遗传多样性与外来物种的成功入侵: 现状和展望

遗传多样性被认为是影响外来种入侵成功的重要因素之一。研究表明, 尽管外来种在入侵过程中可能受到奠基者效应的影响, 但是多次引种、种内或种间杂交等过程使得许多外来种在引入地的遗传多样性水平未必会显著低于原产地, 从而使得外来种可能通过快速进化来适应引入地的新生境。然而, 高水平的遗传多样性并非成功入侵的必要条件, 遗传变异的匮乏对一些外来种的入侵能力没有明显的影响, 甚至在一些生物入侵案例中, 遗传多样性的降低反而促进了入侵成功。针对遗传多样性与入侵成功之间的复杂关系, 本文在评述外来种遗传多样性的研究现状的基础上, 分析了遗传多样性对外来种的短期入侵成功和长期进化的影响机制, 从方法角度探讨了目前研究中存在的若干问题, 并对如何推进入侵生态学研究提出了一些看法。正如一些学者提出的, 入侵生态学需要与生态学其他分支整合起来, 才能加深对生物入侵及其相关的生态和进化过程的理解。     

Predicting invasion success in complex ecological networks

A central and perhaps insurmountable challenge of invasion ecology is to predict which combinations of species and habitats most effectively promote and prevent biological invasions. Here, we integrate models of network structure and nonlinear population dynamics to search for potential generalities among trophic factors that may drive invasion success and failure. We simulate invasions where 100 different species attempt to invade 150 different food webs with 15–26 species and a wide range (0.06–0.32) of connectance. These simulations yield 11 438 invasion attempts by non-basal species, 47 per cent of which are successful. At the time of introduction, whether or not the invader is a generalist best predicts final invasion success; however, once the invader establishes itself, it is best distinguished from unsuccessful invaders by occupying a lower trophic position and being relatively invulnerable to predation. In general, variables that reflect the interaction between an invading species and its new community, such as generality and trophic position, best predict invasion success; however, for some trophic categories of invaders, fundamental species traits, such as having the centre of the feeding range low on the theoretical niche axis (for non-omnivorous and omnivorous herbivores), or the topology of the food web (for tertiary carnivores), best predict invasion success. Across all invasion scenarios, a discriminant analysis model predicted successful and failed invasions with 76.5 per cent accuracy for properties at the time of introduction or 100 per cent accuracy for properties at the time of establishment. More generally, our results suggest that tackling the challenge of predicting the properties of species and habitats that promote or inhibit invasions from food web perspective may aid ecologists in identifying rules that govern invasions in natural ecosystems.     

A conceptual framework for understanding arthropod predator and parasitoid invasions

Arthropods make up the largest group of invasive alien species (IAS) worldwide. Although invasion research has been biased towards alien plants and vertebrates, it has suggested potential mechanisms for the success of IAS and provided a theoretical framework for further investigation. Here we address key concepts from invasion biology that are essential to our understanding of the success of invasive alien arthropod predators and parasitoids including human intervention, environmental characteristics, propagule pressure, biological traits, and biological interactions. To gain a greater understanding of the factors most likely to influence the different stages of invasion (arrival, establishment, and spread) for alien arthropod predators and parasitoids, we use a comparative approach to compare and contrast the differential success of invasions by alien phytophagous and carnivorous arthropods. Insights gained from this comparison suggest that future research will require a multitrophic approach in order to enhance our understanding of invasions at higher trophic levels.     

Biotic indirect effects: a neglected concept in invasion biology

Indirect effects involve more than two species and are defined as how one species alters the effect that another species has on a third. These complex interactions are often overlooked in studies of interactions between alien and native species, and their role in influencing biological invasions has been rarely considered. Based on a comprehensive review of the invasion biology literature, we examine the evidence for the occurrence of four of the most commonly documented indirect effects (apparent competition, indirect mutualism/commensalism, exploitative competition, and trophic cascades) in the invasion process. Studies investigating indirect effects in the context of invasion biology are relatively rare, but have been increasing in recent years, and there are sufficient examples to indicate that this kind of interaction is likely to be more common than is currently recognized. Whether indirect interactions are mediated by an alien or a native species, and whether they occur between ecologically similar or dissimilar alien and native species, depends in part on the type of interaction considered and no predictable patterns were detected in the literature. Further empirical studies will help to elucidate such patterns. At this stage, the inherent unpredictability of indirect interactions means that their impacts in relation to invasions are particularly challenging for land managers to deal with, and their role in invasions is a complex, but is a valuable area of investigation for researchers.     

动物入侵的行为机制

行为特征可在外来动物建立种群和扩张过程中发挥重要作用,因此,要正确理解动物入侵,常常需要仔细研究其行为机制。20世纪80年代以来,随着动物入侵规模在世界各地的迅速加剧,有关其行为机制的研究也受到了广泛关注。最近一些研究表明,一些入侵动物种内攻击和觅食等行为具有可塑性,因此它们能够灵活应对多变的环境条件,这对于种群的建立和维持至关重要;入侵动物与土著物种发生行为互作时,往往占据优势,从而取代土著物种,并有助于其地域扩张;入侵动物长距离扩散可以提高其地域扩张速度,许多行为可与扩散行为结合进一步促进扩张。今后需要加强对入侵动物的行为分析,使之全面地融合到生物入侵的研究之中。这不仅可以提高对外来物种入侵的预警和治理能力,而且为探索动物行为的奥秘以及动物间行为互作在物种进化中的意义提供了独特的机会。     

Invasions by alien plant species of the agro-pastoral ecotone in northern China: Species-specific and environmental determinants

The establishment, reproduction, dispersal, and distribution of alien plants are affected by various factors during the transition from being newly introduced in a habitat to being invasive. In the agro-pastoral ecotone of northern China, comprising farmlands and natural grasslands, the biological characteristics of alien plant species were the key intrinsic factors (propagation characteristics and competitive ability), followed by such extrinsic factors as human interference and environmental heterogeneity. Among biological characteristics, the life form may be an important and useful indicator of the invasive ability of a species, and the risk of invasion is greater from alien species that are poisonous, inedible, and have traits that facilitate wide dispersal. Farmlands may serve as initial shelters for alien species, from which they spread into neighbouring habitats, whereas natural grassland may act as a barrier to plant invasions. Management practices detrimental to grasslands, including overgrazing, reclamation, and road construction, often facilitate the invasions; therefore, counter measures such as reseeding and a ban on grazing need special attention. Environmental factors including precipitation, nutrients, prevailing winds, fires, and topography may be other factors that promote or block the process of invasion. In studying ways of preventing or controlling such invasions, alien plants with short life cycle, prolific seed production, and strong competitiveness, deserve particular attention and so do human activities that may damage the environment and fragile habitats.     

Niche complementarity between an alien predator and native omnivorous fish in the Wilge River,South Africa

Hydrobiologia - A major challenge in biological invasions is to predict community susceptibility to invasion. This study investigated trophic interrelationships between an alien predator,...     

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.     

Geographical and taxonomic biases in invasion ecology

Invasive alien species come from most taxonomic groups, and invasion biology is searching for robust cross-taxon generalizations and principles. An analysis of 2,670 papers dealing with 892 invasive species showed that all major groups of invaders are well studied, but that most information on the mechanisms of invasion has emerged from work on a limited number of the most harmful invaders. A strong geographical bias, with Africa and Asia understudied, inhibits a balanced understanding of invasion, because we might be lacking knowledge of specific invasion mechanisms from poorly studied, regionally specific habitats. International cooperation is required to achieve a more geographically balanced picture of biological invasions. Invasive species with the greatest impact are best studied, but more studies of species that are naturalized but not (yet) invasive are needed to improve understanding of the mechanisms acting during the naturalization phase of invasions and leading to successful invasion.     

What affects the probability of biological invasions in Antarctica? Using an expanded conceptual framework to anticipate the risk of alien species expansion

Successful alien species invasion depends on many factors studied mostly in post invasion habitats, and subsequently summarized in frameworks tailored to describe the studied invasion. We used an existing expanded framework with three groups of contributing factors: habitat invisibility, system context and species invasiveness, to analyze the probability of alien species invasions in terrestrial communities of Maritime Antarctic in the future. We focused on the first two factor groups. We tested if the expanded framework could be used under a different scenario. We chose Point Thomas Oasis on King George Island to perform our analysis. Strong geographical barrier, low potential bioclimatic suitability and resource availability associated with habitat invasibility significantly reduce the likelihood of biological invasion in Antarctica. An almost full enemy release (low pressure of consumers), the high patchiness of the habitat, and the prevalence of open gaps also associated with habitat invasibility increase the possibility of invasion. The dynamics of functional connectivity, propagule pressure and spatio-temporal patterns of propagule arrival associated with human activity and climate change belonging to the system context contribute to an increase in the threat of invasions. Due to the still low land transport activity migration pathways are limited and will reduce the spread of alien terrestrial organisms by land. An effective way of preventing invasions in Antarctica seems to lie in reducing propagule pressure and eliminating alien populations as early as possible. The expanded conceptual framework opens up wider possibilities in analyzing invasions taking place in different systems and with multiple taxa.     

Macroecology of global bryophyte invasions at different invasion stages

In this study we provide the first comprehensive assessment of the environmental and anthropogenic factors driving bryophyte invasions worldwide. We compiled data of alien bryophyte distributions from 82 regions on five continents and oceanic islands and region specific variables. For each species, we collected data on its region‐specific invasion stage, i.e. casual (ephemeral) vs naturalized (persistent) occurrences, and we differentiated between known aliens and those which are likely to be alien (cryptogenic). We used these data to test how species attributes, environmental and socio‐economic conditions of the study areas as well as introduction effort affect invasion probabilities at different invasion stages and of known alien vs cryptogenic species. We collected information on species’ attributes (native range size and location, niche breadth, habitat affiliation), and calculated variables characterising the environmental, biogeographic and socio‐economic features of the native and recepient regions. Subsequently, we related the probability of alien occurrence across different invasion stages with these species‐ and region‐wise predictor variables using generalized linear mixed effects models. Greater native range size raised the likelihood that a species becomes alien or cryptogenic. Islands are more invaded by alien (and cryptogenic) bryophytes than continental regions. Native range size and socio‐economic activity increase the likelihood that a species becomes alien or cryptogenic elsewhere. Interestingly, among alien bryophytes naturalizations occur more frequently in regions of the complementary hemisphere than in regions of their native hemisphere. In general, regions in the Southern Hemisphere have higher numbers of naturalized bryophytes. We conclude that there is a conspicuous change in factors determining bryophyte invasions at different invasion stages. Whereas alien and cryptogenic bryophyte species occurrences are more frequent on islands and depend on native range size, and hence probably propagule pressure, naturalized bryophytes are more frequent in areas which are biogeographically separated but climatically similar to the native ranges.     

Parasites as Drivers and Passengers of Human-Mediated Biological Invasions

We provide an overview of the current state of knowledge of parasites in biological invasions by alien species. Parasites have frequently been invoked as drivers of invasions, but have received less attention as invasion passengers. The evidence to date that parasites drive invasions by hosts is weak: while there is abundant evidence that parasites have effects in the context of alien invasions, there is little evidence to suggest that parasites have differential effects on alien species that succeed versus fail in the invasion process. Particular case studies are suggestive but not yet informative about general effects. What evidence there is for parasites as aliens suggests that the same kind of factors determine their success as for non-parasites. Thus, availability is likely to be an important determinant of the probability of translocation. Establishment and spread are likely to depend on propagule pressure and on the environment being suitable (all necessary hosts and vectors are present); the likelihood of both of these dependencies being favourable will be affected by traits relating to parasite life history and demography. The added complication for the success of parasites as aliens is that often this will depend on the success of their hosts. We discuss how these conclusions help us to understand the likely effects of parasites on the success of establishing host populations (alien or native).

     

新建川藏铁路(雅安-昌都段)沿线外来入侵植物种类及分布特征

横断山区为全球生物多样性热点地区之一, 也是全国生态屏障的重要组成部分。新建川藏铁路雅安至昌都段横跨横断山核心地区, 铁路建设形成的交通网络将沿线生物多样性热点区域与外界相连, 导致生物入侵风险陡增。为获得区域内外来入侵植物的种类及分布特征信息, 为即将开始的铁路工程建设、生态保护及生态修复等工作提供参考, 我们在雅安-昌都段内选择43个位点各进行长度1 km、宽度20 m的样线调查。研究结果显示: 雅安-昌都段共发现外来入侵植物58种, 隶属于18科42属, 其中出现频度最高的种类依次是牛膝菊(Galinsoga parviflora)、秋英(Cosmos bipinnatus)和鬼针草(Bidens pilosa)。从危害等级来看, 其中10种为恶性入侵种, 16种为严重入侵种, 8种为局部入侵种, 15种为一般入侵种, 9种为有待观察种, 超过半数种类具有明显入侵性。原产地分析结果显示美洲是该区域外来入侵植物的主要原产地。基于海拔及主要河流区段的比较研究发现: 入侵植物的种类数量呈现出明显的由东向西、由低海拔向高海拔逐渐递减的趋势, 该分布格局是环境因子和人类活动共同作用的结果。结合铁路沿线入侵现状和生境特征, 本文分析了铁路建设可能造成的外来植物入侵风险, 并针对入侵的防范提出了相应的建议。     

Parasites as Drivers and Passengers of Human-Mediated Biological Invasions

We provide an overview of the current state of knowledge of parasites in biological invasions by alien species. Parasites have frequently been invoked as drivers of invasions, but have received less attention as invasion passengers. The evidence to date that parasites drive invasions by hosts is weak: while there is abundant evidence that parasites have effects in the context of alien invasions, there is little evidence to suggest that parasites have differential effects on alien species that succeed versus fail in the invasion process. Particular case studies are suggestive but not yet informative about general effects. What evidence there is for parasites as aliens suggests that the same kind of factors determine their success as for non-parasites. Thus, availability is likely to be an important determinant of the probability of translocation. Establishment and spread are likely to depend on propagule pressure and on the environment being suitable (all necessary hosts and vectors are present); the likelihood of both of these dependencies being favourable will be affected by traits relating to parasite life history and demography. The added complication for the success of parasites as aliens is that often this will depend on the success of their hosts. We discuss how these conclusions help us to understand the likely effects of parasites on the success of establishing host populations (alien or native).     

可利用资源波动对外来种入侵的抵抗性

生物入侵带来的生态和经济危害引起了人们的广泛关注。在入侵生态学研究方面,生物多样性与生物入侵之间的关系长久以来成为群落可入侵性探讨的焦点。Elton经典假说认为,物种多样性越高对外来种入侵的抵抗能力越强,许多模型或野外试验都支持这一假说。但现在越来越多的试验对此提出了异议,各种假说纷纷出现。究竟生物多样性会不会影响外来种的入侵?假设两种不同的群落结构(功能群),设计6种外来种入侵土著群落的情景分析不同多样性及相同多样性下外来种的入侵状况。结果发现,在多样性相同的情况下,两种群落对外来种入侵的抵抗力不同。外来种成功入侵等比群落,却被倍数群落排斥在系统之外。进一步分析表明这主要是由于可利用资源的波动引起的,即Davis提出的"资源机遇假说"。在相同的物种多样性下,由于倍数群落的特殊结构,整个群落所占有资源远远大于等比群落资源比率。因此,外来种在等比群落中更易找到合适的入侵机会。而在物种多样性不同的情况下,由于物种多样性与已占有资源的变化是成正比的,因此,混淆了多样性与剩余资源可利用性对外来种入侵的影响。     

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。     

气候变化背景下生物入侵研究态势的文献计量分析

生物入侵严重威胁生物多样性与生态系统健康,对全球环境、经济造成极大损失,而快速的气候变化显著影响外来生物的扩散和入侵进程。探讨气候变化背景下生物入侵研究态势能够从宏观上把握该领域的国际研究现状与热点,为深入理解外来种入侵机制和制定合理的防治策略提供参考。该文基于最近27 a间(1990—2016年)科学文献数据库Web of Science中科学引文索引扩展版(SCI-E)数据,利用TDA等统计工具对气候变化下生物入侵方面的研究进行了文献计量分析。结果表明:27 a间共发表论文1 736篇,论文数量整体保持增长态势,2009年开始进入快速发展阶段;该领域的研究涉及环境科学与生态学、生物多样性保护、植物学等多个学科;澳大利亚莫纳什大学Chown SL教授发文量最高(35篇);美国的总发文量(708篇)和高被引、高影响因子论文数量均居世界首位;发文量最多的研究机构是加利福尼亚大学(93篇),中国科学院发文量居世界第10位(27篇);Biological Invasions是刊文量最大的学术杂志;物种分布模型、生物多样性、全球变暖、风险评估等是近年来该领域的研究热点;中国共发表论文52篇,中国科学院是国内最大的发文机构,其中,动物研究所、武汉植物园、植物研究所的发文量居中科院科研系统前三名;中国在气候变化下生物入侵领域的高被引、高影响因子论文数量及国际合作强度亟待提升。未来需重点关注气候变化下生物入侵的预测与风险评估、生物入侵与生物多样性关系、入侵物种的系统进化、入侵生态系统的多营养级关系、海洋生物入侵、生物入侵与人类健康等问题。     

基于分子生物学方法的外来入侵物种入侵历史重构

生物入侵是一个世界性的问题。全球每年因生物入侵造成的损失超过1万亿美元。探究入侵物种在入侵地的入侵历史对了解生物入侵的生物生态学机制、制定阻截及防除措施有重要意义。分子标记方法的兴起和大规模应用打开了入侵生物入侵历史研究的新天地。采用分子标记的方法可鉴定入侵物种的种类、追溯其来源地、回溯其扩散路径、分析扩散模式及探究物种入侵过程中对入侵种群本身的变化及其对生态系统所造成的各种影响。分子标记的应用使得多个入侵物种的入侵历史得以重现。由于分子标记方法重构的入侵历史受采样范围、采用的分子标记的种类及数量等因素的影响,该方法呈现入侵历史是否是真实发生的入侵过程还存在争议。     

Impact of invasions by alien plants on soil seed bank communities: Emerging patterns

Much of our current understanding of the impact of invasive species on plant communities is based on patterns occurring in the above-ground vegetation, while only few studies have examined changes in soil seed banks associated with plant invasions, despite their important role as determinants of vegetation dynamics. Here, we reviewed the literature on the impact of plant invasions on the seed bank and we provide a quantitative synthesis using a meta-analysis approach. Specifically, (1) we quantified the impact of 18 invasive alien plants on (i) species richness and (ii) density of the seed banks of invaded communities, based on 58 pair-wise invaded-uninvaded comparisons (cases); we identified (2) the invasive taxa that are responsible for the largest changes in the seed bank; and (3) the habitats where substantial changes occur. Our study showed three major findings: (1) species richness (68% of cases) and density (58% of cases) were significantly lower in native seed banks invaded by alien plants; (2) species richness and density of native and alien species were remarkably lower in seed banks invaded by large, perennial herbs compared to uninvaded sites; and (3) invaded seed banks were often associated with a larger richness and/or abundance of alien species. This study indicates a need for additional seed bank data in invasion ecology to characterize species-specific and habitat-specific impacts of plant invasions, and to determine whether changes in the seed banks of native and alien species are a symptom of environmental degradation prior to a plant invasion or whether they are its direct result. The findings of this study help improve our capacity to predict the long-term implications of plant invasions, including limitations in the recruitment of native species from the seed bank and the potential for secondary invasions by seeds of other alien species.     

Invasion of the cosmopolitan species Echinochloa colona into herbaceous vegetation of a tropical wetland system

The negative effects of alien plant species on ecosystem structure and functions are increasingly recognised, and efforts to control these species are vital to restore degraded ecosystems and preserve biodiversity. However, we lack a full understanding of factors that determine alien species invasions along spatial gradients in herbaceous vegetation of tropical systems. We therefore examined the effects of community properties, environmental variables and human-related disturbance factors on the invasion of the alien grass Echinochloa colona (L.) Link at small- and large scales in the Kilombero Valley wetland, Tanzania. Generalized additive mixed models showed that E. colona abundance on a small scale was negatively related to above-ground biomass and evenness of resident species, whereas E. colona abundance was positively related to grazing intensity. On a large scale, biomass (negatively related to E. colona abundance) and distance to river (positive) were important in explaining E. colona abundance. These findings support the assertion that different factors may contribute to the invasion of alien plant species at different spatial scales, as also reported in many temperate systems. Overall, our results show that successful invasion of alien species is a function of plant community properties, human-related disturbance and favourable environmental conditions. Effective management strategies should consider mitigations that can increase the biomass and evenness of native species and a reduction of grazing pressure to restore the wetland and conserve biodiversity.