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.

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.

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?

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.

Leaf trait co-ordination in relation to construction cost, carbon gain and resource-use efficiency in exotic invasive and native woody vine species

Background and Aims: Success of invasive plant species is thought to be linked with their higher leaf carbon fixation strategy, enabling them to capture and utilize resources better than native species, and thus pre-empt and maintain space. However, these traits are not well-defined for invasive woody vines. Methods: In a glass house setting, experiments were conducted to examine how leaf carbon gain strategies differ between non-indigenous invasive and native woody vines of south-eastern Australia, by investigating their biomass gain, leaf structural, nutrient and physiological traits under changing light and moisture regimes. Key Results: Leaf construction cost (CC), calorific value and carbon : nitrogen (C : N) ratio were lower in the invasive group, while ash content, N, maximum photosynthesis, light-use efficiency, photosynthetic energyuse efficiency (PEUE) and specific leaf area (SLA) were higher in this group relative to the native group. Trait plasticity, relative growth rate (RGR), photosynthetic nitrogen-use efficiency and water-use efficiency did not differ significantly between the groups. However, across light resource, regression analyses indicated that at a common (same) leaf CC and PEUE, a higher biomass RGR resulted for the invasive group; also at a common SLA, a lower CC but higher N resulted for the invasive group. Overall, trait co-ordination (using pair-wise correlation analyses) was better in the invasive group. Ordination using 16 leaf traits indicated that the major axis of invasive-native dichotomy is primarily driven by SLA and CC (including its components and/or derivative of PEUE) and was significantly linked with RGR. Conclusions: These results demonstrated that while not all measures of leaf resource traits may differ between the two groups, the higher level of trait correlation and higher revenue returned (RGR) per unit of major resource need (CC) and use (PEUE) in the invasive group is in line with their rapid spread where introduced.

Seed persistence of the invasive aquatic plant, Gymnocoronis spilanthoides (Asteraceae)

Seed persistence of Gymnocoronis spilanthoides (D.Don) DC.; Asteraceae (Senegal tea), a serious weed of freshwater habitats, was examined in relation to burial status and different soil moisture regimes over a 3-year period. Seeds were found to be highly persistent, especially when buried. At the end of the experiment, 42.0%, 27.3% and 61.4% of buried seeds were viable following maintenance at field capacity, water logged and fluctuating (cycles of 1 week at field capacity followed by 3 weeks’ drying down) soil moisture conditions, respectively. Comparable viability values for surface-situated seeds were ~3% over all soil moisture regimes. Predicted times to1% viability are 16.2 years for buried seed and 3.8 years for surface-situated seed. Persistence was attributed primarily to the absence of light, a near-obligate requirement for germination in this species, although secondary dormancy was induced in some seeds. Previous work has demonstrated low fecundity in field populations of G. spilanthoides, which suggests that soil seed banks may not be particularly large. However, high levels of seed persistence, combined with ostensibly effective dispersal mechanisms, indicate that this weed may prove a difficult target for regional or state-wide eradication.

Variation in ecophysiology and carbon economy of invasive and native woody vines of riparian zones in south-eastern Queensland

Exotic and invasive woody vines are major environmental weeds of riparian areas, rainforest communities and remnant natural vegetation in coastal eastern Australia, where they smother standing vegetation, including large trees, and cause canopy collapse. We investigated, through glasshouse resource manipulative experiments, the ecophysiological traits that might facilitate faster growth, better resource acquisition and/or utilization and thus dominance of four exotic and invasive vines of South East Queensland, Australia, compared with their native counterparts. Relative growth rate was not significantly different between the two groups but water use efficiency (WUE) was higher in the native species while the converse was observed for light use efficiency (quantum efficiency, AQE) and maximum photosynthesis on a mass basis (Amax mass). The invasive species, as a group, also exhibited higher respiration load, higher light compensation point and higher specific leaf area. There were stronger correlations of leaf traits and greater structural (but not physiological) plasticity in invasive species than in their native counterparts. The scaling coefficients of resource use efficiencies (WUE, AQE and respiration efficiency) as well as those of fitness (biomass accumulated) versus many of the performance traits examined did not differ between the two species-origin groups, but there were indications of significant shifts in elevation (intercept values) and shifts along common slopes in many of these relationships – signalling differences in carbon economy (revenue returned per unit energy invested) and/or resource usage. Using ordination and based on 14 ecophysiological attributes, a fair level of separation between the two groups was achieved (51.5% explanatory power), with AQE, light compensation point, respiration load, WUE, specific leaf area and leaf area ratio, in decreasing order, being the main drivers. This study suggests similarity in trait plasticity, especially for physiological traits, but there appear to be fundamental differences in carbon economy and resource conservation between native and invasive vine species.

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.

Wet-dry cycling extends seed persistence by re-instating antioxidant capacity

Seeds in the field experience wet-dry cycling that is akin to the well-studied commercial process of seed priming in which seeds are hydrated and then re-dried to standardise their germination characteristics. To investigate whether the persistence (defined as in situ longevity) and antioxidant capacity of seeds are influenced by wet-dry cycling, seeds of the global agronomic weed Avena sterilis ssp. ludoviciana were subjected to (1) controlled ageing at 60% relative humidity and 53.5°C for 31 days, (2) controlled ageing then priming, or (3) ageing in the field in three soils for 21 months. Changes in seed viability (total germination), mean germination time, seedling vigour (mean seedling length), and the concentrations of the glutathione (GSH) / glutathione disulphide (GSSG) redox couple were recorded over time. As controlled-aged seeds lost viability, GSH levels declined and the relative proportion of GSSG contributing to total glutathione increased, indicative of a failing antioxidant capacity. Subjecting seeds that were aged under controlled conditions to a wet-dry cycle (to −1 MPa) prevented viability loss and increased GSH levels. Field-aged seeds that underwent numerous wet-dry cycles due to natural rainfall maintained high viability and high GSH levels. Thus wet-dry cycles in the field may enhance seed longevity and persistence coincident with re-synthesis of protective compounds such as GSH.

Lantana camara L. (Verbenaceae) invasion effects on soil physicochemical properties

Lantana camara is a recognized weed of worldwide significance due to its extensive distribution and its impacts on primary industries and nature conservation. However, quantitative data on the impact of the weed on soil ecosystem properties are scanty, especially in SE Australia, despite the pervasive presence of the weed along its coastal and inland regions. Consequently, mineral soils for physicochemical analyses were collected beneath and away from L. camara infestations in four sites west of Brisbane, SE Australia. These sites (hoop pine plantation, cattle farm, and two eucalyptus forests with occasional grazing and a fire regime, respectively) vary in landscape and land-use types. Significant site effect was more frequently observed than effect due to invasion status. Nonetheless, after controlling for site differences, ~50% of the 23 soil traits examined differed significantly between infested and non-infested soils. Moisture, pH, Ca, total and organic C, and total N (but not exchangeable N in form of NO3-) were significantly elevated, while sodium, chloride, copper, iron, sulfur, and manganese, many of which can be toxic to plant growth if present in excess levels, were present at lower levels in soils supporting L. camara compared to soils lacking the weed. These results indicate that L. camara can improve soil fertility and influence nutrient cycling, making the substratum ideal for its own growth and might explain the ability of the weed to outcompete other species, especially native ones.

Contagious dispersal of seeds of synchronously fruiting species beneath invasive and native fleshy-fruited trees

In subtropical Australia, many native and invasive plant species rely on a shared suite of frugivores, largely birds, for seed dispersal. Many native plants fruit during summer in this region, whereas most invasive plants fruit during winter, thus providing the opportunity for contagious dispersal of seeds beneath synchronously fruiting species. We sampled invasive and native seed rain beneath the canopy of a native summer-fruiting tree Guioa semiglauca and an invasive winter-fruiting tree Cinnamomum camphora, in three study sites over the course of a year. In July, during peak fruiting season for C. camphora and other invasive species, seed rain of invasive species was higher beneath C. camphora than G. semiglauca. This was partly due to the invasive tree Ligustrum lucidum, whose seed rain was three times higher beneath C. camphora than beneath the native tree. In February, seed rain of native species was more abundant beneath the canopy of G. semiglauca than beneath C. camphora, despite the fact that C. camphora was also fruiting at this time. This was probably due to the larger fruit crop produced by G. semiglauca at this time of year. Our study provides evidence that the presence of invasive bird-dispersed plants may facilitate contagious seed dispersal of other invaders, and likewise native species may facilitate seed spread of other native plants.

Innovative management of herbicide resistant weeds - seed banks and soil biology

The project will evaluate seed bank depletion of key northern herbicide resistant weeds under different environments, cropping systems, crop agronomies and non-chemical control tactics. The project will also evaluate soil biology and seed bank relationships to explain differences in seed bank persistence.

Invasion impacts on biodiversity: responses of ant communities to infestation by cat’s claw creeper, Macfadyena unguis-cati (Bignoniaceae) in subtropical Australia

Ants are the dominant soil faunal group in many if not most terrestrial ecosystems, and play a key role in soil structure and function. This study documents the impacts of invasion by the exotic cat’s claw creeper vine, Macfadyena unguis-cati (L.) Gentry (Bignoniaceae) on surface-situated (epigaeic) and subterranean (hypogaeic) ant communities in subtropical SE Queensland Australia where it is a major environmental weed of riparian areas, rainforest communities and remnant natural vegetation, smothering standing vegetation and causing canopy collapse. Soil ants were sampled in infested and uninfested areas at eight sites spanning both riparian and non-riparian habitats in subtropical SE Queensland. Patterns of ant species composition and functional grouping in response to patch invasion status, landscape type and habitat stratum were investigated using ANOVA and non-metric multidimensional scaling ordination. The epigaeic and subterranean strata supported markedly different ant assemblages, and ant communities also differed between riparian and non-riparian habitats. However, M. unguis-cati invasion had a surprisingly limited impact. There was a tendency for ant abundance and species richness to be lower in infested patches, and overall species composition was different between infested and uninfested patches, but these differences were relatively small, and did not occur consistently across sites. There were changes in functional group composition that conformed to known functional group responses to environmental change, but these were similarly limited and inconsistent across sites. Our study has shown that ant communities are surprisingly resilient to invasion by M. unguis-cati, and serves as a warning against making assumptions about invasion impacts based on visual appearances.

Modeling population growth and site specific control of the invasive Lantana camara L. (Verbenaceae) under differing fire regimes

It is at the population level that an invasion either fails or succeeds. Lantana camara L. (Verbenaceae) is a weed of great significance in Queensland Australia and globally but its whole life-history ecology is poorly known. Here we used 3 years of field data across four land use types (farm, hoop pine plantation and two open eucalyptus forests, including one with a triennial fire regime) to parameterise the weed’s vital rates and develop size-structured matrix models. Lantana camara in its re-colonization phase, as observed in the recently cleared hoop pine plantation, was projected to increase more rapidly (annual growth rate, λ = 3.80) than at the other three sites (λ 1.88–2.71). Elasticity analyses indicated that growth contributed more (64.6 %) to λ than fecundity (18.5 %) or survival (15.5 %), while across size groups, the contribution was of the order: juvenile (19–27 %) ≥ seed (17–28 %) ≥ seedling (16–25 %) > small adult (4–26 %) ≥ medium adult (7–20 %) > large adult (0–20 %). From a control perspective it is difficult to determine a single weak point in the life cycle of lantana that might be exploited to reduce growth below a sustaining rate. The triennial fire regime applied did not alter the population elasticity structure nor resulted in local control of the weed. However, simulations showed that, except for the farm population, periodic burning could work within 4–10 years for control of the weed, but fire frequency should increase to at least once every 2 years. For the farm, site-specific control may be achieved by 15 years if the biennial fire frequency is tempered with increased burning intensity.

Cat's claw creeper vine, Macfadyena unguis-cati (Bignoniaceae), invasion impacts: comparative leaf nutrient content and effects on soil physicochemical properties

Macfadyena unguis-cati (L.) Gentry (Bignoniaceae) is a major environmental weed in coastal Queensland, Australia. There is a lack of quantitative data on its leaf chemistry and its impact on soil properties. Soils from infested vs uninfested areas, and leaves of M. unguis-cati and three co-occurring vine species (one exotic, two native) were collected at six sites (riparian and non-riparian) in south-eastern Queensland. Effects of invasion status, species, site and habitat type were examined using univariate and multivariate analyses. Habitat type had a greater effect on soil nutrients than on leaf chemistry. Invasion effect of M. unguis-cati on soil chemistry was more pronounced in non-riparian than in riparian habitat. Significantly higher values were obtained in M. unguis-cati infested (vs. uninfested) soils for ~50% of traits. Leaf ion concentrations differed significantly between exotic and native vines. Observed higher leaf-nutrient load (especially nitrogen, phosphorus and potassium) in exotic plants aligns with the preference of invasive plant species for disturbed habitats with higher nutrient input. Higher load of trace elements (aluminium, boron, cadmium and iron) in its leaves suggests that cycling of heavy-metal ions, many of which are potentially toxic at excess level, could be accelerated in soils of M. unguis-cati-invaded landscape. Although inferences from the present study are based on correlative data, the consistency of the patterns across many sites suggests that M. unguis-cati may improve soil fertility and influence nutrient cycling, perhaps through legacy effects of its own litter input.