外来植物成功入侵的生物学特征

外来植物的入侵能力与其性状之间的关系是入侵生态学中的基本问题之一.成功的入侵种常常能占据多样化的生境,具有较强的适应性、繁殖力和散布力.表型可塑性和遗传分化是外来入侵植物对生境异质性的两种适应策略;散布体多态型和散布途径多种化,使外来入侵植物迅速占领入侵生境,并进行远距离扩散;无性生殖和有性生殖并存,并根据生境和入侵阶段权衡的繁育对策不仅使入侵种群大面积暴发成为可能,而且直接影响散布机制并对种群遗传结构具有调节作用.高效的资源利用性竞争,加之以化感作用为基础的干扰性竞争使植物更具入侵性.     

中国东部沿海互花米草种群生活史特征的纬度变异与可塑性

互花米草(Spartina alterniflora)于20世纪70年代被引入中国,目前已在东部沿海盐沼湿地中广泛分布,成为海岸带盐沼中危害严重的入侵植物之一.为了研究互花米草在中国入侵区中的适应机制,揭示遗传分化和表型可塑性在该物种成功入侵中的作用,本研究沿纬度梯度在南起广东(22°N)、北至天津(39°N)的沿海样带上采集了10个种群的样本,通过同质园实验比较了不同纬度来源的种群在,生活史和生长特征方面是否存在遗传分化,并平行设置高低两个水位处理以比较互花米草对水位变化(不同高程生境条件)的可塑性反应.结果表明,在所研究的互花米草17个性状中有12个存在显著的种群间差异.其中,平均开花日期和相对生长率(植株高度)表现出显著的纬度梯度变异:随着纬度的升高,开花时间提前,相对生长速率(植株高度)趋于增加.同时17个性状中有9个在不同水位处理之间存在显著差异.这些结果表明,遗传分化可能是互花米草能够快速占据广阔分布区的重要原因之一,而表型可塑性可能对互花米草在小尺度上占据不同高程环境的过程有重要作用.     

外来种入侵的生物学与生态学基础的若干问题

外来物种的入侵及其所造成的危害已越来越为人们所认识。一个外来物种入侵一个新的生境无疑要具备一定的条件 ,尽管目前对这种条件的研究还只是初步的。首先从其自身来说 ,需要有足够的入侵性 ,这可以认为是入侵种的生物学基础 ,可能表现为具有多倍性、一定的遗传变异、杂合性或表型可塑性 ,以及不同的交配系统等 ;其次一个物种的入侵成功还取决于入侵生境的可入侵性 ,这可以认为是入侵种的生态学基础 ,而新的种间关系的形成 ,尤其是入侵种与其他种类的互惠共生关系的建成是关键。本文主要介绍了有关入侵种的生物学和生态学基础中的一些最新进展 ,希望有助于我国学者对相关研究内容的开展以及对有害外来种入侵的防治     

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。     

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。     

植物的表型可塑性、异速生长及其入侵能力

表型可塑性是指同一个基因型对不同环境响应产生不同表型的特性,特定性状的可塑性本身可以遗传,也可以接受选择而发生进化。植物个体的异速生长是指生物体某一特征的相对生长速率不等于第二种特征的相对生长速率的特性,该特性是由物种的遗传性决定的一种固定特征,植物往往朝着最佳的异速生长曲线进化。植物特定基因型在不同环境下,诸如生物量分配和种群几何学上的一些表型差异,既可由异速生长造成,也可由表型可塑性造成。植物本身的异速生长是一种"外观可塑性",而异速生长曲线的改变才是真正的可塑性。植物的表型可塑性、异速生长对于入侵植物的适应具有重要意义。干扰等异质性生境下表型可塑性成为物种生存扩散的有利性状,表型可塑性强的物种更有可能成为广布种。植物本身的异速生长特性或其异速生长曲线的改变都能影响其入侵能力。     

怒江河谷入侵植物与乡土植物丰富度的分布格局与影响因子

外来物种入侵严重威胁着乡土植物多样性并削弱了生态系统服务功能。本文基于滇西北怒江河谷植被调查的样方数据, 从群落水平研究了乡土和入侵植物多样性的空间分布格局, 以及地形、气候、人类干扰等因子对两种格局的影响。本研究共记录到外来入侵植物26种, 隶属于13科21属; 乡土植物1,145种, 分属于158科628属。沿着怒江河谷, 入侵植物物种丰富度随纬度与海拔的增加而减少; 乡土物种丰富度则随纬度增加而增加, 并在海拔梯度上呈单峰格局。运用广义线性模型分析公路边缘效应(反映生境干扰)、气候、地形和土壤等环境因素对物种丰富度分布格局的影响。等级方差分离的结果显示, 公路两侧的生境干扰对入侵种和乡土种的丰富度格局均具有首要影响。在自然环境因子中, 降水量是入侵植物丰富度的主要限制因子, 而乡土物种丰富度则主要受到地形因子尤其是坡向的影响。结构方程模型的分析结果也表明, 乡土植物和入侵植物丰富度之间的负相关关系反映了二者对环境响应的差异。本文结果支持物种入侵的资源可利用性限制假说, 并强调了人类活动对生物多样性的负面影响; 乡土植物或已较好地适应了干旱河谷气候, 但并没有显示出对外来物种入侵的抵抗作用。     

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。     

天然林和人工林外来入侵和本地植物对比研究: 以弄岗国家级自然保护区为例

外来植物的入侵严重威胁了自然保护区的生物多样性。为探讨自然保护区内天然林和人工林林下外来植物入侵状况及其与本地优势植物的种间关系, 本研究以弄岗国家级自然保护区为研究区域, 结合保护区内4种生境72个样方的野外调查数据, 利用生态位宽度、生态位重叠和种间联结指数比较了不同生境中外来入侵草本植物的多样性和生态位宽度以及外来入侵草本植物与本土优势草本植物的生态位重叠和种间联结。结果表明: 弄岗国家级自然保护区4种生境共记录6种外来入侵草本植物, 隶属于3科6属, 其中菊科物种数量最多。外来入侵植物多样性呈现出秋枫(Bischofia javanica)人工林 > 麻竹(Dendrocalamus latiflorus)次生林 > 人面子(Dracontomelon duperreanum)人工林 > 广西澄广花(Orophea polycarpa)次生林的特征。除广西澄广花次生林生境的外来入侵草本植物丰富度与本土草本植物丰富度呈显著正相关外, 其余生境均无显著的相关关系。三叶鬼针草(Bidens pilosa)的生态位宽度最大, 为4种生境均有分布的外来入侵植物。4种生境的外来入侵植物与本土优势植物生态位重叠和种间联结程度均较低, 种间竞争较弱。本研究明确了弄岗国家级自然保护区天然林和人工林中外来入侵草本植物的分布及其与本土草本植物的种间关系。     

外来植物入侵对生物多样性的影响及本地生物的进化响应

越来越多的证据表明,入侵植物能通过杂交和基冈渐渗等对本地种造成遗传侵蚀,甚至产生新的"基因型"来影响本地种的遗传多样性;通过生境片断化,改变本地种种群内和种群间的基因交流,造成近亲繁殖和遗传漂变,间接影响本地种的遗传多样性.另一方面,本地种能对入侵植物做出适应性进化响应,以减小或消除入侵植物的危害.本地种在与入侵植物的互作过程中产生了一系列的适应进化、物种形成以及灭绝事件,且这些事件不仅局限于地上生态系统,土壤牛物多样性同样受到影响,甚至也能发生进化响应.为更全面地了解外来植物入侵的生态后果和本地生物的适应潜力,本文综述了外来植物入侵对本地(地上和地下)生物(遗传)多样性的影响以及本地生物的进化响应.讨论了外来植物入侵导致的遗传和进化变化与其入侵性的关系,并提出了一些值得研究的课题.如土著种与外来种的协同进化、植物一土壤反馈调节途径和全球变化其他组分与生物入侵的关系等.     

A Comparison of Plastic Responses to Competition by Invasive and Non-invasive Congeners in the Commelinaceae

Evidence supporting an association between phenotypic plasticity and invasiveness across a range of plant taxa is based primarily on comparisons between invasive species and native species whose potential invasiveness is typically unknown. Comparison of invasive and non-invasive exotic species would provide a better test of whether plasticity promotes invasion. Such comparisons should distinguish between adaptive and non-adaptive plasticity because they have different consequences for invasiveness. Adaptive plasticity is expected to promote the invasion of multiple habitats, but non-adaptive plasticity may reflect specialization for invading more favorable habitats only. We grew four invasive and four non-invasive species of the Commelinaceae with and without competitors and compared their putatively adaptive plasticity of three traits related to competitive ability and non-adaptive plasticity in performance. The invasive species grew significantly more than the non-invasive species only in the non-competitive environment. The invasive species had greater plasticity of performance (total biomass) in response to competition than non-invasives, but there was no consistent difference in the plasticities of the traits related to competitive ability. These results are consistent with specialization of these invasive taxa for invading the more productive non-competitive environment rather than a superior ability to invade both competitive and non-competitive environments. A comprehensive understanding of the relationship between plasticity and invasiveness will require many more comparisons of the plasticity of invasive and non-invasive taxa in a range of traits in response to a variety of environments.     

Pre-adaptation or genetic shift after introduction in the invasive species Impatiens glandulifera?

Invasive exotic plants often grow fast, reproduce rapidly and display considerable phenotypic plasticity in their invasive range, which may be essential characteristics for successful invasion. However, it remains unclear whether these characteristics are already present in native populations (pre-adaptation hypothesis) or evolve after introduction (genetic shift hypothesis).To test these hypotheses we compared means and phenotypic plasticity of vegetative and reproductive traits between populations of Impatiens glandulifera collected from either the invasive (Norway) or native range (India). Seeds were sown and the resulting plants were exposed to different experimental environments in a glasshouse. We also tested whether trait means and reaction norms harbored genetic variation, as this may promote fitness in the novel environment.We did not find evidence that invasive populations of I. glandulifera grew more vigorously or produced more seeds than native populations. Phenotypic plasticity did not differ between the native and invasive range, except for the number of nodes which was more plastic in the invasive range. Genetic variation in the slope of reaction norms was absent, suggesting that the lack of change in phenotypic plasticity between native and invasive populations resulted from low genetic variation in phenotypic plasticity initially harbored by this species. Post-introduction evolution of traits thus probably did not boost the invasiveness of I. glandulifera. Instead, the species seems to be pre-adapted for invasion.We suggest that differences in habitat between the native and invasive range, more specifically the higher nutrient availability observed in the new environment, are the main factor driving the invasion of this species. Indeed, plants in the more nutrient-rich invasive range had greater seed mass, likely conferring a competitive advantage, while seed mass also responded strongly to nutrients in the glasshouse. Interactions between habitat productivity and herbivore defense may explain the lack of more vigorous growth in the new range.     

Invasive plants do not display greater phenotypic plasticity than their native or non‐invasive counterparts: a meta‐analysis

Phenotypic plasticity is commonly considered as a trait associated with invasiveness in alien plants because it may enhance the ability of plants to occupy a wide range of environments. Although the evidence of greater phenotypic plasticity in invasive plants is considerable, it is not yet conclusive. We used a meta‐analysis approach to evaluate whether invasive plant species show greater phenotypic plasticity than their native or non‐invasive counterparts. The outcome of such interspecific comparisons may be biased when phylogenetic relatedness is not taken into account. Consequently, species pairs belonged to the same genus, tribe or family. The meta‐analysis included 93 records from 35 studies reporting plastic responses to light, nutrients, water, CO₂, herbivory and support availability. Contrary to what is often assumed, overall, phenotypic plasticity was similar between invasive plants and native or non‐invasive closely related species. The same result was found when separate analyses were conducted for trait plasticity to nutrients, light and water availability. Thus, invasive plant species and their native or non‐invasive counterparts are equally capable of displaying functional responses to environmental heterogeneity. The colonization of a wide range of environments by invasive plants could be due to their capacity to undergo adaptive ecotypic differentiation rather than to their ability to display plastic responses. Alternatively, phenotypic plasticity might play a role in plant invasion, but only during the initial phases, when tolerance of the novel environment is essential for plant survival. Afterwards, once alien plants are identified as invaders, the magnitude of phenotypic plasticity might be reduced after selection of the optimum phenotypes in each habitat. The identification of plant traits that consistently predict invasiveness might be a futile task because different traits favor invasiveness in different environments. Approaches at the local scale, focusing on the ecology of specific invasive plants, could be more fruitful than global macro‐analyses.     

Local adaptation and phenotypic plasticity both occurred in <Emphasis Type="Italic">Wedelia trilobata</Emphasis> invasion across a tropical island

The role of the local adaptation and phenotypic plasticity of invasive species in their invasion of new environments has historically been a debatable issue, particularly at small spatial scales (e.g., different habitats within an island). We selected seven field sites across Hainan Island, Hainan Province, China, to investigate the role of local adaptation and/or phenotypic plasticity in the successful invasion of Wedelia trilobata by a field survey, molecular marker analysis, and common garden experiment. In the field survey, the clonal growth characteristics of W. trilobata showed significant differences among the seven sites, suggesting that the species was able to adapt to different environments. The mean phenotypic plasticity index of W. trilobata was higher than that of other invasive plant species (0.61 vs 0.48). The analysis of the inter-simple sequence repeat molecular markers of 420 individuals from the seven sites revealed a Shannon’s index that was similar to those of other invasive plants (0.29 vs 0.25). The nested analysis of the molecular variance in the genetic diversity of the population showed significant differences among the sites. In the common garden experiment, the growth characteristics of plants grown from the seven sites were significantly affected by light and density treatments but not by soil moisture. However, the responses of plants grown from different sites to light treatment varied. Plants from sunny sites had greater clonal traits than those from shady sites, indicating that local adaptation occurred in plant populations grown at some sites. Overall, our results implied that both phenotypic plasticity and local adaptation contributed to the successful invasion of W. trilobata across Hainan Island.     

Genetic and allelopathic differences between populations of daisy fleabane Erigeron annuus (L.) Pers. (Asteraceae) from disturbed and stable habitats

The interactions between invasive plants and their habitats may vary at different phases of the invasion process and depend on the phenotypic plasticity or local adaptations of each species. In this study, we investigated whether habitat changes during the invasion process are related to variations in the physiological traits (allelopathic properties) and genetic differentiation of daisy fleabane (Erigeron annuus (L.) Pers.). E. annuus is a winter annual invasive species that originated in North America and is now distributed throughout Europe. Genetic and genotypic diversity analyses were performed for 37 populations of E. annuus based on inter simple sequence repeat (ISSR) polymorphisms. In total, 684 plants were analyzed; 342 were from stable habitats and 342 were from disturbed habitats. The genetic differences among the populations from the different habitats were studied using a Bayesian cluster analysis and an analysis of molecular variance (AMOVA) and by calculating the genetic and genotypic diversity parameters. A germination test using the juglone index was employed to examine the potential allelopathic properties of the plants from the different habitats. Bayesian cluster analysis, AMOVA and allelopathic effects evaluation revealed differences in the allelopathic potential and genetic structure of the E. annuus populations from the disturbed and stable habitats. This differentiation of populations could be associated with founder effects or with different selection pressures among habitats.     

Invasion strategies in clonal aquatic plants: are phenotypic differences caused by phenotypic plasticity or local adaptation?

Background and Aims

The successful spread of invasive plants in new environments is often linked to multiple introductions and a diverse gene pool that facilitates local adaptation to variable environmental conditions. For clonal plants, however, phenotypic plasticity may be equally important. Here the primary adaptive strategy in three non-native, clonally reproducing macrophytes (Egeria densa, Elodea canadensis and Lagarosiphon major) in New Zealand freshwaters were examined and an attempt was made to link observed differences in plant morphology to local variation in habitat conditions.

Methods

Field populations with a large phenotypic variety were sampled in a range of lakes and streams with different chemical and physical properties. The phenotypic plasticity of the species before and after cultivation was studied in a common garden growth experiment, and the genetic diversity of these same populations was also quantified.

Key Results

For all three species, greater variation in plant characteristics was found before they were grown in standardized conditions. Moreover, field populations displayed remarkably little genetic variation and there was little interaction between habitat conditions and plant morphological characteristics.

Conclusions

The results indicate that at the current stage of spread into New Zealand, the primary adaptive strategy of these three invasive macrophytes is phenotypic plasticity. However, while limited, the possibility that genetic diversity between populations may facilitate ecotypic differentiation in the future cannot be excluded. These results thus indicate that invasive clonal aquatic plants adapt to new introduced areas by phenotypic plasticity. Inorganic carbon, nitrogen and phosphorous were important in controlling plant size of E. canadensis and L. major, but no other relationships between plant characteristics and habitat conditions were apparent. This implies that within-species differences in plant size can be explained by local nutrient conditions. All together this strongly suggests that invasive clonal aquatic plants adapt to a wide range of habitats in introduced areas by phenotypic plasticity rather than local adaptation.     

The Invasive Wetland Plant Alternanthera philoxeroides Shows a Higher Tolerance to Waterlogging than Its Native Congener Alternanthera sessilis

Plant invasion is one of the major threats to natural ecosystems. Phenotypic plasticity is considered to be important for promoting plant invasiveness. High tolerance of stress can also increase survival of invasive plants in adverse habitats. Limited growth and conservation of carbohydrate are considered to increase tolerance of flooding in plants. However, few studies have examined whether invasive species shows a higher phenotypic plasticity in response to waterlogging or a higher tolerance of waterlogging (lower plasticity) than native species. We conducted a greenhouse experiment to compare the growth and morphological and physiological responses to waterlogging of the invasive, clonal, wetland species Alternanthera philoxeroides with those of its co-occurring, native, congeneric, clonal species Alternanthera sessilis. Plants of A. philoxeroides and A. sessilis were subjected to three treatments (control, 0 and 60 cm waterlogging). Both A. philoxeroides and A. sessilis survived all treatments. Overall growth was lower in A. philoxeroides than in A. sessilis, but waterlogging negatively affected the growth of A. philoxeroides less strongly than that of A. sessilis. Alternanthera philoxeroides thus showed less sensitivity of growth traits (lower plasticity) and higher waterlogging tolerance. Moreover, the photosynthetic capacity of A. philoxeroides was higher than that of A. sessilis during waterlogging. Alternanthera philoxeroides also had higher total non-structural and non-soluble carbohydrate concentrations than A. sessilis at the end of treatments. Our results suggest that higher tolerance to waterlogging and higher photosynthetic capacity may partly explain the invasion success of A. philoxeroides in wetlands.