Is host‐specificity of weed biological control agents likely to evolve rapidly following establishment?

Over 352 herbivore species have now been intentionally introduced into new regions as weed biological control agents. Recent evidence shows that rapid and significant evolution in host-specificity can occur. The risk of non-target use by biological control agents increasing to unacceptable levels through rapid evolution therefore needs to be considered. In addition, weed biological control offers many as yet largely unexploited opportunities for improving our basic understanding of host-specificity and its evolution. We therefore evaluate the evidence that rapid evolution (1) alters the use of existing hosts, and (2) alters the fundamental host-range. Most cited examples of so-called host shifts from weed biological control were not the result of genetic change. There was only limited evidence of genetically altered performance on a non-target host and no evidence of altered fundamental host-range. We conclude, from both theory and the available data, that only altered use of existing hosts (through quantitative genetic changes) needs be considered when evaluating the risk of rapid evolution. Host-specificity testing methodologies can be improved and adapted to better assess the risk of occurrence of post-release evolution.     

杂草生防作用物对本土生物群落的影响

传统生物防治是治理外来入侵杂草危害切实可行的有效策略和途径,近来对传统生物防治的批评主要集注于,引进的生防作用物攻击威胁本土非靶标生物。引进的生防作用物可能对本土非靶标生物产生直接和间接影响,这类影响通过不同营养级生物之间的取食关系,以及通过同一营养级内生物间的竞争关系,而影响本土非靶标生物群落。列举若干杂草生物防治案例对以上影响方式及其发生途径进行了评述。就防范杂草生防作用物对非靶标生物的负面影响,提出了以下对策:(i)把引进天敌防治外来入侵生物作为最后的有效手段;(ii)适当增加对非靶标生物潜在影响的生态学评估;(iii)选择寄主专一性强而且能有效控制靶标杂草的天敌;(iv)加强对杂草传统生物防治的生态学研究。     

Pre-release evaluation of the efficacy of the leaf-sucking bug Carvalhotingis visenda (Heteroptera: Tingidae) as a biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae)

In classical weed biological control, assessing weed response to simulated herbivory is one option to assist in the prioritization of available agents and prediction of their potential efficacy. Previously reported simulated herbivory studies suggested that a specialist herbivore in the leaf-feeding guild is desirable as an effective biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae), an environmental weed that is currently a target for biological control. In this study, we tested (i) whether the results from glasshouse-based simulated herbivory can be used to prioritise potential biological control agents by evaluating the impact of a leaf-sucking tingid bug Carvalhotingis visenda (Drake & Hambleton) (Hemiptera: Tingidae) in quarantine; and (ii) the likely effectiveness of low- and high-densities of the leaf-sucking tingid after its release in the field. The results suggest that a single generation of C. visenda has the potential to reduce leaf chlorophyll content significantly, resulting in reduced plant height and leaf biomass. However, the impact of one generation of tingid herbivory on below-ground plant components, including the roots and tuber size and biomass, were not significant. These findings are consistent with results obtained from a simulated herbivory trial, highlighting the potential role of simulated herbivory studies in agent prioritisation.     

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.     

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.     

Integrating Management Techniques to Restore Sites Invaded by Mile‐a‐Minute Weed,Persicaria perfoliata

Efforts to suppress an invasive weed are often undertaken with the goal of facilitating the recovery of a diverse native plant community. In some cases, however, reduction in the abundance of the target weed results in an increase in other exotic weeds. Mile‐a‐minute weed (Persicaria perfoliata (L.) H. Gross (Polygonaceae)) is an annual vine from Asia that has invaded the eastern United States, where it can form dense monocultures. The host‐specific Asian weevil Rhinoncomimus latipes Korotyaev (Coleoptera: Curculionidae) was first released in the United States in 2004 as part of a classical biological control program. At three sites invaded by mile‐a‐minute weed, biological control was integrated with pre‐emergent herbicide use and two densities of native plantings. After 2 years, native plant cover differed significantly and was greater than 80% in the plots with plantings and pre‐emergent herbicide but less than 30% in the planting treatments without herbicide. Where mile‐a‐minute cover decreased at the two sites with the greatest pressure from exotic plants, plots were dominated by another exotic weed, Microstegium vimineum (Trin.) A. Camus, Japanese stiltgrass. The combination of biocontrol, pre‐emergent herbicide, and revegetation with native plants suppressed mile‐a‐minute weed, prevented invasion by Japanese stiltgrass, and increased the abundance of native plants. The selection of the management strategies used to control mile‐a‐minute weed determined the extent of recovery of the native plant community.     

Benefits of Classical Biological Control for Managing Invasive Plants

As a result of the rapid expansion in international travel and trade over the past few decades, invasive plants have become a problem of global proportions. Plant invasions threaten the existence of endangered species and the integrity of ecosystems, and their ravages cost national economies tens of billions of dollars every year. Strategies for managing the threats posed by plant invasions involve three main tactics: prevention, eradication, and control. The effectiveness of prevention, involving enactment of legislation to prohibit the entry and spread of noxious alien plants, has been questioned. Eradication of all but the smallest, most localized weed infestations generally is not regarded as economically feasible. Conventional weed control techniques, such as mechanical and chemical controls, because they are expensive, energy and labor intensive, and require repeated application, are impractical for managing widespread plant invasions in ecologically fragile conservation areas or low-value habitat, such as rangelands and many aquatic systems. In addition, mechanical means of control disturb the soil and may cause erosion; chemical herbicides have spurred the evolution of resistance in scores of weed species and, further, may pose risks to wildlife and human health. Because of drawbacks associated with conventional weed control methods, classical biological control, the introduction of selective exotic natural enemies to control exotic pests, increasingly is being considered and implemented as a safe, cost-effective alternative to address the invasive plant problem. Worldwide, biological weed control programs have had an overall success rate of 33 percent; success rates have been considerably higher for programs in individual countries. Benefits are several-fold. Biological control is permanent, energy-efficient, nonpolluting, and inexpensive relative to other methods. Economic returns on investment in biological weed control have been spectacular in some cases, and range from an estimated benefit/cost ratio of 2.3 to 4000 or more. Although the risks involved in biological control in general are considered unacceptable by some, biological weed control in particular has had an enviable safety record. Since establishment of the stringent standards and regulatory apparatus currently in place in the United States and elsewhere, there have been no reported cases of biological weed control causing significant harm to nontarget populations or to the environment at large.     

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.     

Field observations on the biology and behaviour of Dasiops caustonae Norrbom and McAlpine (Dipt., Lonchaeidae), as a candidate biocontrol agent of Passiflora mollissima in Hawaii

A newly described lonchaeid from Venezuela, Dasiops caustonae Norrbom and McAlpine, has been recommended as a biological control agent of Passiflora mollissima (H.B.K) Bailey, an aggressive exotic weed of Hawaiian rainforests. In this study, elementary biological information was collected in the field in order to determine the appropriateness of using D. caustonae as a biological control agent against this important weed. Field observations suggested that the host range of D. caustonae is limited to species of the subgenus Tacsonia. Female flies appear to mark flowers with an oviposition deterrent pheromone. This is possibly the first report of marking behaviour outside Tephritidae. The geographical range of this species is wide and comparable with that of the target weed. Life‐cycle studies demonstrated that D. caustonae is multivoltine with an estimated life‐cycle length of 3 to 4 months. Survival to adult was limited by premature flower fall, drought and interspecific competition by a bud and flower feeding Lepidoptera, Pyrausta perelegans Hampson. Additional ecological studies and host range testing of D. caustonae are recommended to determine its impact on the target weed and its safety as a biological control agent.     

An economic assessment of the contribution of biological control to the management of invasive alien plants and to the protection of ecosystem services in South Africa

This study is a first attempt at a holistic economic evaluation of South African endeavours to manage invasive alien plants using biological control. Our focus was on the delivery of ecosystem services from habitats that are invaded by groups of weeds, rather than by each individual weed species. We established the net present value of the weed biological control efforts, and derived benefit:cost ratios by comparing this value (a cost) to the estimated value of ecosystem services protected by weed biological control. We identified four major functional groupings of invading alien plants, and assessed their impact on water resources, grazing and biodiversity. We estimated the area that remained free of invasions due to all historic control efforts in South Africa, and the proportion that remained free of invasion as a result of biological control (which was initiated in 1913). The estimated value of potential ecosystem services amounted to 152 billion South African rands (ZAR—presently, about US$ 19.7 billion) annually. Although an estimated ZAR 6.5 billion was lost every year due to invading alien plants, this would have amounted to an estimated additional ZAR 41.7 billion had no control been carried out, and 5–75% of this protection was due to biological control. The benefit:cost ratios ranged from 50:1 for invasive sub-tropical shrubs to 3,726:1 for invasive Australian trees. Benefit:cost ratios remained positive and our conclusion, that biological control has brought about a considerable level of protection of ecosystem services, remains robust even when our estimates of the economic impacts of key variables (i.e. sensitivity analyses of indeterminate variables) were substantially reduced.     

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.     

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 .     

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.     

Making host specificity testing more efficient: Exploring the use of abridged test plant lists

Testing the specificity of candidate agents is a key component of risk analysis in weed biological control. This step is often time-consuming due to the numerous plant species that need to be tested under quarantine conditions in the invaded country of the weed species. Here, we examined whether an abridged phylogenetically based test list could be used in the weed's native range to quickly screen the host specificity of candidate agents. Ten plant species were used to test the host specificity of a promising candidate for the biological control of Sonchus oleraceus in Australia, the gall midge, Cystiphora sonchi. No-choice and choice tests were carried out in the native Mediterranean range of the midge. The results showed the midge has potential to threaten native Australian species, as those species showed high infestation levels in no-choice tests and produced significantly higher numbers of galls in choice tests. As a result of this approach, C. sonchi was rapidly discarded from the list of agents to be imported into Australian quarantines for further tests. This study demonstrates that testing a few key phylogenetically related species in the native range may save cost and effort in a weed biological control programme.     

Distribution of Insect Attacks in Biological Control of Weeds: Infestation of Centaurea virgata Flowerheads by a Gall Fly

The success of biological control efforts to reduce weed density through release of insects may depend as much on the distribution of insect attacks among individual plants or plant parts as on the mean level of infestation. We used an index of dispersion to describe the distribution of Urophora quadrifasciata (Diptera: Tephritidae) galls among squarrose knapweed (Centaurea virgata) flowerheads at 18 west central Utah sites in the first 5 years following introduction of the biological control agent. Two thirds of the samples showed a significantly aggregated distribution of galls among flowerheads. Statistical analysis showed that site and year accounted for relatively small proportions of the variance in the index of dispersion. The degree of gall aggregation among flowerheads was positively correlated with the mean flowerhead quality (mean number of seeds per flowerhead; P = 0.013) and tended to be negatively correlated with the mean fly density per flowerhead at a site in a given year (P = 0.097). Our data suggest that higher quality flowerheads, and possibly higher quality plants, are preferentially attacked by U. quadrifasciata and therefore are more heavily subject to reduced reproductive potential through biological control. However, an aggregated distribution of fly attacks may undercut the potential of the fly to reduce seed production by the weed population as a whole. Understanding both the distribution of insect attacks among individual plants and the behavioral mechanisms producing such distribution patterns is important to the biological control of weeds.     

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.     

Complementing biological control with plant suppression: Implications for improved management of parthenium weed (Parthenium hysterophorus L.)

Parthenium hysterophorus L. is a weed of global significance that has become a major weed in Australia and many other parts of the world. A combined approach for the management of parthenium weed using biological control and plant suppression, was tested under field conditions over a two-year period in southern central Queensland. The six suppressive plant species, selected for their demonstrably suppressive ability in earlier glasshouse studies, worked synergistically with the biological control agents (Epiblema strenuana Walker, Zygogramma bicolorata Pallister, Listronotus setosipennis Hustache and Puccinia abrupta var. partheniicola) present in the field to reduce the growth (above ground biomass) of parthenium weed, by between 60–86% and 47–91%, in Years 1 and 2, respectively. The biomass of the suppressive plants was between 6% and 23% greater when biological control agents were present than when the biological control agents had been excluded. This shows that parthenium weed can be more effectively managed by combining the current biological control management strategy with selected sown suppressive plant species, both in Australia and elsewhere.     

The prickly pear (Opuntia ficus-indica [Cactaceae]) in South Africa: Utilization of the naturalized weed,and of the cultivated plants

In spite of successful biological control efforts, vast areas of South Africa remain infested with the spiny prickly pear weed, Opuntia ficus-indica. These remnant populations, which vary from sparse to dense and which are more or less stable, are confined mainly to the eastern Cape. They provide a popular fruit and a limited source of income for certain sectors of the population in these areas. In terms of existing legislation, there are, however, restrictions on the large-scale utilization of these fruit. A novel method for mass-rearing the cochineal insect, Dactylopius coccus, for the commercial production of a red dye (carminic acid) has been developed. The cultivated spineless prickly pear has recently been enjoying renewed attention from researchers and growers alike. Whereas it has traditionally been cultivated mainly as a droughtresistant fodder crop, and is still popular as such, it is now increasingly recognised as a fruit, in its own right, with considerable promise as a commercial crop for local and export markets. There is also a need to encourage the use of very young cladodes (nopalitos) as a highly nutritious vegetable for human consumption in South Africa.     

Using molecular methods to determine the origin of weed populations of Pereskia aculeata in South Africa and its relevance to biological control

New biological control agents are required in order to reach and sustain an adequate level of control of the declared environmental weed Pereskia aculeata Miller (Cactaceae) in South Africa. Identifying the origin of weed genotypes has been important in a number of biological control programmes and is likely to be of importance for the control of P. aculeata due to its disjunct native distribution and morphological polymorphisms between plants from different regions of the native and introduced distribution. DNA sequencing of the trnL chloroplastic intron and the phyC nuclear gene indicate that the South African weed population’s origin was in the southern region of native distribution. Inter-Simple Sequence Repeats (ISSRs) confirmed this result and added resolution to the analysis indicating that the native plants with the closest genetic distance to the South African weed population were found in Rio de Janeiro Province, Brazil. The relationship between the South African weed population and garden variety plants as well as the large genetic distance between the South African plants and the native plants suggests that the South African population may be the progeny of escaped garden variety plants that have been cultivated and possibly hybridized. The low levels of genetic variation within the South African population and the monophyly of the South African plants indicates that these plants are the progeny of a single introduction or multiple introductions from the same source. Rio de Janeiro Province in Brazil is the most appropriate region in which to survey for new biological control agents.     

Predicting the host range of Nystalea ebalea: Secondary plant chemistry and host selection by a surrogate biological control agent of Schinus terebinthifolia

The safety of weed biological control depends upon the selection and utilization of the target weed by the agent while causing minimal harm to non-target species. Selection of weed species by biological control agents is determined by the presence of behavioral cues, generally host secondary plant compounds that elicit oviposition and feeding responses. Non-target species that possess the same behavioral cues as found in the target weed may be at risk of damage by classical biological control agents. Here we conducted host range tests and examined secondary plant compounds of several test plant species. We studied the specialist herbivore Nystalea ebalea (Lepidoptera: Notodontidae) a Neotropical species, present in Florida as a surrogate biological control agent of the weed, Brazilian peppertree Schinus terebinthifolia, invasive in Florida and Hawaii. We found that the larvae had the greatest survival when fed the target weed, the Neotropical species Spondias purpurea, the Florida native species Rhus copallinum, and the ornamental Pistacia chinensis. Reduced survival and general larval performance were found on the native species Metopium toxiferum and Toxicodendron radicans. Both the volatiles and the allergen urushiols were chemically characterized for all species but urushiol diversity and concentration best predicted host range of this herbivore species. These results provide insight into host selection and utilization by one oligophagous Schinus herbivore. Other potential biological control agents may also be sensitive to plants that contain urushiols and if so, they may pose minimal risk to these native species.