Niche based distribution modelling of an invasive alien plant: effects of population status, propagule pressure and invasion history

Forecasting the spatial spread of invasive species is important to inform management planning. Niche-based species distribution models offer a well-developed framework for assessing the potential range of species. However, these models assume equilibrium between the species’ distribution and its ecological requirements. During range expansion, invasive species are not in such equilibrium due to both dispersal limitation and frequent casual occurrence in sites unsuitable to persistent populations. In this article we use the example of the invasive annual plant Ambrosia artemisiifolia in Austria to evaluate if model accuracy can be enhanced in such non-equilibrium situations by taking account of propagule pressure and by restricting model calibration to naturalized populations. Moreover, we test if model accuracy increases during invasion history using distribution data from 1984 to 2005. The results suggest that models calibrated with naturalized populations are much more accurate than those based on the total set of records. Proxies of propagule pressure slightly but significantly improve goodness of fit, accuracy, and Type I and II error rates of models calibrated with all available records but have less consistent effects on models of naturalized populations. Model accuracy did not increase during the recent invasion history, probably because the species is still far from an equilibrium distribution. We conclude that even a coarse assessment of population status with records of invasive species delivers important information for predictive modelling and that proxies of propagule pressure should be included into such models at least during early to intermediate stages of the invasion history.     

Divergent effects of land-use,propagule pressure,and climate on woody riparian invasion

Landscape-scale analyses of biological invasion are needed to understand the relative importance of environmental drivers that vary at larger scales, such as climate, propagule pressure, resource availability, and human disturbance. One poorly understood landscape-scale question is, how does human land-use influence riparian plant invasion? To evaluate the relative importance of land-use, climate, propagule pressure, and water availability in riparian invasion, we examined tamarisk (Tamarix ramosissima, T. chinensis, hybrids), Russian olive (Elaeagnus angustifolia), and Siberian elm (Ulmus pumila) occurrence, abundance, and dominance in 238 riparian sites in developed, cultivated, and undeveloped areas of four western USA river basins (281,946 km²). Temperature and propagule pressure from individuals planted nearby largely drove invasive species occurrence, whereas factors likely to affect resource availability (e.g., land-use, precipitation, streamflow intermittency) were more important to abundance and dominance, supporting the argument that species distribution models based on occurrence alone may fail to identify conditions where invasive species have the greatest impact. The role of land-use varied among taxa: urban and suburban land-use increased Siberian elm occurrence, abundance, and dominance, and urban land-use increased Russian olive occurrence, whereas suburban land-use reduced tamarisk dominance. Surprisingly, Siberian elm, which has received scant prior scientific and management attention, occurred as or more frequently than tamarisk and Russian olive (except in undeveloped areas of the Colorado River headwaters) and had higher density and dominance than tamarisk and Russian olive in developed areas. More research is needed to understand the impacts of this largely unrecognized invader on riparian ecosystem services, particularly in urban and suburban areas.     

Decomposing propagule pressure: the effects of propagule size and propagule frequency on invasion success

Propagule pressure quantifies the inflow of individuals to a location and appears to be a key driver of invasion success. It is often defined as the average number of individuals introduced per time unit, or equivalently as the product of the average number of individuals introduced per introduction event (propagule size) and the frequency of introduction events (propagule frequency). Here we study how the influence of propagule size, frequency, and their product depends on the underlying ecological conditions. While previous studies have focused on introductions under environmental heterogeneity or a strong Allee effect, we examine a range of ecological scenarios that differ in the type of density dependence and in the sign of per capita growth rate. Our results indicate that the relative influence of propagule size and frequency depends mainly on the sign of per capita growth rate. Given a certain average number of individuals introduced per time unit, a high propagule frequency accelerates invasions under ecological scenarios with positive average per capita growth rate throughout the invasion process (‘easy’ scenarios). If per capita growth rate is negative throughout the invasion process (‘difficult’ scenarios) or if there is both an easy and a difficult stage (‘mixed scenarios’), a high propagule size leads to a faster invasion than a high propagule frequency. To explain this finding, we argue that for a fixed value of the product of propagule size and frequency, an increase in propagule size leads to an increase in demographic variance, which promotes invasion success in difficult and mixed but not in easy scenarios. However, we also show that in many of these cases, the product of propagule size and frequency still correlates more strongly with invasion success than either of the single components. Finally, we illustrate our approach with empirical examples from the literature.     

Integrating species distribution models and interacting particle systems to predict the spread of an invasive alien plant

Aim We demonstrate how to integrate two widely used tools for modelling the spread of invasive plants, and compare the performance of the combined model with that of its individual components using the recent range dynamics of the invasive annual weed Ambrosia artemisiifolia L. Location Austria. Methods Species distribution models, which deliver habitat‐based information on potential distributions, and interacting particle systems, which simulate spatio‐temporal range dynamics as dependent on neighbourhood configurations, were combined into a common framework. We then used the combined model to simulate the invasion of A. artemisiifolia in Austria between 1990 and 2005. For comparison, simulations were also performed with models that accounted only for habitat suitability or neighbourhood configurations. The fit of the three models to the data was assessed by likelihood ratio tests, and simulated invasion patterns were evaluated against observed ones in terms of predictive discrimination ability (area under the receiver operating characteristic curve, AUC) and spatial autocorrelation (Moran’s I). Results The combined model fitted the data significantly better than the single‐component alternatives. Simulations relying solely on parameterized spread kernels performed worst in terms of both AUC and spatial pattern formation. Simulations based only on habitat information correctly predicted infestation of susceptible areas but reproduced the autocorrelated patterns of A. artemisiifolia expansion less adequately than did the integrated model. Main conclusions Our integrated modelling approach offers a flexible tool for forecasts of spatio‐temporal invasion patterns from landscape to regional scales. As a further advantage, scenarios of environmental change can be incorporated consistently by appropriately updating habitat suitability layers. Given the susceptibility of many alien plants, including A. artemisiifolia, to both land use and climate changes, taking such scenarios into account will increasingly become relevant for the design of proactive management strategies.     

Using maximum entropy to predict the potential distribution of an invasive freshwater snail

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The relative importance of latitude matching and propagule pressure in the colonization success of an invasive forb

Factors that influence the early stages of invasion can be critical to invasion success, yet are seldom studied. In particular, broad pre-adaptation to recipient climate may importantly influence early colonization success, yet few studies have explicitly examined this. I performed an experiment to determine how similarity between seed source and transplant site latitude, as a general indicator of pre-adaptation to climate, interacts with propagule pressure (100, 200 and 400 seeds/pot) to influence early colonization success of the widespread North American weed, St. John's wort Hypericum perforatum . Seeds originating from seven native European source populations were sown in pots buried in the ground in a field in western Montana. Seed source populations were either similar or divergent in latitude to the recipient transplant site. Across seed density treatments, the match between seed source and recipient latitude did not affect the proportion of pots colonized or the number of individual colonists per pot. In contrast, propagule pressure had a significant and positive effect on colonization. These results suggest that propagules from many climatically divergent source populations can be viable invaders.     

Hotspots of plant invasion predicted by propagule pressure and ecosystem characteristics

Aim Biological invasions pose a major conservation threat and are occurring at an unprecedented rate. Disproportionate levels of invasion across the landscape indicate that propagule pressure and ecosystem characteristics can mediate invasion success. However, most invasion predictions relate to species’ characteristics (invasiveness) and habitat requirements. Given myriad invaders and the inability to generalize from single‐species studies, more general predictions about invasion are required. We present a simple new method for characterizing and predicting landscape susceptibility to invasion that is not species‐specific. Location Corangamite catchment (13,340 km²), south‐east Australia. Methods Using spatially referenced data on the locations of non‐native plant species, we modelled their expected proportional cover as a function of a site’s environmental conditions and geographic location. Models were built as boosted regression trees (BRTs). Results On average, the BRTs explained 38% of variation in occupancy and abundance of all exotic species and exotic forbs. Variables indicating propagule pressure, human impacts, abiotic and community characteristics were rated as the top four most influential variables in each model. Presumably reflecting higher propagule pressure and resource availability, invasion was highest near edges of vegetation fragments and areas of human activity. Sites with high vegetation cover had higher probability of occupancy but lower proportional cover of invaders, the latter trend suggesting a form of biotic resistance. Invasion patterns varied little in time despite the data spanning 34 years. Main conclusions To our knowledge, this is the first multispecies model based on occupancy and abundance data used to predict invasion risk at the landscape scale. Our approach is flexible and can be applied in different biomes, at multiple scales and for different taxonomic groups. Quantifying general patterns and processes of plant invasion will increase understanding of invasion and community ecology. Predicting invasion risk enables spatial prioritization of weed surveillance and control.     

Contrasting effects of an invasive ant on a native and an invasive plant

When invasive species establish in new environments, they may disrupt existing or create new interactions with resident species. Understanding of the functioning of invaded ecosystems will benefit from careful investigation of resulting species-level interactions. We manipulated ant visitation to compare how invasive ant mutualisms affect two common plants, one native and one invasive, on a sub-tropical Indian Ocean island. Technomyrmex albipes, an introduced species, was the most common and abundant ant visitor to the plants. T. albipes were attracted to extrafloral nectaries on the invasive tree (Leucaena leucocephala) and deterred the plant’s primary herbivore, the Leucaena psyllid (Heteropsylla cubana). Ant exclusion from L. leucocephala resulted in decreased plant growth and seed production by 22% and 35%, respectively. In contrast, on the native shrub (Scaevola taccada), T. albipes frequently tended sap-sucking hemipterans, and ant exclusion resulted in 30% and 23% increases in growth and fruit production, respectively. Stable isotope analysis confirmed the more predacious and herbivorous diets of T. albipes on the invasive and native plants, respectively. Thus the ants’ interactions protect the invasive plant from its main herbivore while also exacerbating the effects of herbivores on the native plant. Ultimately, the negative effects on the native plant and positive effects on the invasive plant may work in concert to facilitate invasion by the invasive plant. Our findings underscore the importance of investigating facilitative interactions in a community context and the multiple and diverse interactions shaping novel ecosystems.     

The risk of establishment of aquatic invasive species: joining invasibility and propagule pressure

Invasive species are increasingly becoming a policy priority. This has spurred researchers and managers to try to estimate the risk of invasion. Conceptually, invasions are dependent both on the receiving environment (invasibility) and on the ability to reach these new areas (propagule pressure). However, analyses of risk typically examine only one or the other. Here, we develop and apply a joint model of invasion risk that simultaneously incorporates invasibility and propagule pressure. We present arguments that the behaviour of these two elements of risk differs substantially--propagule pressure is a function of time, whereas invasibility is not--and therefore have different management implications. Further, we use the well-studied zebra mussel (Dreissena polymorpha) to contrast predictions made using the joint model to those made by separate invasibility and propagule pressure models. We show that predictions of invasion progress as well as of the long-term invasion pattern are strongly affected by using a joint model.     

The vulnerability of habitats to plant invasion: disentangling the roles of propagule pressure,time and sampling effort

Aim To quantify the vulnerability of habitats to invasion by alien plants having accounted for the effects of propagule pressure, time and sampling effort. Location New Zealand. Methods We used spatial, temporal and habitat information taken from 9297 herbarium records of 301 alien plant species to examine the vulnerability of 11 terrestrial habitats to plant invasions. A null model that randomized species records across habitats was used to account for variation in sampling effort and to derive a relative measure of invasion based either on all records for a species or only its first record. The relative level of invasion was related to the average distance of each habitat from the nearest conurbation, which was used as a proxy for propagule pressure. The habitat in which a species was first recorded was compared to the habitats encountered for all records of that species to determine whether the initial habitat could predict subsequent habitat occupancy. Results Variation in sampling effort in space and time significantly masked the underlying vulnerability of habitats to plant invasions. Distance from the nearest conurbation had little effect on the relative level of invasion in each habitat, but the number of first records of each species significantly declined with increasing distance. While Urban, Streamside and Coastal habitats were over‐represented as sites of initial invasion, there was no evidence of major invasion hotspots from which alien plants might subsequently spread. Rather, the data suggest that certain habitats (especially Roadsides) readily accumulate alien plants from other habitats. Main conclusions Herbarium records combined with a suitable null model provide a powerful tool for assessing the relative vulnerability of habitats to plant invasion. The first records of alien plants tend to be found near conurbations, but this pattern disappears with subsequent spread. Regardless of the habitat where a species was first recorded, ultimately most alien plants spread to Roadside and Sparse habitats. This information suggests that such habitats may be useful targets for weed surveillance and monitoring.     

Recruitment in species-rich grasslands: the effects of functional traits and propagule pressure

Aims The assembly of plant communities is a complex process which combines impacts from the species pool, dispersal and propagule pressure, niche requirements of colonizing species and the niche structure of the community. Recent theory development has incorporated all these aspects, e.g. in 'stochastic niche theory'. We investigated recruitment into a species-rich grassland community, using an experimental approach where we manipulated the trait composition of the community and examined the success of colonizing species entering with various propagule pressure. Specifically, we examined two predictions: (i) colonization success increases with increasing difference between traits of the colonizing species and the trait profile of the community and (ii) colonization success increases with increasing propagule pressure.Methods The examined communities were species-rich semi-natural grasslands located in southern Sweden. After a careful documentation of the composition of the plant communities at the experimental sites, we manipulated the trait profile of species-rich grassland plots based on the plant functional trait specific leaf area (SLA), which is correlated with several key life history functions. In addition to SLA, seed mass was also used to describe the trait profile of grassland plots. Seeds of 12 plant species from the regional species pool, varying in SLA and seed mass, were sown into plots using four different levels of propagule pressure. Recruitment was examined after 1 year. We also planted juvenile 'plug plants' of the same species which allowed us to examine survivorship and growth beyond the seedling stage.Important findings Overall we found very limited evidence for relationships between the traits of the colonizing species and the trait profile of the community and for recruitment after sowing these relationships were contrary to the prediction. Survival of plug plants after two seasons of growth was high irrespective of the trait profile of the community, but growth of plug plants was affected by the trait profile of the surrounding community. For four of the species there was a positive effect of increased propagule pressure on colonization. The results suggest that species assembly in species-rich grasslands is not strongly dependent on the niche structure of the community. However, the finding that colonization of only a third of the species responded positively to increased propagule pressure indicates that there might be niche-related effects that were not captured by our treatments. Overall, our results indicate that the factors determining colonization in this community are species specific. Some species are able to colonize irrespective of niche relationships, provided that the propagule pressure is sufficiently high to overcome stochastic mortality after seed arrival. For other species, however, we cannot exclude that niche assembly occurred, but we failed to identify the relevant niche factor.     

Riparian land-use and in-stream habitat predict the distribution of a critically endangered freshwater mussel

Hydrobiologia - Recent human population growth in North America’s southern Appalachian Mountains has raised concerns about the effects of ex-urban development on its streams and their at-risk...     

Manipulating propagule pressure to test the invasibility of subtidal marine habitats

Global patterns show that estuaries are more invaded than open coasts and artificial habitats are more invaded than natural ones. The contention that artificial habitats in estuaries are more invasible than other habitats may result from variation in propagule supply, however, as artificial habitats are closely linked to vectors of non-native propagules, such as ships and boats. True comparisons of habitat invasibility require manipulations of propagule pressure, which has been historically difficult in marine systems. Using in-situ larval dosing, we delivered propagules of the widespread invasive ascidian Botrylloides violaceus into field mesocosms and assessed how habitat type (floating dock vs. benthic rock), resource availability (occupied vs. unoccupied plates), and propagule number (5, 25 and 50 larvae 225 cm^?2) affected settlement (survival after 24 h) and recruitment (survival after 56 days) success. In-situ larval dosing was successful, and after eight weeks there were significant differences in recruitment due to initial dose-size, habitat type, and space availability. At the habitat scale, despite equal propagule delivery, PVC plates in natural benthic rock were not equally invasible and few propagules survived to recruitment. At the organism scale, increased habitat complexity, through facilitation by established fouling species, rather than freedom from space competitors, appears to be more important for B. violaceus to establish. Our results offer greater mechanistic understanding of broader invasion patterns between artificial and natural habitats. This work extends the possibilities for further research to counteract the confounding issue of unknown and unknowable propagule delivery when attempting to explain variation in invasion success.     

提高生态位模型转移能力来模拟入侵物种 的潜在分布

生态位模型利用物种分布点所关联的环境变量去推算物种的生态需求, 模拟物种的分布。在模拟入侵物种分布时, 经典生态位模型包括模型构建于物种本土分布地, 然后将其转移并投射至另一地理区域, 来模拟入侵物种的潜在分布。然而在模型运用时, 出现了模型的转移能力较低、模拟的结果与物种的实际分布不相符的情况, 由此得出了生态位漂移等不恰当的结论。提高生态位模型的转移能力, 可以准确地模拟入侵物种的潜在分布, 为入侵种的风险评估提供参考。作者以入侵种茶翅蝽(Halyomorpha halys)和互花米草(Spartina alterniflora)为例, 从模型的构建材料(即物种分布点和环境变量)入手, 全面阐述提高模型转移能力的策略。在构建模型之前, 需要充分了解入侵物种的生物学特性、种群平衡状态、本土地理分布范围及物种的生物历史地理等方面的知识。在模型构建环节上, 物种分布点不仅要充分覆盖物种的地理分布和生态空间的范围, 同时要降低物种采样点偏差; 环境变量的选择要充分考虑其对物种分布的限制作用、各环境变量之间的空间相关性, 以及不同地理种群间生态空间是否一致, 同时要降低环境变量的空间维度; 模型构建区域要真实地反映物种的地理分布范围, 并考虑种群的平衡状态。作者认为, 在生态位保守的前提下, 如果模型是构建在一个合理方案的基础上, 生态位模型的转移能力是可以保证的, 在以模型转移能力较低的现象来阐述生态位分化时需要引起注意。