Are some communities of the coastal dune zonation more susceptible to alien plant invasion?

Aims Coastal areas, and in particular coastal dunes, are ecosystems strongly affected by the invasion of alien plants. However, few attempts have ever been made to quantify alien species incidence in different communities along the coastal zonation. This work aims to analyze the distribution of alien plants along the coastal zonation of sandy shores on the Tyrrhenian coast, addressing specifically differences among plant communities in abundance of alien plants.Methods The study was performed on recent dunes (Holocene) of the central western coast of Italy. We selected dune landscapes where invasion processes were particularly evident. Vegetation plots were randomly sampled and through cluster analysis, we identified six plant communities corresponding to the typical zonation described for the Tyrrhenian sandy coast of Central Italy. We evaluated and compared frequency and abundance of invasion in these different communities. Further, we investigated how propagule pressure (measured using as proxy human structures) contributed to the observed invasion patterns.Important findings We found a relatively low total number of aliens but also a differential distribution pattern and strong abundance of some of the aliens in specific sectors of the vegetation zonation. The perennial community of transition dunes appears most affected by invasion processes, related almost exclusively to the frequent and widespread Carpobrotus aff. acinaciformis. This alien species reaches high cover values, apparently lowering cover of native species of transition dune plant communities. Higher levels of invasion in the transition dune can be partially explained because of greater propagule pressure in this section of the dune profile. Our findings thus have important conservation and management implications since transition dune communities with Crucianella maritima are rare and protected (sensu Habitat 92/43/EEC Directive) along the entire Italian coast.     

植物功能性状与外来植物入侵

揭示影响外来植物入侵性的功能性状及其生态机制是入侵植物生态学的核心任务之一。本文综述了植物功能性状与外来植物入侵性的研究进展, 通过分析植物功能性状对外来植物入侵的贡献以及外来植物的不同入侵阶段对其功能性状的需求, 探讨植物功能性状与外来植物入侵的相关性及其入侵机理。迄今研究较多的影响外来植物入侵性的功能性状主要包括形态性状、生长性状、生理性状、繁殖性状、种子性状、克隆性状、表型可塑性和遗传变异等。这些功能性状对外来植物入侵的贡献随着入侵阶段的不同而变化。在传播到达阶段, 种子性状对入侵具有重要影响; 在定居建群阶段, 与植物抗逆性和适应性相关的生理性状和繁殖性状发挥主要作用; 在扩散入侵阶段, 克隆性状和影响植物竞争能力的生理性状对植物成功入侵具有重要贡献。由于植物入侵性是其功能性状和环境因素互作的结果, 且功能性状的作用随环境因素和入侵阶段不同而异, 因此, 结合外来植物入侵阶段, 并考虑功能性状与环境因子的互作, 是入侵生物学中植物功能性状研究的发展趋势。     

外来入侵植物对土壤氮转化主要过程及相关微生物的影响

外来入侵植物不仅影响植物群落组成、生物多样性以及生态系统的结构和功能, 而且显著影响土壤氮(N)的转化过程。外来入侵植物对N循环影响的研究已成为入侵生态学的研究热点。N循环与凋落物的分解和养分释放有关, 外来入侵植物能够改变凋落物的组成与结构, 进而影响土壤的N转化过程。另外, 外来入侵植物的化感作用也会影响土壤N转化过程, 这些作用与土壤微生物的结构与功能变化密不可分。该文主要从凋落物分解与养分释放及外来入侵植物化感作用两个方面综述了外来入侵植物对土壤N转化的影响, 总结了外来入侵植物对土壤N转化相关土壤微生物(尤其是氨氧化细菌与氨氧化古菌)的影响, 探讨了土壤N转化对外来植物入侵的反馈, 并探讨了丛植菌根真菌与外来入侵植物的互相影响。     

Effects of plant invasion on soil nitrogen transformation processes and its associated microbes

外来入侵植物不仅影响植物群落组成、生物多样性以及生态系统的结构和功能, 而且显著影响土壤氮(N)的转化过程。外来入侵植物对N循环影响的研究已成为入侵生态学的研究热点。N循环与凋落物的分解和养分释放有关, 外来入侵植物能够改变凋落物的组成与结构, 进而影响土壤的N转化过程。另外, 外来入侵植物的化感作用也会影响土壤N转化过程, 这些作用与土壤微生物的结构与功能变化密不可分。该文主要从凋落物分解与养分释放及外来入侵植物化感作用两个方面综述了外来入侵植物对土壤N转化的影响, 总结了外来入侵植物对土壤N转化相关土壤微生物(尤其是氨氧化细菌与氨氧化古菌)的影响, 探讨了土壤N转化对外来植物入侵的反馈, 并探讨了丛植菌根真菌与外来入侵植物的互相影响。     

Effects of plant invasion on soil nitrogen transformation processes and it’s associated microbial北大核心CSCD

Invasive alien plants not only influence plant community composition, biodiversity and ecosystem structure and function, but also have severe impacts on soil nitrogen transformation processes. The effects of invasive alien plants on nitrogen (N) cycling have been one of the hot topics in invasion ecology. Litter decomposition and its nutrient release play an important role in nutrient cycling. In addition, invasive alien plants have the potential to influence soil N transformation through allelopathy. All these processes are tightly related to soil microbes. Therefore, this review mainly focuses on litter decomposition and its nutrient release, and allelopathy to understand the effects of plant invasion on soil N transformation. Changes in soil N transformation and soil microbes (esp. Ammonia oxidizing bacteria and Ammonia oxidizing archaea) due to plant invasion, as well as the feed-backs of these changes to further invasion of alien plants were discussed. Finally, the interactions between arbuscular mycorrhizal fungi and plant invasion were reviewed. © Chinese Journal of Plant Ecology     

Release from belowground enemies and shifts in root traits as interrelated drivers of alien plant invasion success: a hypothesis

Our understanding of the interrelated mechanisms driving plant invasions, such as the interplay between enemy release and resource‐acquisition traits, is biased by an aboveground perspective. To address this bias, I hypothesize that plant release from belowground enemies (especially fungal pathogens) will give invasive plant species a fitness advantage in the alien range, via shifts in root traits (e.g., increased specific root length and branching intensity) that increase resource uptake and competitive ability compared to native species in the alien range, and compared to plants of the invader in its native range. Such root‐trait changes could be ecological or evolutionary in nature. I explain how shifts in root traits could occur as a consequence of enemy release and contribute to invasion success of alien plants, and how they could be interrelated with other potential belowground drivers of invasion success (allelopathy, mutualist enhancement). Finally, I outline the approaches that could be taken to test whether belowground enemy release results in increased competitive ability and nutrient uptake by invasive alien plants, via changes in root traits in the alien range.     

潮虫消耗木本植物凋落物的可选择性试验

近年来,越来越多的学者关注外来植物入侵对土壤生态功能的影响效应及其相应反馈机制的探索与研究,然而本地原生土壤生物群落对不同入侵程度下的外来植物种以及本地原生植物种之间是否存在消耗差异却尚不明了.通过等足目潮虫的选择性喂养试验来测试10个本地种、5个非入侵性外来种和5个强入侵性外来种之间的适口性差异,试图求证外来植物的入侵性是否与植物落叶被消耗率呈现必然联系.数据分析结果显示潮虫对本地种、非入侵性外来种和入侵性外来种的消耗并无显著差异;而潮虫对不同生活型下木本植物的取食却存在显著差异,即灌木消耗率显著高于乔木.其次,通过植物初始性状指标(包括木质素、纤维素、半纤维素、碳、氮含量)与相应消耗率的相关比较,消耗量总体上与植物凋落物的氮含量呈正相关关系(R2 =0.358).由此,研究结论强调植物落叶的降解速率并不一定与植物入侵性或入侵阶段呈绝对相互关连,但是氮含量,抑或各种形式的植物氮元素成分都可能在一定程度上参与并影响着外来植物的入侵进程.     

Interactions between alien plant species traits and habitat characteristics in agricultural landscapes in Finland

The survival and success of alien plant species is determined by species traits (i.e., invasiveness) and the characteristics of the habitats in the region of introduction (i.e., invasibility). However, little is known about species traits as related to habitat characteristics. We assessed the characteristics of successful invaders and the interaction of environmental factors and life-history traits for alien plant species. The vascular plants were recorded from 52 agricultural landscapes in Finland. We compared the traits of native and alien plant species with Fisher’s exact test and used a three table ordination analysis, RLQ analysis, to relate species traits to environmental conditions. Species were clustered according to their position on the RLQ axes, and the clusters were tested for phylogenetic independence. The successful alien plant species were associated with life form and preferences for moisture and nitrogen, but the trait composition varied according to the habitat type. Two RLQ axes explained 80.5% of the variation, and the species traits were significantly associated with environmental variables. The clustering showed that the occurrence of alien plant species in agricultural habitats was driven by invasion history, traits related to dispersal (dispersal type, seed mass) and habitat preferences, as well as environmental features, such as geographical location, temperature and the quality and disturbance regime of the habitats. All clusters were phylogenetically non-independent. Thus, the clusters of alien species comprised species of diverse taxonomic affinities, although, they shared the traits explaining their occurrence in particular habitats. This information is useful for understanding the link between species traits and the environmental conditions of the habitats, and complexity of the invasion process.     

Getting the Right Traits: Reproductive and Dispersal Characteristics Predict the Invasiveness of Herbaceous Plant Species

To better understand the effect of species traits on plant invasion, we collected comparative data on 20 reproductive and dispersal traits of 93 herbaceous alien species in the Czech Republic, central Europe, introduced after 1500 A. D. We explain plant invasion success, expressed by two measures: invasiveness, i.e. whether the species is naturalized but non-invasive, or invasive; and dominance in plant communities expressed as the mean cover in vegetation plots. We also tested how important reproductive and dispersal traits are in models including other characteristics generally known to predict invasion outcome, such as plant height, life history and residence time. By using regression/classification trees we show that the biological traits affect invasion success at all life stages, from reproduction (seed production) to dispersal (propagule properties), and the ability to compete with resident species (height). By including species traits information not usually available in multispecies analyses, we provide evidence that traits do play important role in determining the outcome of invasion and can be used to distinguish between alien species that reach the final stage of the invasion process and dominate the local communities from those that do not. No effect of taxonomy ascertained in regression and classification trees indicates that the role of traits in invasiveness should be assessed primarily at the species level.     

Landscape context modulates alien plant invasion in Mediterranean forest edges

Natural habitats in human-altered landscapes are especially vulnerable to biological invasions, especially in their edges. We aim to understand the influence of landscape and local characteristics on biological invasions by exploring the level of plant invasion and alien species traits in forest edges in highly urbanized landscapes. We identified all plant species in 73 paired plots in the edge and 50 m towards the interior of the forest. We explored the association between alien species richness and similarity in species composition between edge and interior plots with landscape and local variables, using generalized linear models and variance partitioning techniques. Then, we performed Fourth-corner analyses to explore the association between alien plant traits and local and landscape variables. In contrast to native species richness, alien species richness was more affected by the surrounding landscape than by the local characteristics of the edge. Road proximity was positively associated with alien species richness and proportion and was its most important correlate, whereas disturbance was negatively associated with native species richness and was its most influential factor. Alien plant traits were also primarily associated with landscape characteristics. For instance, species of Mediterranean origin and introduced for agriculture were associated with higher agriculture use in the landscape. Our findings suggest that risk analyses of habitat vulnerability to invasion must consider the landscape context in order to successfully predict highly invaded areas and identify potentially successful invaders.     

Evaluating differences in the shape of native and alien plant trait distributions will bring new insights into invasions of plant communities

Failure to quantify differences in the shape of inter‐specific trait distributions (e.g., skew, kurtosis) when comparing co‐occurring alien and native plants hinders the integration of biological invasions and plant community ecology. Within a plant community, understanding the circumstances that lead to the shape of the inter‐specific distribution of one or more functional plant traits being unimodal, bimodal, multimodal or skewed has the potential to shed new light on community vulnerability to invasion, subsequent ecosystem impacts and the selection pressures (e.g., stabilizing, directional or disruptive) acting upon native and alien species. Ignoring differences in the shape of inter‐specific trait distributions of alien and native species could miss important insights into plant invasions, including: the existence of unsaturated native plant communities, empty niches, shifting trait optima of species as a result of environmental change and incomplete colonization–extinction processes following invasion. Future comparisons of functional trait differences between native and alien species should include assessment of the shapes of inter‐specific trait distributions since these may differ even when the mean values of traits are similar for native and alien species. The infrequent application of such approaches may explain the limited generalizations regarding the drivers and consequences of plant invasions in plant communities.     

入侵植物的生理生态特性对碳积累的影响

随着国际贸易的发展和人们交往的增加以及全球环境的变化,生物种类在全球扩散的机会也大大增加,从而为生物入侵创造了机会。生物入侵不仅给农林牧生产造成损失,而且具有长期的生态学效应。外来种的成功入侵不是其自身某一个特性决定的,而是其特性与新的生境综合作用的结果。外来入侵种生理生态特性的研究对其预测和防治具有重要的意义。目前对入侵种生理生态特性的研究较少。已有的研究表明,与本地种相比入侵种可能通过提高光合能力、资源利用率、表型可塑性、化感作用,以及降低繁殖成本等增加植株碳积累,促进其入侵。但并不是所有的入侵种都同时具有这些特性。生境不同限制性资源不同,入侵机制就不同。成功的入侵种应该能够高效地利用生境中的限制性资源,并且能够较快地调节自身的生理特性以适应波动的资源环境。     

Recruitment limitation of native species in invaded coastal dune communities

Recruitment limitation may limit the ability of sites to regenerate after disturbances such as weed invasion and weed management. We investigated seed bank constraints and dispersal limitation in coastal dune communities on the east coast of Australia. The ability of sites to regenerate naturally following weed removal was assessed in coastal dune communities invaded by the invasive alien, bitou bush (Chrysanthemoides monilifera subsp. rotundata). To investigate recruitment limitation, seed banks and vegetation of invaded, native, intensively managed (selective application of herbicide and some re-vegetation) and extensively managed (large-scale, non-selective herbicide application) sites were compared. We investigated the dispersal mechanisms of species in the seed bank and vegetation to determine if communities might be dispersal-limited, i.e. contain significant numbers of species with only short-distance dispersal capabilities. Species richness and composition of soil seed banks differed from the vegetation in foredunes and hinddunes. Invasion depleted seed banks further. About half of the species had short-distance dispersal mechanisms indicating the potential for dispersal limitation. Secondary weed invasion following management was evident although alien species occurred in both seed banks and vegetation. Our results indicated that coastal dune communities suffer recruitment limitation. Native, managed and invaded dune communities appear to be both seed bank and dispersal-limited although management and invasion exacerbates recruitment. Regeneration of coastal dune communities will require active reintroduction of species, particularly those with short-distance dispersal mechanisms.     

Are boreal ecosystems susceptible to alien plant invasion? Evidence from protected areas

Although biological invasion by alien species is a major contributor to loss of indigenous biological diversity, few studies have examined the susceptibility of the boreal biome to invasion. Based on studies of other ecosystems, we hypothesized that alien plants will be restricted to disturbed areas near human activity and will not be found in natural areas of boreal ecosystems in Gros Morne National Park (Canada), a protected area experiencing a wide range of disturbance regimes. The distribution of alien plants in the region was evaluated using surveys, and study sites were established in naturally and anthropogenically disturbed habitats that had been invaded. Within study sites, randomization tests evaluated the importance of disturbance to alien plant invasion by examining changes in environmental conditions and species abundance within various disturbance regimes, while the importance of site characteristics limiting the distribution of alien plants were examined using Canonical Correspondence Analysis. Consistent with studies in a variety of biomes, areas of high disturbance and human activity had the greatest abundance of resources and the highest percentage of alien species. However, contrary to our hypothesis, natural areas of boreal ecosystems were found susceptible to alien plant invasion. Vegetation types vulnerable to invasion include forests, riparian areas, fens, and alpine meadows. Natural disturbance occurring in these vegetation types caused increases in bare ground and/or light availability facilitating alien plant invasion. Although high soil pH was associated with alien plants in these areas, disturbance was not found to cause changes in soil pH, suggesting susceptibility to invasion is pre-determined by bedrock geology or other factors influencing soil pH. Moose (Alces alces), a non-native herbivore, acts as the primary conduit for alien plant invasion in GMNP by dispersing propagules and creating or prolonging disturbance by trampling and browsing vegetation. The recurrent nature of disturbance within the boreal biome and its interaction with site conditions and herbivores enables alien plants to persist away from areas of high human activity. Managers of natural lands should monitor such interactions to decrease the invasion potential of alien plants.     

The invasive alien plant species Solidago gigantea alters ecosystem properties across habitats with differing fertility

Question: Invasive alien plants can affect biomass production and rates of biogeochemical cycling. Do the direction and intensity of such effects depend upon the functional traits of native and alien species and upon the properties of the invaded habitat, with the same alien species having differing impacts in different habitats? Location: Lowlands of Switzerland. Methods: Fourteen grassland and wetland sites invaded by Solidago gigantea and widely differing in biomass production and soil P availability were surveyed. To determine whether the impact of the species was related to site fertility, we compared the invaded and native vegetation in terms of biomass, species composition, plant traits and soil properties. Results: S. gigantea generally increased the above‐ground biomass production of the vegetation and soil C content, while reducing nutrient concentrations in biomass and N availability in the soil. However, it had no significant effect on plant species richness, soil respiration, soil pH and P availability. Leaves of S. gigantea had a greater C content than those of native species; other leaf traits and root phosphatase activity did not differ significantly. Conclusions: Our results suggest that a conservative nutrient‐use strategy allows S. gigantea to invade a broad range of habitats. The observed effects of invasion did not vary according to biomass production of the invaded sites, but some effects did depend on soil P availability, being more pronounced at more P‐rich sites. Thus, the full range of invaded habitats should be considered in studying the potential impact of plant invasions on ecosystem processes.     

Impact of alien species on dune systems: a multifaceted approach

We applied a multifaceted approach, in terms of taxonomic, phylogenetic and functional diversity, to study at fine scale how three plant communities occurring in a Mediterranean dune have been affected by the encroachment of alien species. We sampled 81 sites in a Site of Community Importance in Central Italy. Past and present land use/cover data have been derived using GIS and remote sensing tools. Information on plants phylogenesis and functional traits has been gathered from several databases. Ecological variables have been collected. GLMs in conjunction with an Information Based approach were used to model species composition, richness and phylogenetic diversity. Multivariate analysis has been used to study functional diversity. The results outlined how total species richness is related to recent land transformations and to a set of environmental factors. The analyses of functional and phylogenetic diversity support the idea that alien species significantly affect the functional and phylogenetic characteristics of the native plant communities. Habitat filtering seems to be predominant in not-invaded plots, whereas limiting similarity/niche differentiation is predominant in driving community assembly of invaded communities. The attained scenario depicts the spread of a reduced group of alien species phylogenetically and functionally well-differentiated, able to reduce the abundance of native species, not to exclude them though. Ultimately, the multifaceted approach assisted in understanding the community assembly of dune vegetation, and to discern the relative impact of alien species on native plant communities. Such approach represents a crucial step to achieve an efficient management of dune habitats, as useful tool to monitor and to effectively protect their biodiversity and functioning.     

Understanding the influence of urbanization on invasibility: <Emphasis Type="Italic">Carpobrotus edulis</Emphasis> as an exemplar

Coastal dune areas are valuable ecosystems, generally impacted by habitat destruction and invasive alien species. In this study, we assessed how human disturbance and invasion by Carpobrotus edulis impact the soils and the establishment of native flora in the north-western coastal regions of Spain. We compared soil characteristics (pH, conductivity, water content, nutrients and enzymatic activities) and native plant as well as C. edulis fitness correlates (germination and early growth) between uninvaded and invaded soils from urban and natural coastal dune areas. We found that human disturbance impacts coastal soils by increasing organic matter and water content, modifying soil nutrients and cycles, and reducing the pH in urban soils. The presence of invasive C. edulis further increases these impacts. These changes in soil characteristics allow for the establishment of the native, but ruderal, Scolymus hispanicus and non-native C. edulis, both of which are not adapted to the typically limiting conditions of coastal dunes. In some instances, the coastal dune endemic, Malcolmia littorea, showed no fitness effects in response to urbanization or the presence of C. edulis. These results suggest that human disturbed coastal areas might be more easily invaded than natural areas. More broadly, our findings of differential responses of different native species to disturbance and invasion, illustrate the need for multi-taxon approaches when assessing the impacts of invasive species.     

Growth‐form and spatiality driving the functional difference of native and alien aquatic plants in Europe

Trait‐based approaches are widely used in community ecology and invasion biology to unravel underlying mechanisms of vegetation dynamics. Although fundamental trade‐offs between specific traits and invasibility are well described among terrestrial plants, little is known about their role and function in aquatic plant species. In this study, we examine the functional differences of aquatic alien and native plants stating that alien and native species differ in selected leaf traits. Our investigation is based on 60 taxa (21 alien and 39 native) collected from 22 freshwater units of Hungarian and Italian lowlands and highlands. Linear mixed models were used to investigate the effects of nativeness on four fundamental traits (leaf area, leaf dry matter content, specific leaf area, and leaf nitrogen content), while the influence of growth‐form, altitude, and site were employed simultaneously. We found significantly higher values of leaf areas and significantly lower values of specific leaf areas for alien species if growth‐form was included in the model as an additional predictor.We showed that the trait‐based approach of autochthony can apply to aquatic environments similar to terrestrial ones, and leaf traits have relevance in explaining aquatic plant ecology whether traits are combined with growth‐forms as a fixed factor. Our results confirm the importance of traits related to competitive ability in the process of aquatic plant invasions. Alien aquatic plants can be characterized as species producing soft leaves faster. We argue that the functional traits of alien aquatic plants are strongly growth‐form dependent. Using the trait‐based approach, we found reliable characteristics of aquatic plants related to species invasions, which might be used, for example, in conservation management.     

Plant functional diversity enhances associations of soil fungal diversity with vegetation and soil in the restoration of semiarid sandy grassland

The trait‐based approach shows that plant functional diversity strongly affects ecosystem properties. However, few empirical studies show the relationship between soil fungal diversity and plant functional diversity in natural ecosystems. We investigated soil fungal diversity along a restoration gradient of sandy grassland (mobile dune, semifixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China, using the denaturing gradient gel electrophoresis of 18S rRNA and gene sequencing. We also examined associations of soil fungal diversity with plant functional diversity reflected by the dominant species' traits in community (community‐weighted mean, CWM) and the dispersion of functional trait values (FD_is). We further used the structure equation model (SEM) to evaluate how plant richness, biomass, functional diversity, and soil properties affect soil fungal diversity in sandy grassland restoration. Soil fungal richness in mobile dune and semifixed dune was markedly lower than those of fixed dune and grassland (P < 0.05). Soil fungal richness was positively associated with plant richness, biomass, CWM plant height, and soil gradient aggregated from the principal component analysis, but SEM results showed that plant richness and CWM plant height determined by soil properties were the main factors exerting direct effects. Soil gradient increased fungal richness through indirect effect on vegetation rather than direct effect. The negative indirect effect of FDis on soil fungal richness was through its effect on plant biomass. Our final SEM model based on plant functional diversity explained nearly 70% variances of soil fungal richness. Strong association of soil fungal richness with the dominant species in the community supported the mass ratio hypothesis. Our results clearly highlight the role of plant functional diversity in enhancing associations of soil fungal diversity with community structure and soil properties in sandy grassland ecosystems.     

基于功能性状的外来植物入侵预测模型框架构建

预测外来植物的潜在入侵性已成为生物多样性保护研究的重要内容,外来植物与乡土物种间的亲缘关系是预测外来植物能否成功入侵的一个重要途径。然而,达尔文归化难题却预测了两种截然不同的结果(即达尔文归化假说和预适应假说)。该研究解析了达尔文归化难题的内涵,提出了基于功能性状的外来植物与乡土群落间的相似性关系应该是进行外来植物入侵预测的重要切入点,而功能性状的种间分化与种内变异可能是外来植物成功入侵的两种不同生态策略。在此基础上,该研究还通过物种功能性状的多维超体积构建了外来植物与乡土群落间的相似性,提出了基于这种相似性的外来植物入侵预测的研究框架和基本流程。该模型框架的建立有助于理解外来植物的入侵机制,对外来植物的潜在入侵性预测提供了理论依据。当然,要实现外来植物能否成功入侵的准确预测,不仅依赖于功能性状的选择,还要考虑入侵的生境依赖性、空间尺度的重要性以及乡土群落的可入侵性等,未来的研究重点是通过控制实验对该模型进行验证和进一步完善。