Prospects for the biological control of the weedy sporobolus grasses in Australia.

Bill Palmer and colleagues recently published their paper 'Prospects for the biological control of the weedy sporobolus grasses in Australia' in Proceedings of the 16th Australian Weeds Conference. The paper gives a summary of a recent project to find a biological control for the weedy sporobolus grasses, which include giant rat's tail grass. Southern Africa was surveyed for potential agents and two, a leaf smut and a stem wasp, were selected for follow up studies. Unfortunately, they could not rear the stem wasp in the laboratory and the leaf smut infected four of the Australian native Sporobolus spp. and was therefore rejected. This project was one of the first attempts at biological control of a grass.

Applications of CLIMEX modelling leading to improved biological control

Wilmot Senaratne, Bill Palmer and Bob Sutherst recently published their paper 'Applications of CLIMEX modelling leading to improved biological control' in Proceedings of the 16th Australian Weeds Conference. They looked at three examples where modern climate matching techniques using computer software produces decisions and results than might happen using previous techniques such as climadiagrams. Assessment of climatic suitability is important at various stages of a biological control project; from initial foreign exploration, to risk assessment in preparation for the release of a particular agent, through to selection of release sites that maximise the agent´s chances of initial establishment. It is now also necessary to predict potential future distributions of both target weeds and agents under climate change.

Factors influencing the release and establishment of weed biological control agents

Mike Day and colleagues recently published their paper 'Factors influencing the release and establishment of weed biocontrol agents' in Proceedings of the 16th Australian Weeds Conference. The CRC for Australian Weed Management facilitated an investigation into the factors influencing the release and establishment of weed biological control agents on a wide variety of Australian weeds. The investigation improved the understanding of post-release ecology of biocontrol agents and generated recommendations for best practice. Factors affecting successful establishment on the weed include host plant characteristics, size of releases, dispersal power of the agent, predation and parasitism, and climate. A best practice guide was produced by the CRC to assist practitioners in designing robust release strategies to increase rates of establishment.

Cassowary dispersal of the invasive pond apple in a tropical rainforest: the contribution of subordinate dispersal modes in invasion.

Dispersal is a significant determinant of the pattern and process of invasions; however, weed dispersal distances are rarely described and descriptions of dispersal kernels are completely lacking for vertebrate-dispersed weeds. Here, we describe dispersal kernels generated by a native disperser, the endangered southern cassowary (Casuarius casuarius, L.) for an invasive, tropical rainforest plant, pond apple (Annona glabra, L.). Pond apple is primarily water-dispersed and is managed as such. We consider whether cassowary dispersal, as a numerically subordinate dispersal mode, provides an additional dispersal service that may modify the invasion process. In infested areas, pond apple seed was common in cassowary dung. Gut passage had no effect on the probability of single seed germination but deposition in clumps or as whole fruits reduced the probability of germination below that of single seeds. Gut passage times ranged from 65 to 1675 min. Combined with cassowary movement data, this resulted in estimated dispersal distances of 12.5-5212 m, with a median distance of 387 m (quartile range 112-787 m). Native frugivores can be effective dispersers of weeds in rainforest and even terrestrial dispersers can provide long-distance dispersal. Importantly, though pond apple might be expected to be almost entirely dispersed downstream and along the margins of aquatic and marine habitats, cassowaries provide dispersal upstream and between drainages, leading to novel dispersal outcomes. Even through the provision of small quantities of novel dispersal outcomes, subordinate dispersal modes can play a significant role in determining invasion pattern and influence the ultimate success of control programs by providing dispersal to locations unattainable via the primary mode.

Inter-seasonal population dynamics and pest status of Bemisia tabaci (Gennadius) biotype B in an Australian cropping system

Bemisia tabaci, biotype B, commonly known as the silverleaf whitefly (SLW) is an alien species that invaded Australia in the mid-90s. This paper reports on the invasion ecology of SLW and the factors that are likely to have contributed to the first outbreak of this major pest in an Australian cotton cropping system, population dynamics of SLW within whitefly-susceptible crop (cotton and cucurbit) and non-crop vegetation (sowthistle, Sonchus spp.) components of the cropping system were investigated over four consecutive growing seasons (September-June) 2001/02-2004/05 in the Emerald Irrigation Area (EIA) of Queensland, Australia. Based on fixed geo-referenced sampling sites, variation in spatial and temporal abundance of SLW within each system component was quantified to provide baseline data for the development of ecologically sustainable pest management strategies. Parasitism of large (3rd and 4th instars) SLW nymphs by native aphelinid wasps was quantified to determine the potential for natural control of SLW populations. Following the initial outbreak in 2001/02, SLW abundance declined and stabilised over the next three seasons. The population dynamics of SLW is characterised by inter-seasonal population cycling between the non-crop (weed) and cotton components of the EIA cropping system. Cotton was the largest sink for and source of SLW during the study period. Over-wintering populations dispersed from weed host plant sources to cotton in spring followed by a reverse dispersal in late summer and autumn to broad-leaved crops and weeds. A basic spatial source-sink analysis showed that SLW adult and nymph densities were higher in cotton fields that were closer to over-wintering weed sources throughout spring than in fields that were further away. Cucurbit fields were not significant sources of SLW and did not appear to contribute significantly to the regional population dynamics of the pest. Substantial parasitism of nymphal stages throughout the study period indicates that native parasitoid species and other natural enemies are important sources of SLW mortality in Australian cotton production systems. Weather conditions and use of broad-spectrum insecticides for pest control are implicated in the initial outbreak and on-going pest status of SLW in the region.

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.

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.

Will biological control of Lantana camara ever succeed? Patterns, processes & prospects

Despite biocontrol research spanning over 100 years, the hybrid weed, commonly referred to as Lantana camara, is not under adequate control. Host specificity and varietal preference of released agents, climatic suitability of a region for released agents, number of agents introduced and range or area of infestation appear to play a role in limiting biocontrol success. At least one of 41 species of mainly leaf- or flower-feeding insects has been introduced, or spread, to 41 of the 70 countries or regions where lantana occurs. Over half (26) of these species have established, achieving varying levels of herbivory and presumably some degree of control. Accurate taxonomy of the plant and adaptation of potential agents to the host plant are some of the better predictors of at least establishment success. Retrospective analysis of the hosts of introduced biocontrol agents for L. camara show that a greater proportion of agents that were collected from L. camara or Lantana urticifolia established, than agents that were collected from other species of Lantana. Of the introduced agents that had established and were oligophagous, 18 out of 22 established. The proportion of species establishing, declined with the number of species introduced. However, there was no trend when oceanic islands were treated separately from mainland areas and the result is likely an artefact of how introductions have changed over time. A calculated index of the degree of herbivory due to agents known to have caused some damage per country, was not related to land area infested with lantana for mainlands nor for oceanic islands. However, the degree of herbivory is much higher on islands than mainlands. This difference between island and mainland situations may reflect population dynamics in patchy or metapopulation landscapes. Basic systematic studies of the host remain crucial to successful biocontrol, especially of hybrid weeds like L. camara. Potential biocontrol agents should be monophages collected from the most closely related species to the target weed or be phytophages that attack several species of lantana. Suitable agents should be released in the most ideal ecoclimatic area. Since collection of biocontrol agents has been limited to a fraction of the known number of phytophagous species available, biocontrol may be improved by targeting insects that feed on stems and roots, as well as the agents that feed on leaves and flowers.

Predicting risk and benefit a priori in biological control of invasive plant species: a systems modelling approach

While the method using specialist herbivores in managing invasive plants (classical biological control) is regarded as relatively safe and cost-effective in comparison to other methods of management, the rarity of strict monophagy among insect herbivores illustrates that, like any management option, biological control is not risk-free. The challenge for classical biological control is therefore to predict risks and benefits a priori. In this study we develop a simulation model that may aid in this process. We use this model to predict the risks and benefits of introducing the chrysomelid beetle Charidotis auroguttata to manage the invasive liana Macfadyena unguis-cati in Australia. Preliminary host-specificity testing of this herbivore indicated that there was limited feeding on a non-target plant, although the non-target was only able to sustain some transitions of the life cycle of the herbivore. The model includes herbivore, target and non-target life history and incorporates spillover dynamics of populations of this herbivore from the target to the non-target under a variety of scenarios. Data from studies of this herbivore in the native range and under quarantine were used to parameterize the model and predict the relative risks and benefits of this herbivore when the target and non-target plants co-occur. Key model outputs include population dynamics on target (apparent benefit) and non-target (apparent risk) and fitness consequences to the target (actual benefit) and non-target plant (actual risk) of herbivore damage. The model predicted that risk to the non-target became unacceptable (i.e. significant negative effects on fitness) when the ratio of target to non-target in a given patch ranged from 1:1 to 3:2. By comparing the current known distribution of the non-target and the predicted distribution of the target we were able to identify regions in Australia where the agent may be pose an unacceptable risk. By considering risk and benefit simultaneously, we highlight how such a simulation modelling approach can assist scientists and regulators in making more objective decisions a priori, on the value of releasing specialist herbivores as biological control agents.

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.

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.

Review of approaches to evaluate the effectiveness of weed biological control agents

We review key issues, available approaches and analyses to encourage and assist practitioners to develop sound plans to evaluate the effectiveness of weed biological control agents at various phases throughout a program. Assessing the effectiveness of prospective agents before release assists the selection process, while post-release evaluation aims to determine the extent that agents are alleviating the ecological, social and economic impacts of the weeds. Information gathered on weed impacts prior to the initiation of a biological control program is necessary to provide baseline data and devise performance targets against which the program can subsequently be evaluated. Detailed data on weed populations, associated plant communities and, in some instances ecosystem processes collected at representative sites in the introduced range several years before the release of agents can be compared with similar data collected later to assess agent effectiveness. Laboratory, glasshouse and field studies are typically used to assess agent effectiveness. While some approaches used for field studies may be influenced by confounding factors, manipulative experiments where agents are excluded (or included) using chemicals or cages are more robust but time-consuming and expensive to implement. Demographic modeling and benefit–cost analyses are increasingly being used to complement other studies. There is an obvious need for more investment in long-term post-release evaluation of agent effectiveness to rigorously document outcomes of biological control programs.

The host range of Isturgia deeraria, an insect considered for the biological control of Acacia nilotica in Australia

The geometrid caterpillar Isturgia deerraria was imported from Kenya into quarantine facilities in Australia as a potential biological control agent for prickly acacia, Acacia nilotica subsp. indica (Benth.) Brenan (family Mimosaceae). The insect was then tested on 30 plant species presented to neonate larvae as a no-choice cut foliage test and 13 plant species presented as a no-choice potted plant test. In these tests the insect was able to complete its life cycle on 13 native Acacia spp. and also on Acacia farnesiana and the exotic ornamental Delonix regia (family Caesalpiniaceae). The tests supported field observations that the insect has a host range spanning many leguminous species and as such the insect could not be considered for release in Australia.