Simulation to investigate site-based monitoring of pest insect species for trade.

Pest insect surveillance using lures is widely used to support market access requirements for traded articles that are hosts or carriers of quarantine pests. Modeling has been used extensively to guide the design of surveillance to support pest free area claims but is less commonly applied to provide confidence in pest freedom or low pest prevalence within sites registered for trade. Site-based surveillance typically needs to detect pests that are already present in the site or that may be entering the site from surrounding areas. We assessed the ability of site-based surveillance strategies to detect pests originating from within or outside the registered site using a probabilistic trapping network simulation model with random-walk insect movement and biologically realistic parameters. For a given release size, time-dependent detection probability was primarily determined by trap density and lure attractiveness, whereas mean step size (daily dispersal) had limited effect. Results were robust to site shape and size. For pests already within the site, detection was most sensitive using regularly spaced traps. Perimeter traps performed best for detecting pests moving into the site, although the importance of trap arrangement decreased with time from release, and random trap placement performed relatively well compared to regularly spaced traps. High detection probabilities were achievable within 7 days using realistic values for lure attractiveness and trap density. These findings, together with the modeling approach, can guide the development of internationally agreed principles for designing site-based surveillance of lure-attractant pests that is calibrated against the risk of non-detection.

Species identification, phylogenetic analysis and detection of herbicide-resistant biotypes of Amaranthus based on ALS and ITS.

The taxonomically challenging genus Amaranthus (Family Amaranthaceae) includes important agricultural weed species that are being spread globally as grain contaminants. We hypothesized that the ALS gene will help resolve these taxonomic challenges and identify potentially harmful resistant biotypes. We obtained 153 samples representing 26 species from three Amaranthus subgenera and included in that incorporated ITS, ALS (domains C, A and D) and ALS (domains B and E) sequences. Subgen. Albersia was well supported, but subgen. Amaranthus and subgen. Acnida were not. Amaranthus tuberculatus, A. palmeri and A. spinosus all showed different genetic structuring. Unique SNPs in ALS offered reliable diagnostics for most of the sampled Amaranthus species. Resistant ALS alleles were detected in sixteen A. tuberculatus samples (55.2%), eight A. palmeri (27.6%) and one A. arenicola (100%). These involved Ala122Asn, Pro197Ser/Thr/Ile, Trp574Leu, and Ser653Thr/Asn/Lys substitutions, with Ala122Asn, Pro197Thr/Ile and Ser653Lys being reported in Amaranthus for the first time. Moreover, different resistant mutations were present in different A. tuberculatus populations. In conclusion, the ALS gene is important for species identification, investigating population genetic diversity and understanding resistant evolution within the genus Amaranthus.

Sellers' Revisited: A Big Data Reassessment of Historical Outbreaks of Bluetongue and African Horse Sickness due to the Long-Distance Wind Dispersion of Culicoides Midges.

The possibility that outbreaks of bluetongue (BT) and African horse sickness (AHS) might occur via long-distance wind dispersion (LDWD) of their insect vector (Culicoides spp.) was proposed by R. F. Sellers in a series of papers published between 1977 and 1991. These investigated the role of LDWD by means of visual examination of the wind direction of synoptic weather charts. Based on the hypothesis that simple wind direction analysis, which does not allow for wind speed, might have led to spurious conclusions, we reanalyzed six of the outbreak scenarios described in Sellers' papers. For this reanalysis, we used a custom-built Big Data application ("TAPPAS") which couples a user-friendly web-interface with an established atmospheric dispersal model ("HYSPLIT"), thus enabling more sophisticated modeling than was possible when Sellers undertook his analyzes. For the two AHS outbreaks, there was strong support from our reanalysis of the role of LDWD for that in Spain (1966), and to a lesser degree, for the outbreak in Cyprus (1960). However, for the BT outbreaks, the reassessments were more complex, and for one of these (western Turkey, 1977) we could discount LDWD as the means of direct introduction of the virus. By contrast, while the outbreak in Cyprus (1977) showed LDWD was a possible means of introduction, there is an apparent inconsistency in that the outbreaks were localized while the dispersion events covered much of the island. For Portugal (1956), LDWD from Morocco on the dates suggested by Sellers is very unlikely to have been the pathway for introduction, and for the detection of serotype 2 in Florida (1982), LDWD from Cuba would require an assumption of a lengthy survival time of the midges in the air column. Except for western Turkey, the BT reanalyses show the limitation of LDWD modeling when used by itself, and indicates the need to integrate susceptible host population distribution (and other covariate) data into the modeling process. A further refinement, which will become increasingly important to assess LDWD, will be the use of virus and vector genome sequence data collected from potential source and the incursion sites.

Modelling seasonal habitat suitability for wide-ranging species: Invasive wild pigs in northern Australia.

Invasive wildlife often causes serious damage to the economy and agriculture as well as environmental, human and animal health. Habitat models can fill knowledge gaps about species distributions and assist planning to mitigate impacts. Yet, model accuracy and utility may be compromised by small study areas and limited integration of species ecology or temporal variability. Here we modelled seasonal habitat suitability for wild pigs, a widespread and harmful invader, in northern Australia. We developed a resource-based, spatially-explicit and regional-scale approach using Bayesian networks and spatial pattern suitability analysis. We integrated important ecological factors such as variability in environmental conditions, breeding requirements and home range movements. The habitat model was parameterized during a structured, iterative expert elicitation process and applied to a wet season and a dry season scenario. Model performance and uncertainty was evaluated against independent distributional data sets. Validation results showed that an expert-averaged model accurately predicted empirical wild pig presences in northern Australia for both seasonal scenarios. Model uncertainty was largely associated with different expert assumptions about wild pigs' resource-seeking home range movements. Habitat suitability varied considerably between seasons, retracting to resource-abundant rainforest, wetland and agricultural refuge areas during the dry season and expanding widely into surrounding grassland floodplains, savanna woodlands and coastal shrubs during the wet season. Overall, our model suggested that suitable wild pig habitat is less widely available in northern Australia than previously thought. Mapped results may be used to quantify impacts, assess risks, justify management investments and target control activities. Our methods are applicable to other wide-ranging species, especially in data-poor situations.

The Relative Importance of Genetic Diversity and Phenotypic Plasticity in Determining Invasion Success of a Clonal Weed in the USA and China.

Phenotypic plasticity has been proposed as an important adaptive strategy for clonal plants in heterogeneous habitats. Increased phenotypic plasticity can be especially beneficial for invasive clonal plants, allowing them to colonize new environments even when genetic diversity is low. However, the relative importance of genetic diversity and phenotypic plasticity for invasion success remains largely unknown. Here, we performed molecular marker analyses and a common garden experiment to investigate the genetic diversity and phenotypic plasticity of the globally important weed Alternanthera philoxeroides in response to different water availability (terrestrial vs. aquatic habitats). This species relies predominantly on clonal propagation in introduced ranges. We therefore expected genetic diversity to be restricted in the two sampled introduced ranges (the USA and China) when compared to the native range (Argentina), but that phenotypic plasticity may allow the species' full niche range to nonetheless be exploited. We found clones from China had very low genetic diversity in terms of both marker diversity and quantitative variation when compared with those from the USA and Argentina, probably reflecting different introduction histories. In contrast, similar patterns of phenotypic plasticity were found for clones from all three regions. Furthermore, despite the different levels of genetic diversity, bioclimatic modeling suggested that the full potential bioclimatic distribution had been invaded in both China and USA. Phenotypic plasticity, not genetic diversity, was therefore critical in allowing A. philoxeroides to invade diverse habitats across broad geographic areas.

Are high-impact species predictable? An analysis of naturalised grasses in northern Australia.

Predicting which species are likely to cause serious impacts in the future is crucial for targeting management efforts, but the characteristics of such species remain largely unconfirmed. We use data and expert opinion on tropical and subtropical grasses naturalised in Australia since European settlement to identify naturalised and high-impact species and subsequently to test whether high-impact species are predictable. High-impact species for the three main affected sectors (environment, pastoral and agriculture) were determined by assessing evidence against pre-defined criteria. Twenty-one of the 155 naturalised species (14%) were classified as high-impact, including four that affected more than one sector. High-impact species were more likely to have faster spread rates (regions invaded per decade) and to be semi-aquatic. Spread rate was best explained by whether species had been actively spread (as pasture), and time since naturalisation, but may not be explanatory as it was tightly correlated with range size and incidence rate. Giving more weight to minimising the chance of overlooking high-impact species, a priority for biosecurity, meant a wider range of predictors was required to identify high-impact species, and the predictive power of the models was reduced. By-sector analysis of predictors of high impact species was limited by their relative rarity, but showed sector differences, including to the universal predictors (spread rate and habitat) and life history. Furthermore, species causing high impact to agriculture have changed in the past 10 years with changes in farming practice, highlighting the importance of context in determining impact. A rationale for invasion ecology is to improve the prediction and response to future threats. Although our study identifies some universal predictors, it suggests improved prediction will require a far greater emphasis on impact rather than invasiveness, and will need to account for the individual circumstances of affected sectors and the relative rarity of high-impact species.

Development of microsatellite markers for the invasive weed Parthenium hysterophorus (Asteraceae).

PREMISE OF THE STUDY: Microsatellite primers were developed for Parthenium hysterophorus to investigate its genetic structure and genetic diversity. METHODS AND RESULTS: Using the combined biotin capture method, 15 microsatellite primer sets were isolated and characterized. All markers showed polymorphism, and the number of alleles per locus ranged from two to nine across 60 individuals from two populations. The observed and expected heterozygosities ranged from 0.117 to 0.750 and from 0.182 to 0.835, respectively. CONCLUSIONS: These markers will be useful for investigating the invasion history of this weed globally and to help characterize its invasiveness.

Differential influence of clonal integration on morphological and growth responses to light in two invasive herbs.

BACKGROUND AND AIMS: In contrast to seeds, high sensitivity of vegetative fragments to unfavourable environments may limit the expansion of clonal invasive plants. However, clonal integration promotes the establishment of propagules in less suitable habitats and may facilitate the expansion of clonal invaders into intact native communities. Here, we examine the influence of clonal integration on the morphology and growth of ramets in two invasive plants, Alternanthera philoxeroides and Phyla canescens, under varying light conditions. METHODS: In a greenhouse experiment, branches, connected ramets and severed ramets of the same mother plant were exposed under full sun and 85% shade and their morphological and growth responses were assessed. KEY RESULTS: The influence of clonal integration on the light reaction norm (connection×light interaction) of daughter ramets was species-specific. For A. philoxeroides, clonal integration evened out the light response (total biomass, leaf mass per area, and stem number, diameter and length) displayed in severed ramets, but these connection×light interactions were largely absent for P. canescens. Nevertheless, for both species, clonal integration overwhelmed light effect in promoting the growth of juvenile ramets during early development. Also, vertical growth, as an apparent shade acclimation response, was more prevalent in severed ramets than in connected ramets. Finally, unrooted branches displayed smaller organ size and slower growth than connected ramets, but the pattern of light reaction was similar, suggesting mother plants invest in daughter ramets prior to their own branches. CONCLUSIONS: Clonal integration modifies light reaction norms of morphological and growth traits in a species-specific manner for A. philoxeroides and P. canescens, but it improves the establishment of juvenile ramets of both species in light-limiting environments by promoting their growth during early development. This factor may be partially responsible for their ability to successfully colonize native plant communities.

Phenotypic plasticity influences the size, shape and dynamics of the geographic distribution of an invasive plant.

Phenotypic plasticity has long been suspected to allow invasive species to expand their geographic range across large-scale environmental gradients. We tested this possibility in Australia using a continental scale survey of the invasive tree Parkinsonia aculeata (Fabaceae) in twenty-three sites distributed across four climate regions and three habitat types. Using tree-level responses, we detected a trade-off between seed mass and seed number across the moisture gradient. Individual trees plastically and reversibly produced many small seeds at dry sites or years, and few big seeds at wet sites and years. Bigger seeds were positively correlated with higher seed and seedling survival rates. The trade-off, the relation between seed mass, seed and seedling survival, and other fitness components of the plant life-cycle were integrated within a matrix population model. The model confirms that the plastic response resulted in average fitness benefits across the life-cycle. Plasticity resulted in average fitness being positively maintained at the wet and dry range margins where extinction risks would otherwise have been high ("Jack-of-all-Trades" strategy JT), and fitness being maximized at the species range centre where extinction risks were already low ("Master-of-Some" strategy MS). The resulting hybrid "Jack-and-Master" strategy (JM) broadened the geographic range and amplified average fitness in the range centre. Our study provides the first empirical evidence for a JM species. It also confirms mechanistically the importance of phenotypic plasticity in determining the size, the shape and the dynamic of a species distribution. The JM allows rapid and reversible phenotypic responses to new or changing moisture conditions at different scales, providing the species with definite advantages over genetic adaptation when invading diverse and variable environments. Furthermore, natural selection pressure acting on phenotypic plasticity is predicted to result in maintenance of the JT and strengthening of the MS, further enhancing the species invasiveness in its range centre.

Tackling contentious invasive plant species: a case study of buffel grass in Australia.

Introduced plants that have both production values and negative impacts can be contentious. Generally they are either treated as weeds and their use prohibited; or unfettered exploitation is permitted and land managers must individually contend with any negative effects. Buffel grass (Cenchrus ciliaris) is contentious in Australia and there has been no attempt to broadly and systematically address the issues surrounding it. However, recent research indicates that there is some mutual acceptance by proponents and opponents of each others' perspectives and we contend that this provides the basis for a national approach. It would require thorough and on-going consultation with stakeholders and development of realistic goals that are applicable across a range of scales and responsive to regional differences in costs, benefits and socio-economic and biophysical circumstances. It would be necessary to clearly allocate responsibilities and ascertain the most appropriate balance between legislative and non-legislative mechanisms. A national approach could involve avoiding the introduction of additional genetic material, countering proliferation in regions where the species is sparse, preventing incursion into conservation reserves where it is absent, containing strategically located populations and managing communities to prevent or reduce dominance by buffel grass. This approach could be applied to other contentious plant species.

Phenotypic divergence during the invasion of Phyla canescens in Australia and France: evidence for selection-driven evolution.

Rapid adaptive evolution has been advocated as a mechanism that promotes invasion. Demonstrating adaptive evolution in invasive species requires rigorous analysis of phenotypic shifts driven by selection. Here, we document selection-driven evolution of Phyla canescens, an Argentine weed, in two invaded regions (Australia and France). Invasive populations possessed similar or higher diversity than native populations, and displayed mixed lineages from different sources, suggesting that genetic bottlenecks in both countries might have been alleviated by multiple introductions. Compared to native populations, Australian populations displayed more investment in sexual reproduction, whereas French populations possessed enhanced vegetative reproduction and growth. We partitioned evolutionary forces (selection vs. stochastic events) using two independent methods. Results of both analyses suggest that the pattern of molecular and phenotypic variability among regions was consistent with selection-driven evolution, rather than stochastic events. Our findings indicate that selection has shaped the evolution of P. canescens in two different invaded regions.

Interaction between seed dormancy-release mechanism, environment and seed bank strategy for a widely distributed perennial legume, Parkinsonia aculeata (Caesalpinaceae).

BACKGROUND AND AIMS: Parkinsonia aculeata (Caesalpinaceae) is a perennial legume with seeds that have hard-seeded (physical) dormancy and are potentially very long-lived. Seed dormancy is a characteristic that can both help maximize the probability of seedling establishment and spread the risk of recruitment failure across years (bet-hedging). In this study, dormancy-release patterns are described across the diverse environments in which this species occurs in order to test whether wet heat (incubation under wet, warm-to-hot, conditions) alone can explain those patterns, and in order to determine the likely ecological role of physical dormancy across this species distribution. METHODS: A seed burial trial was conducted across the full environmental distribution of P. aculeata in Australia (arid to wet-dry tropics, uplands to wetlands, soil surface to 10 cm deep). KEY RESULTS: Wet heat explained the pattern of dormancy release across all environments. Most seeds stored in the laboratory remained dormant throughout the trial (at least 84 %). Dormancy release was quickest for seeds buried during the wet season at relatively high rainfall, upland sites (only 3 % of seeds remained dormant after 35 d). The longest-lived seeds were in wetlands (9 % remained dormant after almost 4 years) and on the soil surface (57 % after 2 years). There was no consistent correlation between increased aridity and rate of dormancy release. CONCLUSIONS: The results suggest that physical dormancy in P. aculeata is a mechanism for maximizing seedling establishment rather than a bet-hedging strategy. However, seed persistence can occur in environmental refuges where dormancy-release cues are weak and conditions for germination and establishment are poor (e.g. under dense vegetation or in more arid micro-environments) or unsuitable (e.g. when seeds are inundated or on the soil surface). Risks of recruitment failure in suboptimal environments could therefore be reduced by inter-year fluctuations in microclimate or seed movement.

Wet-season dormancy release in seed banks of a tropical leguminous shrub is determined by wet heat.

BACKGROUND AND AIMS: Hard-seeded (physical) dormancy is common among plants, yet mechanisms for dormancy release are poorly understood, especially in the tropics. The following questions are asked: (a) whether dormancy release in seed banks of the tropical shrub Parkinsonia aculeata (Caesalpiniaceae) is determined by wet heat (incubation under wet, warm to hot, conditions); and (b) whether its effect is modified by microclimate. METHODS: A seed burial trial was conducted in the wet-dry tropics (northern Australia) to compare dormancy release across different habitats (open, artificial cover, ground cover and canopy cover), burial depths (0, 3 and 20 cm) and burial durations (2, 6 and 14 weeks). Results were compared with predictions using a laboratory-derived relationship between wet heat and dormancy release, and microclimate data collected during the trial. KEY RESULTS: Wet heat (defined as the soil temperature above which seeds were exposed to field capacity or higher for a cumulative total of 24 h) was 43.6 degrees C in the 0 cm open treatment, and decreased with increasing shade and depth to 29.5 degrees C at 20 cm under canopy cover. The dormancy release model showed that wet heat was a good predictor of the proportion of seeds remaining dormant. Furthermore, dormancy release was particularly sensitive to wet heat across the temperature range encountered across treatments. This resulted in a 16-fold difference in dormancy levels between open (<5 % of seeds still dormant) and covered (82 %) microhabitats. CONCLUSIONS: These results demonstrate that wet heat is the principal dormancy release mechanism for P. aculeata when conditions are hot and wet. They also highlight the potential importance of microclimate in driving the population dynamics of such species.