The leaf-tying moth Hypocosmia pyrochroma (Lep., Pyralidae), a host-specific biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia

Cat's claw creeper, Macfadyena unguis-cati, a major environmental weed in coastal and sub-coastal areas of Queensland and New South Wales, Australia is a target for classical biological control. Host specificity of Hypocosmia pyrochroma Jones (Lep., Pyralidae), as a potential biological control agent was evaluated on the basis of no-choice and choice larval feeding and survival, and adult oviposition preference tests, involving 38 plant species in 10 families. In no-choice tests, larval feeding and development occurred only on cat's claw creeper. In choice tests, oviposition and larval development was evident only on cat's claw creeper. The results support the host-specificity tests conducted in South Africa, and suggest that H. pyrochroma is a highly specific biological control agent that does not pose any risk to non-target plants tested in Australia. This agent has been approved for field release by relevant regulatory authorities in Australia.

Improving the agent selection, release and evaluation process: the role of bioclimatic modelling

The value of CLIMEX models to inform biocontrol programs was assessed, including predicting the potential distribution of biocontrol agents and their subsequent population dynamics, using bioclimatic models for the weed Parkinsonia aculeata, two Lantana camara biocontrol agents, and five Mimosa pigra biocontrol agents. The results showed the contribution of data types to CLIMEX models and the capacity of these models to inform and improve the selection, release and post release evaluation of biocontrol agents. Foremost among these was the quality of spatial and temporal information as well as the extent to which overseas range data samples the species’ climatic envelope. Post hoc evaluation and refinement of these models requires improved long-term monitoring of introduced agents and their dynamics at well selected study sites. The authors described the findings of these case studies, highlighted their implications, and considered how to incorporate models effectively into biocontrol programs.

Biology and host range of Ophiomyia camarae Spencer (Diptera: Agromyzidae), a potential biocontrol agent for Lantana spp. (Verbenaceae) in Australia

The life history and host range of the herringbone leaf-mining fly Ophiomyia camarae, a potential biological control agent for Lantana spp., were investigated. Eggs were deposited singly on the underside of leaves. Although several eggs can be laid on a single leaf and a maximum of three individual mines were seen on a single leaf, only one pupa per leaf ever developed. The generation time (egg to adult) was about 38 days. Females (mean 14 days) lived longer than males (mean 9 days) and produced about 61 mines. Oviposition and larval development occurred on all five lantana phenotypes tested. Eleven plant species representing six families were tested to determine the host range. Oviposition and larval development occurred on only lantana and another nonnative plant Lippia alba (Verbenaceae), with both species supporting populations over several generations. A CLIMEX model showed that most of the coastal areas of eastern Australia south to 30°16' S (Coffs Harbour) would be suitable for O. camarae. O. camarae was approved for release in Australia in October 2007 and mines have been observed on plants at numerous field sites along the coast following releases.

Specificity of Carvalhotingis visenda (Hemiptera: Tingidae) as a biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia

Biological control is considered the most suitable management option for cat's claw creeper, Macfadyena unguis-cati, a major environmental weed in coastal and sub-coastal areas of Queensland and New South Wales, Australia. The potential host range of the leaf-sucking bug, Carvalhotingis visenda (Hemiptera: Tingidae) was evaluated on the basis of nymphal survival and development, adult feeding and survival, and oviposition preference using choice and no-choice tests involving 38 plant species in 10 families. In no-choice tests, although adults survived on a few of the non-target plants, no eggs were laid on any of the non-target plants. In no-choice condition, the tingid oviposits and completes nymphal development only on M. unguis-cati. There was also no visible feeding damage on any of the non-target plants. In choice tests, adults showed distinct preference for M. unguis-cati, and the preference level increased over time as the tingids moved away from the non-target plants. At the end of the trial no adults were evident on any of the non-target plants. Host specificity tests confirm that the tingid is a highly host specific biocontrol agent, and does not pose risk to any non-target plants in Australia. This agent has been approved for field release by the relevant regulatory authorities in Australia.

The host range and biology of Cometaster pyrula; a biocontrol agent for Acacia nilotica subsp. indica in Australia

Prickly acacia, Acacia nilotica subsp. indica (Benth.) Brenan, a major weed of the Mitchell Grass Downs of northern Queensland, Australia, has been the target of biological control projects since the 1980s. The leaf-feeding caterpillar Cometaster pyrula (Hopffer) was collected from Acacia nilotica subsp. kraussiana (Benth.) Brenan during surveys in South Africa to find suitable biological control agents, recognised as a potential agent, and shipped into a quarantine facility in Australia. Cometaster pyrula has a life cycle of approximately 2 months during which time the larvae feed voraciously and reach 6 cm in length. Female moths oviposit a mean of 339 eggs. When presented with cut foliage of 77 plant species, unfed neonates survived for 7 days on only Acacia nilotica subsp. indica and Acacia nilotica subsp. kraussiana. When unfed neonates were placed on potted plants of 14 plant species, all larvae except those on Acacia nilotica subsp. indica and Acacia nilotica subsp. kraussiana died within 10 days of placement. Cometaster pyrula was considered to be highly host specific and safe to release in Australia. Permission to release C. pyrula in Australia was obtained and the insect was first released in north Queensland in October 2004. The ecoclimatic model CLIMEX indicated that coastal Queensland was climatically suitable for this insect but that inland areas were only marginally suitable.

A systematic approach to biological control agent exploration and prioritisation for prickly acacia (Acacia nilotica ssp. indica)

Agent selection for prickly acacia has been largely dictated by logistics and host specificity. Given that detailed ecological information is available on this species in Australia, we propose that it is possible to select agents based on agent efficacy and desired impact on prickly acacia demography. We propose to use the 'plant genotype' and 'climatic' similarities as filters to identify areas for future agent exploration; and plant response to herbivory and field host range as 'predictive' filters for agent prioritisation. Adopting such a systematic method that incorporates knowledge from plant population ecology and plant-herbivore interactions makes agent selection decisions explicit and allow more rigorous evaluations of agent performance and better understanding of success and failure of agents in weed biological control.

Temporal patterns in incidence and abundance of Aconophora compressa (Hemiptera: Membracidae), a biological control agent for Lantana camara, on target and nontarget plants

The membracid Aconophora compressa Walker, a biological control agent released in 1995 to control Lantana camara (Verbenaceae) in Australia, has since been collected on several nontarget plant species. Our survey suggests that sustained populations of A. compressa are found only on the introduced nontarget ornamental Citharexylum spinosum (Verbenaceae) and the target weed L. camara. It is found on other nontarget plant species only when populations on C. spinosum and L. camara are high, suggesting that the presence of populations on nontarget species may be a spill-over effect. Some of the incidence and abundance on nontarget plants could have been anticipated from host specificity studies done on this agent before release, whereas others could not. This raises important issues about predicting risks posed by weed biological control agents and the need for long-term postintroduction monitoring on nontarget species.

Studies on the rust Prospodium tuberculatum, a new classical biological control agent released against the invasive alien weed Lantana camara in Australia. 2. Host range

A strain of the rust Prospodium tuberculatum from Brazil was screened as a potential biocontrol agent against 40 Australian Lantana camara forms and 52 closely related, non-target plant species. Results under glasshouse conditions showed that the Brazilian rust strain is pathogenic to only two flower colour forms: pink and pink-edged red. Macro- and microsymptoms were recorded using 11 assessment categories and four susceptibility ratings. No macrosymptoms were observed on any of the non-target plants.

A scientific approach to agent selection

The prioritisation of potential agents on the basis of likely efficacy is an important step in biological control because it can increase the probability of a successful biocontrol program, and reduce risks and costs. In this introductory paper we define success in biological control, review how agent selection has been approached historically, and outline the approach to agent selection that underpins the structure of this special issue on agent selection. Developing criteria by which to judge the success of a biocontrol agent (or program) provides the basis for agent selection decisions. Criteria will depend on the weed, on the ecological and management context in which that weed occurs, and on the negative impacts that biocontrol is seeking to redress. Predicting which potential agents are most likely to be successful poses enormous scientific challenges. 'Rules of thumb', 'scoring systems' and various conceptual and quantitative modelling approaches have been proposed to aid agent selection. However, most attempts have met with limited success due to the diversity and complexity of the systems in question. This special issue presents a series of papers that deconstruct the question of agent choice with the aim of progressively improving the success rate of biological control. Specifically they ask: (i) what potential agents are available and what should we know about them? (ii) what type, timing and degree of damage is required to achieve success? and (iii) which potential agent will reach the necessary density, at the right time, to exert the required damage in the target environment?

How widespread is Parthenium hysterophorus and its biological control agent Zygogramma bicolorata in South Asia?

Parthenium hysterophorus is a weed of global significance causing severe economic, environmental, human and animal health problems in Asia, Africa, Australia and the Pacific. In South Asia, P. hysterophorus occurs in India, Pakistan, Sri Lanka, Bangladesh and Nepal. A host-specific leaf-feeding beetle Zygogramma bicolorata from Mexico was introduced into India in 1984, as a biological control agent for P. hysterophorus. In this study, a GIS-based distribution map of P. hysterophorus and its biological control agent Z. bicolorata in South Asia based on meta-analysis is presented. The map highlights the limited published information on P. hysterophorus incidence in many of the states and territories in India, as well as in neighbouring Bangladesh, Bhutan, Nepal and Pakistan. Incidence of Z. bicolorata was recorded as three distinct clusters, covering many states in India. In Pakistan, Z. bicolorata was recorded in the Punjab region bordering India. A CLIMEX model based on the current distribution of Z. bicolorata in India suggests that the geographic range of this agent in India and Pakistan can extend to other P. hysterophorus-infested areas in the region. The CLIMEX model also suggests that all of Bangladesh and Sri Lanka, and parts of Nepal are climatically suitable for Z. bicolorata.

Thermal tolerance and potential distribution of Carvalhotingis visenda (Hemiptera: Tingidae), a biological control agent for cat's claw creeper, Macfadyena unguis-cati (Bignoniaceae)

The specialist tingid, Carvalhotingis visenda, is a biological control agent for cat's claw creeper, Macfadyena unguis-cati (Bignoniaceae). Cat's claw creeper is an invasive liana with a wide climatic tolerance, and for biological control to be effective the tingid must survive and develop over a range of temperatures. We evaluated the effect of constant temperatures (0-45°C) on the survival and development of C. visenda. Adults showed tolerance for wider temperature ranges (0-45°C), but oviposition, egg hatching and nymphal development were all affected by both high (>30°C) and low (<20°C) temperatures. Temperatures between 20°C and 30°C are the most favourable for adult survival, oviposition, egg hatching and nymphal development. The ability of adults and nymphs to survive for a few days at high (40°C and 45°C) and low (0°C and 5°C) temperatures suggest that extreme temperature events, which usually occur for short durations (hours) in cat's claw creeper infested regions in Queensland and New South Wales states are not likely to affect the tingid population. The potential number of generations (egg to adult) the tingid can complete in a year in Australia ranged from three to eight, with more generations in Queensland than in New South Wales.

Introduction and establishment of Carvalhotingis visenda (Hemiptera: Tingidae) as a biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia

Carvalhotingis visenda (Hemiptera: Tingidae) is the first biological control agent approved for release against cat’s claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia. The mass-rearing and field releases of C. visenda commenced in May 2007 and since then more than half a million individuals have been released at 72 sites in Queensland and New South Wales. In addition, community groups have released over 11,000 tingid-infested potted cat’s claw creeper plants at 63 sites in Queensland. Establishment of C. visenda was evident at 80% of the release sites after three years. The tingid established on the two morphologically distinct ‘long-pod’ and ‘short-pod’ cat’s claw creeper varieties present in Australia. Establishment was more at sites that received three or more field releases (83%) than at sites that received two or less releases (73%); and also at sites that received more than 5000 individuals (82%) than at sites that received less than 5000 individuals (68%). In the field, the tingid spread slowly (5.4 m per year), and the maximum distance of C. visenda incidence away from the initial release points ranged from 6 m to approximately 1 km.

Refining the process of agent selection through understanding plant demography and plant response to herbivory

Understanding plant demography and plant response to herbivory is critical to the selection of effective weed biological control agents. We adopt the metaphor of 'filters' to suggest how agent prioritisation may be improved to narrow our choices down to those likely to be most effective in achieving the desired weed management outcome. Models can serve to capture our level of knowledge (or ignorance) about our study system and we illustrate how one type of modelling approach (matrix models) may be useful in identifying the weak link in a plant life cycle by using a hypothetical and an actual weed example (Parkinsonia aculeata). Once the vulnerable stage has been identified we propose that studying plant response to herbivory (simulated and/or actual) can help identify the guilds of herbivores to which a plant is most likely to succumb. Taking only potentially effective agents through the filter of host specificity may improve the chances of releasing safe and effective agents. The methods we outline may not always lead us definitively to the successful agent(s), but such an empirical, data-driven approach will make the basis for agent selection explicit and serve as testable hypotheses once agents are released.

Relative host plant species use by the lantana biological control agent Aconophora compressa (Membracidae) across its native and introduced ranges

Aconophora compressa Walker (Hemiptera: Membracidae) was released in 1995 against the weed lantana in Australia, and is now found on multiple host plant species. The intensity and regularity at which A. compressa uses different host species was quantified in its introduced Australian range and also its native Mexican range. In Australia, host plants fell into three statistically defined categories, as indicated by the relative rates and intensities at which they were used in the field. Fiddlewood (Citharexylum spinosum L.: Verbenaceae) was used much more regularly and at higher densities than any other host sampled, and alone made up the first group. The second group, lantana (Lantana camara L.: Verbenaceae; pink variety) and geisha girl (Duranta erecta L.: Verbenaceae), were used less regularly and at much lower densities than fiddlewood. The third group, Sheena’s gold (another variety of D. erecta), jacaranda (Jacaranda mimosifolia D. Don: Bignoniaceae) and myoporum (Myoporum acuminatum R. Br.: Myoporaceae), were used infrequently and at even lower densities. In Mexico, the insect was found at relatively low densities on all hosts relative to those in Australia. Densities were highest on L. urticifolia, D. erecta and Tecoma stans (L.) Juss. ex Kunth (Bignoniaceae), which were used at similar rates to one another. It was found also on a few other verbenaceous and non-verbenaceous host species but at even lower densities. The relative rate at which Citharexylum spp. and L. urticifolia were used could not be assessed in Mexico because A. compressa was found on only one plant of each species in areas where these host species co-occurred. The low rate at which A. compressa occurred on fiddlewood in Mexico is likely to be an artefact of the short-term nature of the surveys or differences in the suites of Citharexylum and Lantana species available there. These results provide further incentive to insist on structured and quantified surveys of non-target host use in the native range of potential biological control agents prior to host testing studies in quarantine.