传统杂草生防的目标杂草有4种,紫茎泽兰、空心莲子草、豚草和水葫芦,其中,空心莲子草的生物防治获得成功。共引进天敌14种,输出天敌23种,与世界上生物防治先进的国家比尚有距离。中国应充分借鉴国际成功经验,对外来杂草开展生物防治。中国的生物多样性在世界上占有十分独特的地位,将在生物多样性保护中发挥重要作用。     

The effects of gibberellin A3 on white clover during floral development

Hokkanen & Pimentel (1984) proposed a novel approach for the selection of biocontrol agents. They advocated the selection of agents from a relative of the weed plant rather than from the target weed species itself. The new relationship that such agents would have with the weed would be characterised by a relative lack of homeostasis compared with the old herbivore-plant associations traditionally used in weed biocontrol, and would consequently be more effective. There are few examples to support these contentions because of the traditional use of old herbivore-plant associations in weed biocontrol. In the present study, herbivore-plant associations in agricultural situations, which are analogous to agent-weed associations, are examined to assess the potential of new associations for weed biocontrol. The herbivores on 14 introduced crop plants which have salient similarities to the major weeds in the south-western Cape were surveyed: (a) 68% of the 188 insect and mite herbivores are indigenous species in new associations with these host plants, and (b) of the five most damaging pests on each of 13 of the crop plants, 53% are in new associations with the plants. Of the 40 most important agricultural pests in South Africa, 58% are in new associations, confirming these results. About 50% of the insect and mite herbivores in new associations with their host plants are oligophagous, indicating that new associations are not necessarily characterised by polyphagy and hence unsafe for use in weed biocontrol. We conclude that new associations between herbivore species and host plants have strong potential in weed biocontrol because (a) their frequency in agriculture indicates that they can easily be established in weed biocontrol situations, (b) they are as damaging as old associations and (c) they are not necessarily unsafe as regards host specificity. We therefore advocate the use of both the classical approach and that of Hokkanen & Pimentel (1984). Our support for Hokkanen & Pimentel (1984) is, however, based on evidence and rationale clearly different to theirs, and we provide novel guidelines, which can be routinely and practically applied in the selection of agents for weed biocontrol.     

Experts know more than just facts: eliciting functional understanding to help prioritise weed biological control targets

Prioritising investments in classical weed biological control (biocontrol) is a common decision-making challenge: biocontrol programmes can yield substantial benefits but are typically long-term and costly, and the outcome uncertain. Experts are often relied upon to help, but their role is generally restricted to providing facts and judgements to populate an existing prioritisation model, which in turn receives little scrutiny. We developed and applied a new prioritisation framework to guide biocontrol investment decisions by livestock industries that required eliciting experts’ functional understanding (including their in-depth knowledge of the theoretical and practical drivers of weed biocontrol programmes). This consultative and transparent framework drew on expertise from most biocontrol practitioners in Australia through a structured workshop, and the literature. Each of the 75 weed taxa considered was placed in a matrix according to their impact (current or potential) and the prospects of biocontrol achieving pre-defined management goals. There was considerable knowledge uncertainty regarding potential impacts, which is of concern when making pre-emptive investments. Feasibility (likelihood of finding host-specific agents) and likelihood of success (management goals being met, assuming that host-specific agents are available) of biocontrol were both assessed as low for 51 % of taxa. Predicted barriers to successful biocontrol were diverse and idiosyncratic, suggesting that application of more quantitative prioritisation approaches would be challenging. A short-list of 13 weed taxa was identified for further consideration as biocontrol targets, based on the trade-off between potential impact and prospects for biocontrol. Research priorities emerged from the prioritisation process that would maximise investment outcomes for each taxon. Only two short-listed taxa are new targets, reflecting the maturity of the biocontrol discipline targeting weeds of livestock industries in Australia. Accessing the in-depth functional understanding of experts resulted in explicit characterisation of the barriers to successful biocontrol and if/how they might be overcome, improved characterisation of uncertainty, and provided directed guidance for investment. Such an approach would be readily applicable to analogous decision-making challenges in other sectors and countries.     

Constraints in weed biological control: contrasting responses by implementing nations

The compilation of papers in this Special Issue (SI) derives from a Symposium at the 25th International Congress of Entomology, in 2016, entitled “Rise or demise? A global outlook on the future of classical biological weed control”. In the SI-opening-paper, a summary of the 5th edition of the world catalogue of weed biocontrol agents and their target weeds provides a comprehensive international perspective. Weed biocontrol implementation is beleaguered by perceptions of risk and restrictive regulatory procedures, notably in the USA, and less so in Canada. Thus, most of the papers in this SI comprise accounts of innovative responses to these challenges from scientists in the USA. Political and funding issues have inhibited weed biocontrol in Australia over the past decade, but there appears to be a gradual reversal of this trend in recent years. In contrast, in New Zealand and in South Africa, the practice is flourishing, and there are significant recent initiatives in Europe. Overall, the contributions in the SI suggest an optimistic prognosis for weed biological control.     

Apparent competition can compromise the safety of highly specific biocontrol agents

Despite current concern about the safety of biological control of weeds, assessing the indirect impacts of introduced agents is not common practice. Using 17 replicate food webs, we demonstrate that the use of a highly host-plant specific weed biocontrol agent, recently introduced into Australia, is associated with declines of local insect communities. The agent shares natural enemies (predators and parasitoids) with seed herbivore species from native plants, so apparent competition is the most likely cause for these losses. Both species richness and abundance in insect communities (seed herbivores and their parasitoids) were negatively correlated with the abundance of the biocontrol agent. Local losses of up to 11 species (dipteran seed herbivores and parasitoids) took place as the biocontrol agent abundance increased. Ineffective biocontrol agents that remain highly abundant in the community are most likely to have persistent, indirect negative effects. Our findings suggest that more investment is required in pre-release studies on the effectiveness of biocontrol agents, as well as in post-release studies assessing indirect impacts, to avoid or minimize the release of potentially damaging species.     

Ecology and management of Melaleuca quinquenervia,an invader of wetlands in Florida,U.S.A.

Invasive weeds are potent agents of environmental change. Wetlands are valuable environments that frequently are impinged by a variety of threats including invasive weeds. Melaleuca quinquenervia (Cav.) S.T. Blake (broad-leaved paperbark), though experiencing major diminishment of native populations in Australia, is naturalized and highly invasive in most wetland habitats of south Florida, U.S.A., where it infests more than 202,000 ha. Wetlands in south Florida, including such renowned areas as the Everglades, are being transformed into M. quinquenervia swamps, with major environmental and economic impacts. Current management methods include herbicides, mechanical or hand removal of plants, flooding, and prescribed burning. Insufficient information, high costs, non-target impacts, and the resilience of M. quinquenervia (trunk and root sprouts and massive canopy seed banks) greatly constrain the effectiveness of these control methods. Biological control offers long-term management potential, most likely by reducing the rate of spread and the vitality and growth rate of plants, thus rendering them more vulnerable to other environmental stresses and control methods. The leaf weevil Oxyops vitiosa Pascoe, a natural enemy of M. quinquenervia in Australia, will likely be the first biocontrol agent released against the weed in Florida. More information is needed, especially ecological data, to better understand the invasiveness of M. quinquenervia in Florida and to facilitate its management there.     

Making weed biological control predictable,safer and more effective: perspectives from New Zealand

A persistent problem in weed biocontrol is how to reliably predict whether a plant that supports development in laboratory host-specificity testing will be utilized in field conditions, and this is undoubtedly preventing releases of safe and effective agents. Moreover, the potential for unanticipated undesirable indirect effects of weed biocontrol on ecological networks has raised concerns by policy-makers and the general public. The key to minimizing risks of non-target impacts is prioritizing candidate agents that are both host-specific and effective, such that the number of agents required to bring the weed under control is minimized. As a consequence both the weed and its biocontrol agents become minor components of the local biota. Here we review recent attempts in New Zealand to improve the predictive ability of host-range testing, to avoid potentially safe and effective agents being rejected. Research in New Zealand aimed at predicting whether an agent is likely to experience enemy-release (i.e. reduced parasitism and predation) could assist agent prioritization, potentially making biocontrol both environmentally safer and more effective.     

The status of biological control and recommendations for improving uptake for the future

Classical and augmentative biological control of insect pests and weeds has enjoyed a long history of successes. However, biocontrol practices have not been as universally accepted or optimally utilised as they could be. An International Organisation for Biological Control (IOBC) initiative brought together practitioners and researchers from widely diverse fields to identify the main limitations to biocontrol uptake and to recommend means of mitigation. Limitations to uptake included: risk averse and unwieldy regulatory processes; increasingly bureaucratic barriers to access to biocontrol agents; insufficient engagement and communication with the public, stakeholders, growers and politicians of the considerable economic benefits of biocontrol; and fragmentation of biocontrol sub-disciplines. In this contribution we summarise a range of recommendations for the future that emphasise the need for improved communication of economic, environmental and social successes and benefits of biological control for insect pests, weeds and plant diseases, targeting political, regulatory, grower/land manager and other stakeholder interests. Political initiatives in some countries which augur well for biocontrol in the future are discussed.     

Decomposition dynamics of invasive alligator weed compared with native sedges in a Northland lake

Invasive weeds have been shown to alter ecosystem processes such as decomposition and nutrient cycling. However, little is known about the effects of introduced biocontrol agents on these processes. This study examined the effects of alligator weed (Alternanthera philoxeroides) and its biocontrol agent, the alligator weed flea beetle (Agasicles hygrophila), on nutrient cycling in a northern New Zealand lake. Alligator weed litter decomposed significantly faster than either of two native sedge species (Schoenoplectus tabernaemontani, Isolepis prolifer) in a litterbag experiment. In addition, the presence of the alligator weed flea beetle resulted in large amounts of decaying alligator weed litter entering the lake in early summer. Both the timing and magnitude of this litter input were uncharacteristic of seasonal biomass dynamics of the native sedges. Combined with alligator weed?s rapid decomposition, this indicates altered patterns of nutrient cycling at the lake, with potential flow-on effects including facilitation of further weed invasion.     

Ecological,practical, and political inputs into selection of weed targets: What makes a good biological control target?

The topic of ecological, practical, and political considerations in the selection of weed targets for biological control has been widely discussed during the past two decades, mostly from the perspective of insect herbivores. For conceptual and practical purposes, plant pathogens have been treated in these discussions as if they are a subset of inoculative biocontrol agents, with little said about the inherent differences between pathogens and insects as biocontrol agents or the selection of weed targets for control by the inundative, bioherbicide strategy. Herein, I attempt to address the question of what makes a good biological control target for plant pathogens used as inoculative as well as inundative agents, basing my analysis on examples from the past three decades. Despite the small number of examples available for this analysis, the following generalizations can be made: (1) Weeds with robust capacity for vegetative regeneration are more difficult to control with pathogens than those that lack this trait. (2) A plant’s growth habit is not a reliable guide for target selection; weeds that have been successfully controlled include annual and biennial herbs, perennial shrubs, perennial vines, and trees, while numerous failures have been reported irrespective of the target’s growth habit or reproductive mode. (3) It is more challenging to control species with genetic heterogeneity and capacity for introgression than genetically homogeneous and reproductively conserved species. (4) Matching the target host’s susceptibility with the candidate pathogen’s virulence is of utmost importance for biocontrol success since host–pathogen interactions at the species and subspecies levels are often governed by single-gene differences (e.g., varietal specificity). (5) Practical and political considerations are central to the selection of targets for control with pathogens. (6) Demand from influential stakeholders for control and/or for a nonchemical or economically sustainable control typically drives the initiative as well as the continuance of biocontrol projects to their completion. (7) In the case of inundative, bioherbicide agents, the continuity and ultimate implementation of a project will be dictated by the prospects of economic returns from developing and using a pathogen. (8) The stakeholders’ perceptions of the effectiveness of a biocontrol program can be unpredictable, leading to conflicting views of “success.” In the final analysis, a good weed target for control by a pathogen is one that has strong stakeholder backing and the list of available pathogens for the target suggests a possibility of acceptable control at a cost that is competitive with those of other control options. While this conclusion is also applicable to target selection for insect biocontrol agents, it is more relevant for pathogens because of limited funding and personnel available for development of pathogens and the added cost and technological complexity of implementing bioherbicides compared to classical biocontrols.     

Rearing beneficial insects for biological control purposes in resource poor areas of africa

Biological control of insect pest and weeds using beneficial insects in resource poor areas is not very well supported. Poor funding has affected in particular the importation of classical biological control agents, quarantine, rearing, research facilities, and the research programmes. Donor agencies, commercial and semi-commercial enterprises in a number of African countries have, however, been able to contribute to biological control efforts using beneficial insects by providing some of the resources needed. This has led to biological control becoming a real possibility to control insect pests and weeds in several resource poor countries. Examples are provided of the “spin-offs” of such programmes for resource poor areas in South Africa, Zambia, Kenya, Malawi, Benin and Nigeria. Conventional insect rearing, insect conservation and habitat management as an aid to biological control programmes are discussed.     

The role of pre-release efficacy assessment in selecting classical biological control agents for weeds—applying the Anna Karenina principle

The goals in selecting classical biological control agents for weeds are to identify agents that will be both safe for release and effective in controlling their target plants. The release of ineffective agents should be avoided, as these add to the costs and risks of biological control without contributing to its benefits. While the principles of host-specificity testing and risk assessment for weed biological control agents have been extensively debated and refined, there has been less attention given to assessing the probable efficacy of agents prior to release. This reluctance to undertake pre-release efficacy assessment (PREA) is probably based on concerns that it will both add to the cost of screening biological control agents and introduce a risk of wrongly rejecting effective agents. We used a project simulation model to investigate the implications of using PREA as an additional filter in the agent selection process. The results suggest that, if it can be done at a lower cost than host-specificity testing, the use of PREA as the first filter can make agent selection more cost-effective than screening based on host-specificity alone. We discuss examples of PREA and potential approaches. The impact of biocontrol agents is a function of their range, abundance, and per-capita damage. While it will always be difficult to predict the post-release abundance of biological control agents from pre-release studies, some estimates of potential range can be obtained from studies of climatic adaptation. For agents that affect the vegetative growth or survival of their target weeds, experimental measurement of per-capita damage is feasible and can contribute to a reduction in the numbers of ineffective agents released. The Anna Karenina principle states that success in complex undertakings does not depend on a single factor but requires avoiding many separate causes of failure. We suggest that, in biological control of weeds, the use of agents that are not sufficiently damaging is one such cause that can be partially avoided by the use of pre-release efficacy assessment.     

Integration of Herbicides with Arthropod Biocontrol Agents for Weed Control

Classical biological control of weeds using arthropods is being attempted on a large scale in a number of countries, sometimes with spectacularly successful outcomes. However, in many cases biocontrol is not completely effective and use of herbicides on weeds continues to occur, either in the presence of biocontrol agents or as an alternative to them. The ways in which the two techniques may interact are discussed, including direct toxicity of herbicides to biocontrol agents, responses to death of host plants and responses to sublethal changes caused by herbicides with different modes of action. A literature review for selected weed taxa showed that the great majority of publications relate to either chemical or to biological control techniques separately, with integration of the two seldom addressed. Possible reasons for this situation are discussed and some suggestions for future priorities are made.