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.

Selecting replacements for invasive plants to support frugivores in highly modified sites: a case study focusing on Lantana camara

Invasive plants are regarded as a major threat to biodiversity worldwide. Yet, in some cases, invasive plants now perform important ecological functions. For example, fleshy-fruited invasive plants provide food that supports indigenous frugivore populations. How can the disparate goals of conservation versus invasive weed control be managed? We suggest using the fruit characteristics of the invasive plant to select replacement indigenous plants that are functionally similar from the perspective of frugivores. These could provide replacement food resources at sites where plants with these characteristics are part of the goal plant community and where such plants would not otherwise regenerate. Replacement plants could also redirect seed dispersal processes to favour indigenous, rather than invasive, plant species. We investigated the utility of this approach by ranking all indigenous fleshy-fruited plant species from a region using a simple model that scored species based upon measures of fruit phenology, morphology, conspicuousness and accessibility relative to a target invasive species, Lantana (Lantana camara). The model successfully produced high scores for indigenous plant species that were used by more of the frugivores of Lantana than a random selection of plants, suggesting that this approach warrants further investigation.

Reproductive ecology of invasive Ochna serrulata (Ochnaceae) in south-eastern Queensland

We investigated aspects of the reproductive ecology of Ochna serrulata (Hochst.) Walp., an invasive plant in eastern Australia. O. serrulata drupes were similar in size to fleshy fruits of other local invasive plants, but showed some distinct differences in quality, with a very high pulp lipid content (32.8% of dry weight), and little sugar and water. Seeds were dispersed by figbirds, Sphecotheres viridis Vieillot, a locally abundant frugivore, and comprised between 10 and 50% of all non-Ficus spp. fruit consumed during October and November. The rate of removal of O. serrulata drupes was greater in bushland than suburban habitats, indicating that control in bushland habitats should be a priority, but also that suburban habitats are likely to act as significant seed sources for reinvasion of bushland. Germination occurred under all seed-processing treatments (with and without pulp, and figbird gut passage), suggesting that although frugivores are important for dispersal, they are not essential for germination. Recruitment of buried and surface-sown seed differed between greenhouse and field experiments, with minimal recruitment of surface-sown seed in the field. Seed persistence was low, particularly under field conditions, with 0.75% seed viability after 6 months and 0% at 12 months. This provides an opportunity to target control efforts in south-eastern Queensland in spring before fruit set, when there is predicted to be few viable seeds in the soil.

Lantana (Lantana camara) Seed Bank Dynamics: Seedling Emergence and Seed Survival.

Seed persistence is poorly quantified for invasive plants of subtropical and tropical environments and Lantana camara, one of the world's worst weeds, is no exception. We investigated germination, seedling emergence, and seed survival of two lantana biotypes (Pink and pink-edged red [PER]) in southeastern Queensland, Australia. Controlled experiments were undertaken in 2002 and repeated in 2004, with treatments comprising two differing environmental regimes (irrigated and natural rainfall) and sowing depths (0 and 2 cm). Seed survival and seedling emergence were significantly affected by all factors (time, biotype, environment, sowing depth, and cohort) (P < 0.001). Seed dormancy varied with treatment (environment, sowing depth, biotype, and cohort) (P < 0.001), but declined rapidly after 6 mo. Significant differential responses by the two biotypes to sowing depth and environment were detected for both seed survival and seedling emergence (P < 0.001). Seed mass was consistently lower in the PER biotype at the population level (P < 0.001), but this variation did not adequately explain the differential responses. Moreover, under natural rainfall the magnitude of the biotype effect was unlikely to result in ecologically significant differences. Seed survival after 36 mo under natural rainfall ranged from 6.8 to 21.3%. Best fit regression analysis of the decline in seed survival over time yielded a five-parameter exponential decay model with a lower asymptote approaching −0.38 (% seed survival = [( 55 − (−0.38)) • e (k • t)] + −0.38; R2 = 88.5%; 9 df). Environmental conditions and burial affected the slope parameter or k value significantly (P < 0.01). Seed survival projections from the model were greatest for buried seeds under natural rainfall (11 yr) and least under irrigation (3 yr). Experimental data and model projections suggest that lantana has a persistent seed bank and this should be considered in management programs, particularly those aimed at eradication.

A review of current knowledge of the weedy species Themeda quadrivalvis (grader grass)

This review of grader grass (Themeda quadrivalvis) attempts to collate current knowledge and identify knowledge gaps that may require further research. Grader grass is a tropical annual grass native to India that is now spread throughout many of the tropical regions of the world. In Australia, it has spread rapidly since its introduction in the 1930s and is now naturalised in the tropical areas of Queensland, the Northern Territory and Western Australia and extends south along the east coast to northern New South Wales. It is a vigorous grass with limited palatability, that is capable of invading native and improved pastures, cropping land and protected areas such as state and national parks. Grader grass can form dense monocultures that reduce biodiversity, decrease animal productivity and increase the fire hazard in the seasonally dry tropics. Control options are based on herbicides, grazing management and slashing, while overgrazing appears to favour grader grass. The effect of fire on grader grass is inconclusive and needs to be defined. Little is known about the biology and impacts of grader grass in agricultural and protected ecosystems in Australia. In particular, information is needed on soil seed bank longevity, seed production, germination and growth, which would allow the development of management strategies to control this weedy grass.

Weeds of Australian rangelands

Despite recognition that non-native plant species represent a substantial risk to natural systems, there is currently no compilation of weeds that impact on the biodiversity of the rangelands within Australia. Using published and expert knowledge, this paper presents a list of 622 non-native naturalised species known to occur within the rangelands. Of these, 160 species (26%) are considered a current threat to rangeland biodiversity. Most of these plant species have been deliberately introduced for forage or other commercial use (e.g. nursery trade). Among growth forms, shrubs and perennial grasses comprise over 50% of species that pose the greatest risk to rangeland biodiversity. We identify regions within the rangelands containing both high biodiversity values and a high proportion of weeds and recommend these areas as priorities for weed management. Finally, we examine the resources available for weed detection and identification since detecting weeds in the early stages of invasion is the most cost effective method of reducing further impact.

Response of an invasive liana to simulated herbivory: implications for its biological control

Pre-release evaluation of the efficacy of biological control agents is often not possible in the case of many invasive species targeted for biocontrol. In such circumstances simulating herbivory could yield significant insights into plant response to damage, thereby improving the efficiency of agent prioritisation, increasing the chances of regulating the performance of invasive plants through herbivory and minimising potential risks posed by release of multiple herbivores. We adopted this approach to understand the weaknesses herbivores could exploit, to manage the invasive liana, Macfadyena unguis-cati. We simulated herbivory by damaging the leaves, stem, root and tuber of the plant, in isolation and in combination. We also applied these treatments at multiple frequencies. Plant response in terms of biomass allocation showed that at least two severe defoliation treatments were required to diminish this liana's climbing habit and reduce its allocation to belowground tuber reserves. Belowground damage appears to have negligible effect on the plant's biomass production and tuber damage appears to trigger a compensatory response. Plant response to combinations of different types of damage did not differ significantly to that from leaf damage. This suggests that specialist herbivores in the leaf-feeding guild capable of removing over 50% of the leaf tissue may be desirable in the biological control of this invasive species.

Metabolite mobilization in the stem galls of Parthenium hysterophorus induced by Epiblema strenuana inferred from the signatures of isotopic carbon and nitrogen and concentrations of total non-structural carbohydrates

Parthenium hysterophorus L. (Asteraceae) is a weed of national significance in Australia. Among the several arthropod agents introduced into Australia to control populations of P. hysterophorus biologically, Epiblema strenuana Walker (Lepidoptera: Tortricidae) is the most widespread and abundant agent. By intercepting the normal transport mechanisms of P. hysterophorus, the larvae of E. strenuana drain nutrients, other metabolic products, and energy, and place the host plant under intense metabolic stress. In this study, determinations of total non-structural carbohydrates (TNC) levels and carbon and nitrogen isotope ratios of fixed products in different parts of the plant tissue, including the gall, have been made to establish the function of gall as a sink for the nutrients. Values of δ13C and δ15N in galls were significantly different than those in proximal and distal stems, whereas the TNC levels were insignificant, when measured in the total population of P. hysterophorus, regardless of plant age. However, carbon, nitrogen, and TNC signatures presented significant results, when assayed in different developmental stages of P. hysterophorus. Carbon isotope ratios in galls were consistently more negative than those from the compared plant organs. Nitrogen isotope ratios in galls, on the contrary, were either similar to or less negative than the compared plant organs, especially within a single host-plant stage population (i.e., either rosette, preflowering, or flowering stage). TNC levels varied within compared plant populations. The stem distal to the gall functioned more efficiently as a nodal channel than the stem proximal to the gall, especially in the translocation of nitrogenous nutrients. Our findings indicate that the gall induced by E. strenuana functions as a sink for the assayed nutrients, although some variations have been observed in the patterns of nutrient mobilization. By creating a sink for the nutrients in the gall, E. strenuana is able to place the overall plant metabolism under stress, and this ability indicates E. strenuana has the necessary potential for use as a biological-control agent.

Use of CLIMEX modelling to identify prospective areas for exploration to find new biological control agents for prickly acacia

Selection of biocontrol agents that are adapted to the climates in areas of intended release demands a thorough analysis of the climates of the source and release sites. We present a case study that demonstrates how use of the CLIMEX software can improve decision making in relation to the identification of prospective areas for exploration for agents to control the woody weed, prickly acacia Acacia nilotica ssp. indica in the arid areas of north Queensland.

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?

Clinging on: a review on the biological control of cat’s claw creeper

Cat's claw creeper, a native of South America and an escaped ornamental, is a serious environmental weed in Australia, South Africa and also several other countries. This paper reviews the efforts made over the last decade to bring about its biological control. The paper describes the weed status of cat's claw creeper in both Australia and South Africa. These two countries have cooperated in developing biocontrol projects and insect agents have now been released in both countries. These insects and other potential agents are described and suggestions made for the future direction of the projects.

Simulating the evolution of glyphosate resistance in grains farming in northern Australia.

Background and Aims: The evolution of resistance to herbicides is a substantial problem in contemporary agriculture. Solutions to this problem generally consist of the use of practices to control the resistant population once it evolves, and/or to institute preventative measures before populations become resistant. Herbicide resistance evolves in populations over years or decades, so predicting the effectiveness of preventative strategies in particular relies on computational modelling approaches. While models of herbicide resistance already exist, none deals with the complex regional variability in the northern Australian sub-tropical grains farming region. For this reason, a new computer model was developed. Methods: The model consists of an age- and stage-structured population model of weeds, with an existing crop model used to simulate plant growth and competition, and extensions to the crop model added to simulate seed bank ecology and population genetics factors. Using awnless barnyard grass (Echinochloa colona) as a test case, the model was used to investigate the likely rate of evolution under conditions expected to produce high selection pressure. Key Results: Simulating continuous summer fallows with glyphosate used as the only means of weed control resulted in predicted resistant weed populations after approx. 15 years. Validation of the model against the paddock history for the first real-world glyphosate-resistant awnless barnyard grass population shows that the model predicted resistance evolution to within a few years of the real situation. Conclusions: This validation work shows that empirical validation of herbicide resistance models is problematic. However, the model simulates the complexities of sub-tropical grains farming in Australia well, and can be used to investigate, generate and improve glyphosate resistance prevention strategies.

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.

Fruit traits of vertebrate-dispersed alien plants: smaller seeds and more pulp sugar than indigenous species

Vertebrates play a major role in dispersing seeds of fleshy-fruited alien plants. However, we know little of how the traits of alien fleshy fruits compare with indigenous fleshy fruits, and how these differences might contribute to invasion success. In this study, we characterised up to 38 fruit morphology, pulp nutrient and phenology traits of an assemblage of 34 vertebrate-dispersed alien species in south-eastern Queensland, Australia. Most alien fruits were small (81%\15 mm in mean width), and had watery fruit pulps that were high in sugars and low in nitrogen and lipids. When compared to indigenous species, alien fruits had significantly smaller seeds. Further, alien fruit pulps contained more sugar and more variable (and probably greater) nitrogen per pulp wet weight, and species tended to have longer fruiting seasons than indigenous species. Our analyses suggest that fruit traits could be important in determining invasiveness and could be used to improve pre- and post-border weed risk assessment.