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。     

外来种入侵与物种多样性

在入侵生态学研究方面 ,物种多样性与生物入侵之间的关系已成为当前研究和争论的焦点。自Elton的经典假说提出以来 ,物种丰富度高的群落比物种贫乏的群落更能抵抗外来种入侵的观点得到广泛接受。一些理论模型和多样性处理实验支持了该假说。但现在越来越多的野外观测和实验研究开始对这一经典假说提出异议 ,甚至反对。同时 ,在入侵生态学广泛受到关注的今天 ,大量的实验研究也提出了一些新的观点。本文对Elton经典假说提出以来全球有关物种多样性与生物入侵关系的主要研究及其观点进行了评述 ,以期为我国有关研究工作的开展提供参考。     

我国外来水生生物入侵现状及其影响

生物入侵已经被公认是造成生物多样性丧失的重要因素之一，它对生态系统的结构和功能的严重影响已经引起全世界的高度重视。外来水生生物是生物入侵的重要组成部分：因此，从生物入侵的基本概念及入侵物种的传播途径着手，分析我国水生外来物种及其对水生生态系统和渔业可持续发展的影响。     

“主要入侵生物生态危害评估与防制修复技术示范研究”项目介绍

正随着国际交流的增加,物种扩散到新的地理区域更为便捷,一些外来物种在入侵地对当地的生态环境、经济、社会造成危害。《全球生物多样性展望》(2014)认为生物入侵是导致生物多样性丧失的第二大因素。我国是遭受生物入侵最严重的国家之一,已发现外来入侵生物560种,其中属于世界自然保护联盟(IUCN)公布的全球100种最具威胁的外来入侵生物的     

我国外来入侵生物防控现状、问题和对策

外来物种入侵已经威胁全球多个国家和地区,严重影响农林牧渔业生产,威胁生态系统稳定,是当前全球生物多样性丧失的主要原因之一,开展入侵物种防控已成为生物多样性保护与农业绿色发展的重点工作。我国已经成为全球遭受生物入侵威胁与损失最为严重的国家之一,截至2018年底,入侵我国的外来物种有近800种,已确认入侵农林生态系统的有638种,全国31个省（区、市）均有外来生物入侵发生并带来危害,半数以上县域都有入侵物种分布,几乎涉及所有类型的生态系统。本文对全球主要发达国家外来入侵物种发生情况及防控进行梳理发现,从国家层面立法开展外来入侵生物防控成为主流,制定长期防控战略并增加投入是入侵物种防控成功的关键。近些年,我国在外来入侵物种防控方面初步建立了工作机制,发布了重点管理外来入侵物种名录,完成了重点入侵物种的调查监测,开展了局部地区的防控措施,积极推动立法工作。根据国外生物入侵防治经验与我国实际情况,本研究提出我国外来入侵物种防控对策建议,包括加强法制建设、开展本底调查和启动重大防控工程等,为我国外来侵入生物的防控提供借鉴。     

四川攀西地区云南松群落物种多样性和谱系多样性对紫茎泽兰入侵的影响

外来植物入侵严重威胁着入侵地本土植物多样性和生态系统功能, 认识外来物种的入侵机制有助于提高对入侵植物的防控能力。本文以攀西地区云南松(Pinus yunnanensis)林下外来入侵植物紫茎泽兰(Ageratina adenophora)为研究对象, 基于大量野外群落调查, 从群落可入侵性入手, 分析了环境因子和群落物种多样性、谱系多样性等群落生态学特征对紫茎泽兰入侵的影响。结果表明: 海拔、坡向和火烧强度等环境因子和冠层郁闭度、灌木层盖度等生物因子对研究区紫茎泽兰入侵强度没有显著影响(P > 0.05); 但群落灌木层物种多样性和草本层组分种与紫茎泽兰的亲缘关系显著影响紫茎泽兰的入侵强度(P < 0.05), 说明灌木层对光照等环境资源的占用和草本层(同层)物种对相似资源的竞争能够在很大程度上抑制紫茎泽兰的入侵。     

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。     

群落组成及物种间相互作用对外来植物入侵的影响

外来植物入侵已成为严重的环境和社会问题,了解外来植物的入侵机制是有效控制其入侵的前提。生物阻抗假说认为,入侵地本地植物群落中的许多生物因子及生物过程能够抵御外来植物入侵。但关于群落抵抗外来种入侵的主要机制,目前还没有确定的结论。本文综述了群落中物种功能特征的多样性以及与外来种功能特征的相似度、植物与动物、植物与植物以及植物与土壤微生物间的相互作用等因素对外来植物入侵的影响,以及以前研究存在的不足。未来研究应该注重不同条件下植物与植物间的相互作用;不同竞争强度下,植物与食草动物的相互作用;植物、动物及土壤微生物三者之间的相互作用对外来植物入侵的影响。这些研究不仅能够丰富和完善入侵生态学理论,而且对于预测外来植物未来的扩散范围,合理有效地管理生态系统,防止外来植物入侵,保护本地生物多样性具有重要的实际意义。     

中国外来入侵生物的空间分布格局及其影响因素

不同地理区域影响生物分布格局的因子不同,对外来入侵物种也是如此。在区域尺度上分析外来入侵生物的空间分布格局及其影响因子对预测生物入侵的影响及入侵种的控制管理具有重要意义。本研究应用中国外来入侵物种数据库、自然环境数据库和社会人文环境数据库,分析了我国外来入侵动植物的空间格局;并运用主成分分析（PCA）和典范对应分析（CCA）探讨了自然环境和人类活动等因素对外来入侵动植物分布格局的影响;同时研究了外来入侵物种多样性与本地物种多样性之间的关系。结果表明,我国现有外来入侵动物138种、入侵植物384种,其数量和密度都呈现出由东南沿海向西北内陆减少的趋势,且入侵动物和入侵植物空间格局基本一致;降水（MAP）是决定我国外来入侵动植物分布格局的主要自然环境因子,国民生产总值（GDP）是主要社会经济影响因子。在全国尺度上,外来入侵物种多样性与本地物种多样性之间呈显著的正相关关系,但地域间存在较大差异。不同区域外来入侵物种与本土物种多样性的相关关系表现出不同,与研究尺度有密切关系。     

历史因素对土壤微生物群落与外来细菌入侵间关系的影响

群落的组成和结构如何影响其可入侵性一直是入侵生态学的研究热点。然而关于群落可入侵性和群落特征间关系的认知却很不统一。采用交叉互换的试验方法,首先将野外采集的两种长期不同施肥土壤(有机肥和化肥)进行灭菌并回接已方和对方的土壤悬液,研究土壤环境(历史非生物因素)和土壤微生物群落(历史生物因素)对重建土壤微生物群落特征的相对贡献。随后将用红色荧光蛋白标记的青枯菌作为外来种接入重建的土壤中,探究不同土壤微生物群落特征对外来细菌存活数量(前期入侵潜力)和存活时间(后期入侵潜力)的影响。结果表明,历史生物因素对重建土壤的原生动物数量、革兰氏阴性与阳性细菌比等群落特征和外来细菌的存活数量有影响;历史非生物因素对土壤微生物活性、细菌物种多样性和功能多样性等群落特征以及外来细菌入侵土壤后总的存活时间有影响;外来细菌入侵前期状况仅与原生动物数量、革兰氏阴性与阳性细菌比相关,而入侵后期的状况则仅与微生物活性、细菌物种多样性和功能多样性相关。总之,外来细菌在土壤中各时期的入侵潜力和土著微生物群落特征的相关性主要取决于二者是否由同种历史影响因素控制。本研究对于阐明生物群落结构与群落可入侵性之间关系,及指导土壤外来病原菌的防控均具有重要意义。     

Resource limitation,biodiversity, and competitive effects interact to determine the invasibility of rock pool microcosms

The success of species invasions depends on both the characteristics of the invaded habitat and the traits of the invasive species. At local scales biodiversity may act as a barrier to invasion; however, the mechanism by which biodiversity confers invasion resistance to a community has been the subject of considerable debate. The purpose of this study was to test the hypothesis that productivity and diversity affected the ability of a regionally available species to colonize communities from which it is absent. We hypothesized that the invasibility of rock pool invertebrate communities would increase with increasing nutrients and decrease with increasing diversity. We tested this possibility using naturally invaded outdoor aquatic microcosms. We demonstrated that the invasibility of an experimental multi-trophic aquatic community by a competitive native midge species (Ceratopogonidae: Dasyhelea sp.) was determined by an interaction between resource availability, diversity, and the densities of two competitive ostracods species. Nutrient enrichment increased invasion success; however, within nutrient-enriched microcosms, invasion success was highest in the low-diversity treatments. Our results suggest that resource availability may in fact be the principal mechanism determining invasibility at local scales in multi-trophic rock pool communities; however resource availability can be determined by both nutrient input as well as by the diversity of the biotic community.     

Restoration of invaded Cape Floristic Region riparian systems leads to a recovery in foliage-active arthropod alpha- and beta-diversity

The Cape Floristic Region of South Africa is a global biodiversity hotspot threatened by invasive alien plants (IAPs). We assessed the effect of plant invasions, and their subsequent clearing, on riparian arthropod diversity. Foliage-active arthropod communities were collected from two native and one invasive alien tree species. Alpha- and beta-diversity of their associated arthropod communities were compared between near pristine, Acacia-invaded and restored sites. Arthropod alpha-diversity at near pristine sites was higher than at restored sites, and was lowest at invaded sites. This was true for most arthropod taxonomic groups associated with all native tree species and suggests a general trend towards recovery in arthropod alpha-diversity after IAP removal. Overall, arthropod species turnover among sites was significantly influenced by plant invasions with communities at near pristine sites having higher turnover than those at restored and invaded sites. This pattern was not evident at the level of individual tree species. Although arthropod community composition was significantly influenced by plant invasions, only a few significant differences in arthropod community composition could be detected between restored and near pristine sites for all tree species and arthropod taxonomic groups. Assemblage composition on each tree species generally differed between sites with similar degrees of plant invasion indicating a strong turnover of arthropod communities across the landscape. Results further suggest that both arthropod alpha- and beta-diversity can recover after IAP removal, given sufficient time, but catchment signatures must be acknowledged when monitoring restoration recovery.     

Aerial arthropod communities of native and invaded forests, Robinson Crusoe Island, Chile

Invasive species significantly contribute to biological change and threaten biodiversity, with a growing body of evidence that plant invasions affect higher trophic levels. We explored the relative importance of plant invasion and forest structure on aerial arthropod abundance, diversity, and composition on Robinson Crusoe Island, Chile. We used flight intercept traps to sample aerial arthropods within distinct canopy strata of native and invaded forests over 3-mo periods in 2006 and 2007. Arthropod abundance and diversity were higher in native than invaded forest, and arthropod communities were distinct between forest types. In both forest types, arthropod abundance was highest in the lower canopy, and canopy strata exhibited some differences in arthropod community composition. Several morphospecies were distinctly associated with each forest type. The strong differences in aerial arthropod communities associated with the invasion of native forest by non-native plants may affect other trophic levels, such as insectivorous birds. Steps to stop invasive plant spread and to restore native forest composition and structure are needed to safeguard the integrity of native communities, from plants to higher-level consumers.     

Pine invasion decreases density and changes native tree communities in woodland Cerrado

ABSTRACT

Background: Invasive plants can negatively impact native communities, but the majority of the effects of these invasions have been demonstrated only for temperate ecosystems. Tropical ecosystems, including the Cerrado, a biodiversity hotspot, are known to be invaded by numerous non-native species, but studies of their impacts are largely lacking.

Aims: Our research aimed at quantifying how Pinus spp. presence and density affected Cerrado plant communities.

Methods: We sampled areas invaded and non-invaded by Pinus spp. to determine if pine invasion affected native tree richness, diversity, evenness, and density. We also evaluated if community composition differed between invaded and non-invaded sites.

Results: We found invaded plots had lower native tree densities than non-invaded plots and that Pinus spp. invasions changed native tree communities by reducing native species abundances.

Conclusion: Invasive pines had negative impacts on the native Cerrado tree community by reducing native plant density and changing species abundances. Reduced density and abundance at early invasion stages can result in reduction in biodiversity in the long term.     

紫茎泽兰入侵对土壤细菌的群落组成和多样性的影响

外来生物入侵可能对生物群落结构和生态系统功能产生多种影响, 但入侵植物与土壤微生物群落组成和多样性的关系尚不清楚。为了揭示外来植物紫茎泽兰(Eupatorium adenophorum)入侵对土壤化学性质和细菌群落组成及多样性的影响, 本研究利用第二代高通量测序技术, 比较了紫茎泽兰不同入侵程度的生境(本地植物群落、紫茎泽兰与本地植物混生群落、紫茎泽兰单优群落)土壤中细菌群落的差异。土壤化学性质分析表明, 土壤pH值、有机质、全N和全K随着紫茎泽兰的入侵而逐渐降低, 而土壤全P则在入侵程度最高的生境土壤中最高。通过测序共获得7,755个细菌OUT (operational taxonomic unit)。结果表明, 紫茎泽兰入侵对土壤的细菌多样性影响较小, ACE和Chao指数在3种不同生境间的差异不显著。细菌在紫茎泽兰与本地植物混生群落中的Shannon指数最低, 即细菌的多样性在中等入侵程度的生境最低。此外, 紫茎泽兰入侵改变了土壤细菌组成和结构, 酸杆菌门(Acidobacteria)和疣微菌门(Verrucomicrobia)的相对丰度, 从本地植物群落、混合群落到紫茎泽兰单优群落, 呈现出先增加后减少的趋势。可见, 紫茎泽兰入侵一定程度上改变了土壤微生物的多样性和群落结构, 并改变了土壤的化学性质。     

Resource availability and its role in development of invasion processes

The state of biotic communities inhabiting different water bodies is analyzed aiming to determination of conditions facilitating development of biological invasions. Mass-balance and dynamic models are used to reveal the factors ensuring the invasion of zebra mussel, Dreissena polymorpha, in Naroch Lakes (Byelorussia), and smelt, Osmerus eperlanus, in Lake Syamozero (Karelia). The results obtained indicate that for invasions leading to trophic chain lengthening, the main factor of successful invasion is the availability of resources that are not utilized by aboriginal species. Highest chances for successful invasion belong to invading species feeding on those trophic groups which contribute to the most extent to forming of 'excess' (i.e., available for the invader) production. It is shown that establishment of an alien species in a native community appears to be possible only when total amount of such 'excess' production is higher then some threshold value. Under conditions of spatio-temporal environmental heterogeneity, the timing factor, which determines the exact moment of an alien species intrusion, becomes important as well.     

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.     

Disentangling the effects of plant species invasion and urban development on arthropod community composition

Urban development and species invasion are two major global threats to biodiversity. These threats often co‐occur, as developed areas are more prone to species invasion. However, few empirical studies have tested if both factors affect biodiversity in similar ways. Here we study the individual and combined effects of urban development and plant invasion on the composition of arthropod communities. We assessed 36 paired invaded and non‐invaded sample plots, invaded by the plant Antigonon leptopus, with half of these pairs located in natural and the other half in developed land‐use types on the Caribbean island of St. Eustatius. We used several taxonomic and functional variables to describe community composition and diversity. Our results show that both urban development and A. leptopus invasion affected community composition, albeit in different ways. Development significantly increased species richness and exponential Shannon diversity, while invasion had no effect on these variables. However, invasion significantly increased arthropod abundance and caused biotic homogenization. Specifically, uninvaded arthropod communities were distinctly different in species composition between developed and natural sites, while they became undistinguishable after A. leptopus invasion. Moreover, functional variables were significantly affected by species invasion, but not by urban development. Invaded communities had higher community‐weighted mean body size and the feeding guild composition of invaded arthropod communities was characterized by the exceptional numbers of nectarivores, herbivores, and detritivores. With the exception of species richness and exponential Shannon diversity, invasion influenced four out of six response variables to a greater degree than urban development did. Hence, we can conclude that species invasion is not just a passenger of urban development but also a driver of change.     

Curvilinear Effects of Invasive Plants on Plant Diversity: Plant Community Invaded by Sphagneticola trilobata

The effects of invasive plants on the species diversity of plant communities are controversial, showing either a positive or negative linear relationship. Based on community data collected from forty 5 m×5 m plots invaded by Sphagneticola trilobata in eight cities across Hainan Island, China, we found S. trilobata decreased plant community diversity once its cover was beyond 10%. We demonstrated that the effects of invasive/native plants on the plant diversity of communities invaded by S. trilobata were curvilinear. These effects, which showed peaks under different degrees of vegetation cover, appeared not only for S. trilobata and all invasive plants, but also for all native plants. Invasive plants primarily had negative effects on plant diversity when they became abundant at a much lower cover level (less than 35%), compared with the native plants (over 60%). Thus, it is necessary to distinguish a range for assessing the effects of plants, especially invasive plants. Our results also confirmed that the invasion intensity of invasive alien plants increased with the intensity of local economic development. We highlight and further discuss the critical importance of curvilinear effects of biological invasion to provide ideas regarding the conservation of local biodiversity and the management of invasive plants.