Predicting the invasion success of Mediterranean alien plants from their introduction characteristics

We describe a model to predict the invasion success of alien plants on Mediterranean islands, according to their mode, frequency and time since introduction and simple species traits. The model was developed using canonical discriminant analysis to identify categories of low- and high-risk species. Whilst classifying ca 80% of the data set correctly, the type II error rate remained moderately high, which limits its value in practical terms. This level of accuracy is typical of similar models, and critical analysis of the weaknesses is necessary to develop long-term improvements in methodology. Sensitivity analysis indicates that, overall, mode of introduction had an important influence on invasiveness, with species introduced for public amenity being more likely to attain high-risk status than species introduced privately to gardens. Frequency and time since introduction had a relatively minor influence on the model outcome, but remained highly confounded with other variables. Although invasion success, as measured by the number of islands colonized, is expected to increase over time, it was not possible to estimate the rates of expansion from historical trends, which are complicated by changes in import fashions over the centuries. Many older introductions were crops, whereas there has been a recent tendency towards exotic ornamentals and weedy species. Despite a fundamental influence on the results, similar historical influences are seldom taken into account adequately during trait analyses. Unravelling such inter-correlations between predictors is therefore an important challenge for the future of screening protocols.     

How do introduction characteristics influence the invasion success of Mediterranean alien plants?

Invasive plant species are becoming increasingly widespread following accelerated anthropogenic activity in the Mediterranean region. Humans have played a central role in the expansion process, and it is important to incorporate such considerations into management plans. Using generalized linear models, our first aim was to describe how the invasion success of 862 prominent alien plant species on Mediterranean islands is related to characteristics of the introduction process: introduction frequency, date and region of origin, range size and purpose of import. The importance of each was measured by the numbers of species present and their average invasiveness. The main findings were: (a) accidental imports and ornamentals accounted for a high proportion of all aliens, although neither group had particularly high average invasiveness; (b) introduction frequency had a comparatively modest influence, with the most commonly-introduced species naturalized only three times more widely than those rarely-introduced; (c) rates of species introduction appear to have increased dramatically in the last century, although aliens which have been present in the region for more than 200 years were most widespread, indicating that it may be centuries before some species fill their potential range; (d) there were small tendencies for successful invaders to originate in the Neotropics or in regions with Mediterranean climate biomes and to have large range sizes. Our second aim was to determine whether the number or average invasiveness of species introduced via a given pathway had the most influence on the overall probability of invasion on a given island. An elasticity analysis suggested that the number of species was substantially the best predictor of the two. This finding arises largely because invasion events are rare and remain unpredictable, and has significant implications for assessing invasion risk. We discuss how substantial sources of error and intrinsic variability in invasiveness within species groups limit the potential for developing accurate risk models.     

外来植物入侵的全境性研究进展与展望

由于外来种入侵地一般都远离其自然分布区(原产地), 如果只在其入侵地或者原产地进行研究, 很难真正发现其入侵性形成和成功入侵的根本原因。目前, 许多学者开始关注和倡导对入侵种在原产地和入侵地的表现同时进行研究, 即入侵种的全境性研究(whole-range studies), 为入侵生物现有地理分布格局的形成原因和入侵机制等提供解释。本文结合国内外关于入侵植物全境性研究的进展和成果, 分别针对研究的主要目的、内容、意义等进行了全面的阐述, 探讨了存在的问题与不足, 并对未来相关研究进行了展望。目前已有的全境性研究主要是通过野外直接观测和同质种植园实验来比较入侵种在入侵地和原产地的生长、繁殖和生理生态等表型性状的差异, 以及应用分子标记方法比较入侵地种群和原产地种群遗传多样性的差异, 进行入侵植物的分子系统地理学研究, 从而有效检验生物入侵机制的理论和假说, 深入阐明植物入侵的机制, 为制定入侵植物的防控策略提供指导。值得注意的是, 由于外来植物入侵的全境性研究起步较晚, 现有研究的方法和内容还不够完善, 今后需要在加强国际合作的基础上进一步改善。     

Herbivory in a Mediterranean forest: browsing impact and plant compensation

The compensatory response of plants to defoliation is likely to have important effects on plant–ungulate equilibria in forested ecosystems. We investigated the responses of six species of Mediterranean bushes to defoliation by wild ungulates, comparing an index of browsing impact with the productivity of plants in both open and exclusion plots. The data revealed a great diversity of plant responses to herbivory: Rubus ulmifolius was able to over-compensate and replace the lost tissues. Phillyrea latifolia exhibited a similar, albeit less evident, pattern, while Cistus salvifolius was severely damaged by browsing. Other species, such as Quercus ilex, Juncus acutus and Erica arborea, were not attacked to a large extent and suffered little or no harm. The results strongly suggest that Mediterranean ecosystems may tolerate large stocking rates of ungulates. However, the reduction of plant biomass due to browsing was very different in the six studied species, suggesting that when herbivory becomes severe the structure of the ecosystem will change with the more tolerant plants becoming more abundant. We can apply these results to improve management and conservation of relict coastal forests in the Mediterranean basin which are usually of small size and where decision-makers have to compromise between the conservation of plants and that of large mammals.     

Herbivory in a Mediterranean forest: browsing impact and plant compensation

The compensatory response of plants to defoliation is likely to have important effects on plant–ungulate equilibria in forested ecosystems. We investigated the responses of six species of Mediterranean bushes to defoliation by wild ungulates, comparing an index of browsing impact with the productivity of plants in both open and exclusion plots. The data revealed a great diversity of plant responses to herbivory: Rubus ulmifolius was able to over-compensate and replace the lost tissues. Phillyrea latifolia exhibited a similar, albeit less evident, pattern, while Cistus salvifolius was severely damaged by browsing. Other species, such as Quercus ilex, Juncus acutus and Erica arborea, were not attacked to a large extent and suffered little or no harm. The results strongly suggest that Mediterranean ecosystems may tolerate large stocking rates of ungulates. However, the reduction of plant biomass due to browsing was very different in the six studied species, suggesting that when herbivory becomes severe the structure of the ecosystem will change with the more tolerant plants becoming more abundant. We can apply these results to improve management and conservation of relict coastal forests in the Mediterranean basin which are usually of small size and where decision-makers have to compromise between the conservation of plants and that of large mammals.     

Feral horses dung piles as potential invasion windows for alien plant species in natural grasslands

Small scale disturbances could act as patches that provide sites for the colonization of competitively inferior species, promoting the establishment of non-native species in some cases. We analyzed the vegetation associated with feral horse dung piles in montane pampas grasslands in Mid-East Argentina and described the changes following their abandonment, evaluating whether dung piles act as invasion windows, allowing the entrance of alien plant species. We estimated the portion of the study area directly covered by horse manure and dung height was used to estimate the time elapsed after the abandonment of each pile. Vegetation replacement on dung piles of different ages was assessed and compared with grassland controls using discriminant analysis. We used regression analysis to look for changes in vegetation cover, species richness, species diversity and evenness in response to height (age) of the dung piles, and principal component analyses (PCA) to identify groups of plants associated with different successional stages. We compared cover of alien plant species on dung piles with grassland controls using one-way ANOVA. On average, 2.5% of the study area was covered by horse dung. Total vegetation cover, species richness, diversity and evenness increased after the piles were abandoned. Characteristic plant groups were associated with initial, middle and last phases of the studied succession. Vegetation on the dung piles significantly differed from that in grassland controls and two species were consistently associated with dung piles: the invasive Red Star Thistle, Centaurea calcitrapa, and a native grazing-intolerant grass, Nassella clarazii. Non-native species cover was also higher in dung piles than in control plots. Dung piles cover a significant portion of grassland area in our study site, produce significant changes in the vegetation and are associated with some invasive alien plants that could eventually colonize more pristine areas in the vicinity. On the other hand, they might represent refuges for palatable species, since horses seem to avoid them for grazing.     

Observed and dark diversity of alien plant species in Europe: estimating future invasion risk

Invasion by alien species is a growing concern for nature conservation. We estimated the level of invasion by alien plant species and future invasion risks at the European scale. We used a pan-European atlas and eight regional plant atlases to determine the distribution of alien and native plant richness. In addition, we estimated alien and native dark diversity (species currently absent from a site but present in the surrounding region and able to colonize the site). We used relative diversity metrics to indicate current and future risks by alien species: relative alien richness (compared to native species), alien and native completeness (log-ratio of observed to dark diversity) and completeness difference between alien and native species. Observed and relative richness of alien species were greatest in NW Europe; this suggests that sites in NW Europe could be more disturbed. Observed alien and native species richness show clear regional hotspots; the distribution of completeness values is dispersed, indicating local hotspots. Northern Europe has relatively lower alien completeness, likely because potential invaders inhabit the region but have not yet reached many localities, thereby suggesting a risk of future invasion. A greater number of potential alien species in the region increases the probability that some alien species could have detrimental impacts. Both alien richness and completeness are positively correlated with native richness and completeness, respectively, indicating that both groups share similar distribution patterns. Alien species diversity metrics in Europe are related positively to human population density and agricultural land-use. We suggest that the dark diversity concept can broaden our understanding of alien species diversity and future invasion risks.     

Fire-driven alien invasion in a fire-adapted ecosystem

Disturbance plays a key role in many alien plant invasions. However, often the main driver of invasion is not disturbance per se but alterations in the disturbance regime. In some fire-adapted shrublands, the community is highly resilient to infrequent, high-intensity fires, but changes in the fire regime that result in shorter fire intervals may make these communities more susceptible to alien plant invasions. This study examines several wildfire events that resulted in short fire intervals in California chaparral shrublands. In one study, we compared postfire recovery patterns in sites with different prefire stand ages (3 and 24 years), and in another study we compared sites that had burned once in four years with sites that had burned twice in this period. The population size of the dominant native shrub Adenostoma fasciculatum was drastically reduced following fire in the 3-year sites relative to the 24-year sites. The 3-year sites had much greater alien plant cover and significantly lower plant diversity than the 24-year sites. In a separate study, repeat fires four years apart on the same sites showed that annual species increased significantly after the second fire, and alien annuals far outnumbered native annuals. Aliens included both annual grasses and annual forbs and were negatively correlated with woody plant cover. Native woody species regenerated well after the first fire but declined after the second fire, and one obligate seeding shrub was extirpated from two sites by the repeat fires. It is concluded that some fire-adapted shrublands are vulnerable to changes in fire regime, and this can lead to a loss of native diversity and put the community on a trajectory towards type conversion from a woody to an herbaceous system. Such changes result in alterations in the proportion of natives to non-natives, changes in functional types from deeply rooted shrubs to shallow rooted grasses and forbs, increased fire frequency due to the increase in fine fuels, and changes in carbon storage.     

资源脉冲对外来植物入侵影响的研究进展和展望

全球变化背景下, 诸如营养、水分等资源的波动是非稳态的, 往往以脉冲的形式出现, 呈现出频率低、强度高和持续时间短的特征。资源脉冲往往会打破植物群落固有的平衡状态, 进而影响全球变化的另一重要组分——外来植物入侵。目前, 全球变化对外来植物入侵影响的研究往往关注资源的稳态变化, 忽略了资源的波动性, 特别是脉冲的作用。该文通过综述资源脉冲对外来植物入侵影响的研究, 简要评述了资源脉冲的形成原因、类型及影响, 讨论了不同类型的资源脉冲对外来植物入侵的作用。此外, 该文根据现有的研究进展提出了一些未来可能的研究方向, 如资源脉冲的不同属性, 多种资源脉冲交互作用对植物入侵的影响及其机制等。     

Using management to determine drivers of alien plant invasion and limits to native restoration

Ascertaining whether invasive species are the drivers or passengers of ecological change is crucial for restoration and for optimizing management. Smith et al. (this issue) show that failed control of an invasive forb limits restoration, regardless of whether native species are actively planted. Management‐based experiments that target the complementary processes of invasion and community assembly can help optimize restoration.     

The joint influence of environmental and anthropogenic factors on the invasion of two alien caulerpae in northwestern Mediterranean

Biological Invasions - Many factors are known to influence the invasion of alien species depending on the spatial scales considered. Caulerpa taxifolia and Caulerpa cylindracea are two green...