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。     

群落可入侵性及环境胁迫

群落的可入侵性研究是入侵生态学的中心内容。从群落进化历史,群落结构,繁殖体压力。干扰和胁迫等5个影响入侵的主要因素综述了近年来群落可入侵性研究的工作进展和成果。其中重点阐述了群落结构对入侵的影响。包括群落物种组成,物种多样性和物种功能群多样性的诸多实验研究和相关假说,并用理论模型结合具体实验研究探讨了环境胁迫和可入侵性的关系。针对目前群落可入侵性研究存在的问题提出了自己的见解。     

生物入侵机制研究进展

本文从外来种本身的生物学特性,外来种与新栖息地土著种之间的相互作用,群落生物多样性对入侵种的抵抗能力,新栖息地生态环境变化对入侵种的影响等方面综述了近年来有关生物入侵机制的研究进展,并探讨了研究方法的发展和亟待解决的问题,大量的研究表明生物入侵机制是极为多样的,可能没有一般的,通用的机制,阐明生物入侵的机制,对推动生态学理论的发展和控制有害外来种有重要意义。     

植物群落的生物多样性及其可入侵性关系的实验研究

 生物入侵已经成为一个普遍性的环境问题,并为许多学者所关注。尽管一些理论研究和观察表明生物多样性丰富的群落不容易受到外来种的入侵,但后来有些实验研究并没能证实两者的负相关性,多样性 可入侵性假说仍然是入侵生态学领域争论比较多的一个焦点。人为构建不同物种多样性和物种功能群多样性（C3 禾本科植物、C4植物、非禾本科草本植物和豆科植物）梯度的小尺度群落,把其它影响可入侵性的外在因子和多样性效应隔离开来,研究入侵种喜旱莲子草(Alternanthera philoxeroides)在不同群落里的入侵过程来验证多样性 可入侵性及其相关假说。研究结果显示,物种功能群丰富的群落可入侵程度较低,功能群数目相同而物种多样性不同的群落可入侵性没有显著性差异,功能群特征不同的群落也表现出可入侵性的差异,生活史周期短的单一物种群落和有着生物固氮功能的豆科植物群落可入侵程度较高,与喜旱莲子草属于同一功能群且有着相似生态位的土著种莲子草(A. sessilis)对入侵的抵抗力最强。实验结果表明,物种多样性和群落可入侵性并没有很显著的负相关,而是与物种特性基础上的物种功能群多样性呈负相关,群落中留给入侵种生态位的机会很可能是决定群落可入侵性的一个关键因子。     

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。     

草地物种多样性对群落可入侵性影响的研究进展

综述草地群落入侵实验中物种多样性和群落可入侵性关系的研究进展。目前物种多样性与群落可入侵性主要出现了对立的关系模式,被普遍接受的解释机制为尺度依赖。但其它研究中出现了更为复杂的关系,提出物种特性、植物更新、种间关系变化和群落构建机制等其它因素可能是导致物种多样性与群落可入侵性出现复杂关系的原因。建议未来研究中应注意的几个问题,即物种多样性与群落可入侵性关系在不同营养级适用性,与群落构建机制变化间的联系和时间尺度对物种多样性与群落可入侵性关系的影响。     

Species turnover drives grassland community to phylogenetic clustering over long-term grazing disturbance

放牧通过改变草地群落物种组成和生物多样性,进而影响草地群落结构,对草地生态系统服务和功能产生深远的影响。然而,有关系统发育多样性和系统发育群落结构对长期放牧干扰的响应和适应的研究仍然很少,尤其是对于分布在极端环境中的生态系统。我们在青藏高原高寒草地上开展了多放牧强度的试验,探讨放牧干扰对植物系统发育多样性和群落结构的影响。研究发现,放牧干扰增加了植物群落的物种丰富度,促进了群落物种周转,从而改变了群落物种组成。低强度放牧对系统发育多样性和群落结构没有显著影响,而高强度放牧促使群落结构由分散向聚集变化。高强度放牧通过强烈的环境过滤作用,选择了一些耐牧的草地植物物种。在高强度放牧条件下,草地群落的聚集结构由近缘种的入侵和远缘种的丢失共同驱动。在植物功能性状水平上,我们发现与低强度放牧相比,高强度放牧通过改变根系深度对物种入侵产生影响,在很大程度上提升了物种的入侵性。我们的研究强调,仅仅利用物种丰富度和多样性并不能全面反映放牧干扰对草地群落的影响,而且在以后的放牧生态学研究中应该更加关注物种周转对群落系统发育多样性和群落结构的影响。     

外来生物入侵中的达尔文归化谜团

针对外来生物入侵问题,达尔文在《物种起源》中提出了相反的两个假说,分别主张和本地物种亲缘关系近和远的外来物种更容易归化和入侵.这两个假说综合在一起现在被称为达尔文归化谜团.在沉寂了百余年后,随着系统发育和功能性状分析在生态学中的广泛运用,该谜团在近30年内迅速成为生物入侵领域研究的热点和前沿问题.但由于不同的研究往往得出矛盾的结论,错综复杂的研究结果使得该谜团的和解仍面临很大困难.本文梳理了达尔文归化谜团的内涵和外延,回顾了该谜团研究的发展历程和脉络,总结了近年来从空间尺度、入侵阶段、差异度量方式和控制性实验等角度尝试和解这一谜团的探索及其进展.本文认为,充分考虑环境梯度和人类活动的影响,综合分析入侵进程中的本地群落特征和动态,选择恰当的统计方法和零模型,并将该谜团的研究思路和理念引入到疾病生态学等诸多相关领域,是未来达尔文归化谜团研究的重点方向.对这一长达160余年谜团的探索,不仅有助于发展和完善入侵生态学理论,也将为外来生物入侵的防控提供重要科学指导.     

外来种入侵的过程、机理和预测

生物入侵是指某种生物从原来的分布区域扩展到一个新的（通常也是遥远的）地区,在新的区域里,其后代可以繁殖、扩散并持续维持下去,生物入侵成功的原因,即与入侵者本身的生物学,生态学特征有关,也与群落的脆弱性有关,入侵者可能较本地种的竞争能力强,更适应当地的环境,有的入侵者还可以改变环境,使之对已有利,而不利于本地种。缺乏天敌制约。群落的稳定性低和异常的环境扰动往往导致生物入侵,生物入侵的预测包括哪一种外来种会变成入侵种？哪些生态系统区域会被入侵？影响程度如何？入侵种的扩散态势如何等内容,对有关的理论和模型作了评介。     

基于现代物种共存理论的入侵生态学概念框架

在入侵生态学60多年的发展历程中, 生态学家提出了多种多样的假说来解释生物入侵的机制。这些纷繁复杂的假说在丰富我们对生物入侵认知的同时, 也给入侵生态学概念的整合带来了困难。其中, 外来种和土著种是否存在生态学差异, 以及这种差异如何影响生物入侵, 是入侵生态学研究和争论的焦点问题。现代物种共存理论通过将外来种和土著种的生态学差异划分为生态位差异和适合度差异, 为入侵生态学概念的整合提供了新的视角。依据该理论, 外来种可以通过两种策略实现成功入侵: 一是扩大与土著种的生态位差异, 二是提高自身相较于土著种的适合度优势。因此, 外来种-土著种的生态位差异和适合度差异共同决定了入侵的成败与危害程度。通过对经典入侵假说进行梳理, 我们发现大部分假说都可以在该理论框架下进行解读, 不同假说的主要差别在于强调不同生态学过程对生态位和适合度差异的影响。同时, 这一理论框架很好地解释了为什么外来种-土著种的亲缘关系和性状差异会对生物入侵产生复杂的影响, 为达尔文归化谜团的和解以及外来种-土著种功能性状的比较研究提供了新的思路。目前, 现代物种共存理论还处于快速发展的阶段, 依旧存在很多不足, 但将其运用到生物入侵的实证研究中将是入侵生态学今后一个重要的发展方向。     

Plant diversity increases resistance to invasion in the absence of covarying extrinsic factors

Biological invasion is a widespread, but poorly understood phenomenon. Elton's hypothesis, supported by theory, experiment, and anecdotal evidence, suggests that an important determinant of invasion success is resident biodiversity, arguing that high diversity increases the competitive environment of communities and makes them more difficult to invade. Observational studies of plant invasions, however, find little support for this hypothesis and argue strongly against it. Lack of control of extrinsic factors (e.g., disturbance, climate, or soil fertility) that covary with biodiversity and invasion in observational studies makes it difficult to determine if their findings truly refute Elton's hypothesis. We examined performance of Crepis tectorum (an invasive, annual composite weed) in experimental prairie grassland plots and greenhouse plant assemblages in which resident species richness was directly manipulated. Under these conditions, unlike observational studies, no covarying extrinsic factors could interfere with interpreting results. We found a strong inverse association between resident diversity and invader performance as predicted by Elton's hypothesis. Higher resident diversity increased crowding, decreased available light, and decreased available nutrients all of which increased the competitive environment of diverse plant assemblages and reduced C. tectorum success. Examination of individual resident species impacts on C. tectorum performance demonstrated that this diversity effect was not due to the sampling effect. These results suggest that both Elton's hypothesis and its competitive mechanism may operate in nature, but covarying extrinsic factors may obscure the negative impact of diversity on invader success.     

Forest Invasibility in Communities in Southeastern New York

While biological invasions have been the subject of considerable attention both historically and recently, the factors controlling the susceptibility of communities to plant invasions remain controversial. We surveyed 44 sites in southeastern New York State to examine the relationships between plant community characteristics, soil characteristics, and nonnative plant invasion. Soil nitrogen mineralization and nitrification rates were strongly related to the degree of site invasion (F= 30.2, P < 0.0001 and F= 11.8, P < 0.005, respectively), and leaf C : N ratios were negatively correlated with invasion (R ²= 0.22, P < 0.0001). More surprisingly, there was a strong positive relationship between soil calcium levels and the degree of site invasion (partial r= 0.70, P < 0.01), and there were also positive relationships between invasion and soil magnesium and phosphorus. We found, in addition, a positive factor-ceiling relationship between native species diversity and invasive species diversity. This positive relationship between native and invasive diversity contradicts earlier hypotheses concerning the relationships between species diversity and invasion, but supports some recent findings. Cluster analysis distinguished two broad forest community types at our sites: pine barrens and mixed hardwood communities. Invaders were significantly more abundant in mixed hardwood than in pine barrens communities (Mann–Whitney U = 682.5, P < 0.0001). Even when evaluating the mixed hardwood communities alone, invasion remained significantly positively correlated with soil fertility (calcium, magnesium, and net nitrogen mineralization rates). Soil texture and pH were not useful predictors of the degree to which forests were invaded. Nitrogen and calcium are critical components of plant development, and species better able to take advantage of increased nutrient availability may out-perform others at sites with higher nutrient levels. These results have implications for areas such as the eastern United States, where anthropogenic changes in the availability of nitrogen and calcium are affecting many plant communities.     

Giant invasive Heracleum persicum: Friend or foe of plant diversity?

The impact of invasion on diversity varies widely and remains elusive. Despite the considerable attempts to understand mechanisms of biological invasion, it is largely unknown whether some communities’ characteristics promote biological invasion, or whether some inherent characteristics of invaders enable them to invade other communities. Our aims were to assess the impact of one of the massive plant invaders of Scandinavia on vascular plant species diversity, disentangle attributes of invasible and noninvasible communities, and evaluate the relationship between invasibility and genetic diversity of a dominant invader. We studied 56 pairs of Heracleum persicum Desf. ex Fisch.‐invaded and noninvaded plots from 12 locations in northern Norway. There was lower native cover, evenness, taxonomic diversity, native biomass, and species richness in the invaded plots than in the noninvaded plots. The invaded plots had nearly two native species fewer than the noninvaded plots on average. Within the invaded plots, cover of H. persicum had a strong negative effect on the native cover, evenness, and native biomass, and a positive association with the height of the native plants. Plant communities containing only native species appeared more invasible than those that included exotic species, particularly H. persicum. Genetic diversity of H. persicum was positively correlated with invasibility but not with community diversity. The invasion of a plant community by H. persicum exerts consistent negative pressure on vascular plant diversity. The lack of positive correlation between impacts and genetic diversity of H. persicum indicates that even a small founder population may cause high impact. We highlight community stability or saturation as an important determinant of invasibility. While the invasion by H. persicum may decrease susceptibility of a plant community to further invasion, it severely reduces the abundance of native species and makes them more vulnerable to competitive exclusion.     

Community diversity and invasion resistance: An experimental test in a grassland ecosystem and a review of comparable studies

The relationship between community diversity and invasion resistance in a grassland was examined using experimental plant assemblages that varied in species richness and composition. The assemblages were weeded for three seasons to remove unsown species and we used the number of weeded seedlings, their total biomass and the number of species removed as indicators of community resistance and susceptibility to invasion. In general, we found a positive relationship between invasion resistance and increasing community diversity. Similar patterns of establishment were observed at the end of the fourth field season after several months without weeding. Increased invasion resistance with higher diversity appears to come through reduced levels of several above- and belowground resources, although these did not fully explain the effects of species richness in the studys analyses. Experimental increases and reductions of litter biomass within the studys experimental plant assemblages did not modify these patterns significantly. A review of comparable studies of invasion across directly manipulated diversity gradients revealed similar patterns. Positive effects of species diversity on invasion resistance were found in experimental manipulations of plant diversity conducted in the field and in the glasshouse, from studies with aquatic microcosms and in a marine system. Although some exceptions to this pattern were found in both terrestrial plant systems and aquatic microcosms, it was concluded that in biodiversity manipulation experiments more diverse communities are generally more resistant to invasion.     

Biological invasions and the neutral theory

Aim  The invasion of natural communities by alien species represents a serious threat, but creates opportunities to learn about community functions. Neutral theory proposes that the niche concept may not be needed to explain the assemblage and diversity of natural communities, challenging the classical view of community ecology and generating a lasting debate. Biological invasions, when considered as natural experiments, can be used to contrast some of the predictions of neutral and classic niche theories.
Location  Global.
Methods  We use data from biological invasions as natural experiments to contrast some of the fundamental predictions of neutral theory.
Results  Some emerging patterns did not differ from neutral model expectations (e.g. the relationship between native and exotic species richness, invasibility of resource-rich habitats, and the relationship between propagule release and invasion success). Nevertheless, other patterns (e.g. experimental evidence of the relationship between diversity and susceptibility to invasion, the invasion of communities with a low resource availability, invasiveness related to species traits) contrasted with the predictions that can be inferred from neutral theory.
Main conclusions  Neutral theory correctly highlights the need to include randomness in models of community structure. Biological invasion patterns show that neutral forces are important in structuring natural communities, but the patterns differ from those inferred from a complete neutral model. For biodiversity-conservation purposes, the implications of accepting or not accepting neutral theory as a process-based theory are very important.     

Making remote sense of biodiversity: What grassland characteristics make spectral diversity a good proxy for taxonomic diversity?

Aim

The spectral variability hypothesis (SVH) predicts that spectral diversity, defined as the variability of radiation reflected from vegetation, increases with biodiversity. While confirmation of this hypothesis would pave the path for use of remote sensing to monitor biodiversity, support in herbaceous ecosystems is mixed. Methodological aspects related to scale have been the predominant explanation for the mixed support, yet biological characteristics that vary among herbaceous systems may also affect the strength of the relationship. Therefore, we examined the influence of three biological characteristics on the relationship between spectral and taxonomic diversity: vegetation density, spatial species turnover and invasion by non-native species. We aimed to understand when and why spectral diversity may serve as an indicator of taxonomic diversity and be useful for monitoring.

Location

Continental U.S.A.

Time Period

Peak greenness in 2017.

Major Taxa Studied

Grassland and herbaceous ecosystems.

Methods

For nine herbaceous sites in the National Ecological Observatory Network, we calculated taxonomic diversity from field surveys of 20 m × 20 m plots and derived spectral diversity for those same plots from airborne hyperspectral imagery with a spatial resolution of 1 m. The strength of the taxonomic diversity–spectral diversity relationship at each site was subsequently assessed against measurements of vegetation density, spatial species turnover and invasion.

Results

We found a significant relationship between taxonomic and spectral diversity at some, but not all, sites. Spectral diversity was more strongly related to taxonomic diversity in sites with high species turnover and low invasion, but vegetation density had no effect on the relationship.

Main Conclusions

Using spectral diversity as a proxy for taxonomic diversity in grasslands is possible in some circumstances but should not just be assumed based on the SVH. It is important to understand the biological characteristics of a community prior to considering spectral diversity to monitor taxonomic diversity.     

吉林灌木群落物种多样性与气候和局域环境因子的关系

为了研究气候和局域环境因子对物种多样性的相对作用大小,以及验证两种均匀度地理格局的假说在半湿润地区次生灌丛的适用性,对吉林东、南部地区的灌木群落进行了研究。共调查森林破坏后形成的次生灌丛样方45个,结合气候数据和局域环境因子数据,研究了气候、局域环境因子对群落、灌木层、草本层的物种丰富度、均匀度的影响,以及对不同水分生态型(旱生、旱中生、湿中生)灌木影响的差异。结果表明:1)吉林次生灌丛的群落、草本层物种丰富度,以及草本层均匀度,随纬度增加而显著上升。2)对物种多样性和气候、局域环境因子的分析表明,群落、草本层物种数主要受局域环境因子而不是气候的影响;其物种丰富度与纬度的反常关系,是由于灌木层盖度随降水增加而上升,从而导致物种数下降。灌木层物种数与纬度、气候因子的相关性不显著,则是由于不同水分生态型对气候梯度的响应不一致,反映出功能群对多样性格局的影响。3)群落、灌木层均匀度主要受气候因子的影响;而草本层均匀度主要受局域环境因子的影响,降水同样通过对灌木层盖度的影响间接作用于草本均匀度。但群落、灌木和草本层的结果,都支持均匀度随着环境条件改善而增加的假说,而不支持随着生产力增加、竞争加剧,从而导致均匀度下降的假说。结果表明,物种丰富度和均匀度的影响机制存在很大差异,但二者都受到局域环境因子的强烈影响。气候通过局域生物因素(如盖度、生活型)间接作用于多样性格局,是气候对多样性影响的一个重要方面,但尚未得到应有的重视。由于局域生物因素也随气候而变化,仅研究多样性和气候的表面关系,将无法准确预测气候变化对多样性的影响。     

Ecological explanations for successful invasion of exotic plants

The intentional introduction of exotic species can increase the level of local biodiversity, enrich people's material lives, and bring significant social and economic benefits that are also the symbols of human progress. However, along with the frequent intercourse among countries and regions, the frequency of uncontrolled crossregional migration of species is increased and there is a lack of scientific management strategy for the intentional introduction of exotic species. Exotic species invasion, which is behind habitat fragmentation, has become the second largest threatening factor to the maintenance of the global-scale level of biological diversity. Exotic species invasion can destroy the structure of an ecosystem, disturb the economic life of a society, and do harm to human health. In this paper, the authors review some of the ecological explanations for issues such as "what causes or mechanisms have led to the successful invasion of exotic species", including the "ideal weeds characteristics", "biodiversity resistance hypothesis", "enemies release hypothesis", "evolution of increased competitive ability hypothesis", "niche opportunity hypothesis", and "novel weapon hypothesis". The authors also analyze and evaluate the background and theoretical basis of the hypotheses, providing explanations for some phenomena, as well as the deficiencies of these explanations.