网络理论在入侵生物学中的应用研究现状

生物入侵是一个动态有序的过程,其发生和危害存在异质性,通常由来源地、入侵地和它们之间的连接构成的系统中的自然、生物、社会等因素所决定。网络理论是研究复杂系统的一种新方法,本质是从复杂的信息中抽象出规律、揭示系统的结构特征共性。近20年,网络理论已被应用于生物入侵研究。本研究综述了网络理论在生物入侵研究中的应用进展,明确了主要的研究方向和前沿热点,认为:2000年以来国际上已开展的研究集中在评估外来物种入侵风险和入侵后对生态系统影响2个方面;外来物种随运输网络入侵的风险评估和景观连接性对入侵物种扩散的影响、外来物种入侵对本地物种间互作网络的影响及生态群落可入侵性是网络理论应用的热点;研究热点具有明显的时间发展特征,2013年以前多是对生态系统的影响,近10年来主要是风险评估。我国利用网络理论研究外来物种入侵较少且集中于对生态系统的危害,未来应加强对外来物种的时空定量传入和扩散风险评估,为我国制定和提升外来入侵物种早期监测预警、阻止新的入侵、抑制进一步扩散的管理措施提供依据。     

海洋外来物种入侵生态学研究

海洋外来物种入侵已成为最为严重的全球性环境问题之一。海洋生态系统类型多样、环境复杂,其生物入侵的监测、控制与管理难度相对较大。我国对陆地外来生物的入侵已开展了较为深入的研究,但对于海洋外来生物的入侵研究仍处于起步阶段,对其入侵监测、入侵机制、入侵危害的程度以及防治等问题缺乏基础数据。本文在分析国内外海洋外来生物入侵现状的基础上,概述其入侵生态学研究形势及相关成果,包括海洋外来物种的入侵途径、入侵过程、入侵生态效应以及全球变化对入侵的影响等。海洋外来生物的入侵可能对海洋生态系统造成直接或间接的影响,如种间竞争破坏生态环境、与土著种杂交造成遗传污染、病原生物及有毒藻类导致海洋生态灾害加剧等。此外,从政策和法规、入侵风险评估、监测和公共宣传教育、生物信息系统和有效管理机制等方面提出对我国海洋外来物种入侵的防治策略。本研究为我国海洋外来物种的进一步研究提供了参考。     

中国公布的3 批外来入侵植物种类特征与入侵现状分析

生物入侵是当今世界各国发展的一大挑战。外来入侵物种已对各国经济造成了不同程度的影响, 中国也未能幸免。为了防止外来物种入侵, 减轻其对我国造成的影响, 国家环保总局先后于2003 年、2010 年及2014 年公布了3 批中国外来入侵物种名单。据统计, 中国公布的3 批外来物种名单中的外来入侵植物共29 种, 占半数以上, 其中菊科和禾本科植物合计20 种, 约占3 批外来入侵植物总种数的68.96%; 生活型为草本的植物最多, 占96.55%; 原产地为美洲的占93.10%。3 批外来入侵植物中62.07%的植物是非有意引入的, 现今已全部建立种群, 且在全国各地大范围扩散。     

微生物入侵影响因素研究进展

探明生物入侵的影响因素能够有效防治生物入侵,减少其对生态系统的破坏,但现有的入侵研究主要集中于动植物入侵,对微生物入侵关注较少。本文综述了外来微生物自身属性、入侵地生物和非生物状况以及外来种和土著种之间差异等对微生物入侵的影响,比较了微生物入侵与动植物入侵、微生物定殖之间的差别,提出了今后需要进一步加强的研究内容。外来微生物入侵受其自身种系、形态、大小等特性,入侵地生物多样性、天敌、有效资源等生物或非生物因子,以及外来-本地种谱系距离、生态位差异、相对适应性差异等三方面因素共同影响,但不同因素之间的关联度、交互性及相对贡献仍不清楚。外来微生物入侵过程与动植物入侵过程、微生物定殖过程有诸多类似之处,但同时也存在不少差异。今后,需加强研究外来微生物独特性状(如:持留状态和群体感应等)对其入侵影响,关注微生物进入新环境的存活数量、扩散范围和影响程度,加强利用定殖研究中常用的微生物作为入侵物种验证经典入侵理论,以及注重观察全球变化下外来微生物的入侵趋势和区分微生物在不同入侵阶段的主要影响因素。     

ISSR分子标记在入侵植物研究中的应用

外来生物入侵是对全球生物多样性最为严重的威胁之一,对经济安全、生态安全、社会安全、国际利益和国际贸易都具有重要影响.入侵种群的分子标记分析是外来入侵生物研究的重要途径.其中,简单序列重复区间标记(inter-simple sequence repeat,ISSR)是一种基于微卫星序列发展起来的新型分子标记,具有简便、快捷、结果稳定和DNA多态性高等优点.本文系统地介绍了ISSR分子标记的原理、技术特征及其实验操作,并简要地阐述了ISSR分子标记在外来入侵植物的群体遗传结构分析、遗传多样性检测、入侵来源推测、入侵植物的分布模式及其亲缘关系分析、入侵植物的繁育特性检测等方面的应用及其研究进展.     

中国入侵生物学学科的构建与发展

自英国生态学家查尔斯.艾尔顿1958年撰写的《动植物的入侵生态学》出版至今,半个世纪已经过去,这部著作被公认为是生物入侵在科学研究方面的开端。这期间生物入侵研究经历了萌芽期（20世纪80年代之前）、成长期（20世纪80年代）和快速发展期（20世纪90年代末期至今）。在这个过程中,越来越多的概念、假说、方法和技术被提出和整合到生物入侵研究之中,由此催生了一门生态学领域的新兴学科——入侵生物学。本文在对近50年来生物入侵专著和论文的统计分析基础上,介绍了国际生物入侵研究的发展脉络和现状。同时综述了中国入侵生物学在基础和应用研究方面的相关进展,着重阐述了主要科学问题（＂入侵潜力与成功入侵的关系＂、＂入侵种种群的扩张与扩散＂、＂入侵种的生态适应性与进化＂及＂本地生态系统对入侵的响应及可入侵性＂）和预防与控制的技术体系（风险评估与早期预警、检测与监测、狙击与灭除、生物防治、生态修复与干扰调控）,并进一步介绍了中国入侵生物学学科体系的构建和框架,提出入侵生物学是研究外来物种的入侵性与生态系统的可入侵性,以及外来物种预防与控制的科学,是一门多领域交叉的学科。最后展望和讨论了中国入侵生物学学科发展可能遇到的一些问题。     

外来养殖鱼类的入侵风险评估及防控对策——以尼罗罗非鱼为例

[目的]筛选合适的指标建立一套外来养殖鱼类的生物入侵风险评估体系,并对外来养殖鱼类的生物入侵防控提出对策建议。[方法]通过文献资料的收集和整理,对外来养殖鱼类中典型入侵物种的入侵过程、影响危害和入侵生物学特性进行分析和归纳,从适应能力、繁殖能力、扩散能力3方面指示其入侵性;从对生物的影响和对环境的影响2方面指示其生态影响;从自然因素和人为因素2方面指示环境可入侵性,以上述3方面为框架进行评估体系构建。[结果]筛选20个指标构建了外来养殖鱼类的生物入侵风险评估体系,并举例说明该评估体系的应用。从法规政策、科学研究、治理技术和公众参与等4个方面针对性地提出外来养殖鱼类生物入侵风险防控对策。[结论]防范和治理入侵生物是一个系统工程。对于外来养殖鱼类的管理,既不能只考虑经济效益而置生态风险于不顾,也不能片面放大外来养殖良种的入侵风险。科学管控的关键在于完善制度建设、加强风险评估、发展防治手段、促进公众参与,使外来鱼类养殖业在严格受控的前提下发挥其经济效益,将其潜在的生态危害效应降到最低。     

重要外来入侵植物随南水北调工程传入京津冀受水区的风险评估

【目的】外来物种在入侵地的扩张蔓延是其造成危害的关键。南水北调工程可为已入侵外来物种的进一步扩张蔓延提供通道和驱动力。风险评估可为预防入侵植物通过该途径的扩散蔓延提供依据。【方法】以国家重点管理外来入侵物种名录中的入侵植物为研究对象,根据生物入侵的发生过程(传入、定殖、扩散、危害)构建了包括4个层次29个指标的外来物种入侵综合风险评估的指标体系,规范了风险指数的计算方法,系统评估了24种入侵植物沿南水北调输水通道向北扩散而入侵京津冀受水区的风险。【结果】紫茎泽兰和互花米草的入侵风险值(R)相对较低(R0.6),其余22种入侵植物的风险值均较高(R0.6),其中,喜旱莲子草、凤眼莲和大薸3种外来水生植物的入侵风险等级最高(R0.8)。【结论】南水北调工程可能会促进入侵植物向北扩张蔓延并最终入侵京津冀受水区。建议尽早开展监测预警工作,以控制入侵物种随南水北调工程的扩张蔓延,进而阻止或降低其对京津冀尤其是华北最大水源地——白洋淀和雄安新区生态环境的威胁。     

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。     

植物入侵机制研究进展

生物入侵对人类的生产、生活及生态系统的稳定性造成了严重的影响。本文主要介绍植物入侵相关假说、入侵过程中的进化改变及其对生物入侵防控的意义。     

Molecular genetic approaches to elucidate the ecological and evolutionary issues associated with biological invasions

Biological invasions may cause serious damage to the native environments and threaten the native biodiversity. Molecular genetic approaches have been found to be powerful tools for investigating the ecological and evolutionary aspects of biological invasions because the genetic structure and level of genetic variation of an invasive species are changed following its invasion. The present article reviews the use of molecular markers in addressing various aspects of invasive species. The application of these techniques has shown that many invasive species are actually "cryptic" species – species whose uniqueness is only recognizable at the genetic level. An estimation of the actual number of invasive species is essential when evaluating its ecological and economic impacts. Molecular genetic approaches have also enabled the source populations of invasive species to be identified. Reconstructions of invasion histories are crucial to preventing future invasions and conserving the native biodiversity, while comparisons of genetic variations between the native and introduced populations provide valuable opportunities to elucidate the mechanisms of rapid adaptation demonstrated by many invasive species.     

分子标记技术在昆虫入侵研究中的应用

遗传变异与种群持续性及其进化潜力密切相关,而生物入侵导致种群遗传变异或遗传多样性的改变为研究自然界中各种生态和进化问题提供了理想模式。分子标记技术是调查种群遗传变异的重要工具,揭示了入侵种的入侵过程和结果,并预测未来的发生情况。本综述归纳了分子标记技术在昆虫入侵机制研究中的应用,以典型的研究个案为例,分别综述了分子标记技术在隐蔽入侵的监测应用,分子标记技术在重构入侵历史研究中的推算方式,分子标记技术在探索种群遗传变异与成功入侵机制方面取得的重要进展,并进一步介绍了高分辨率熔解曲线（high-resolution melting, HRM）分析在昆虫入侵研究中的应用前景。     

Deciphering the worldwide invasion of the Asian long‐horned beetle: A recurrent invasion process from the native area together with a bridgehead effect

Retracing introduction routes is crucial for understanding the evolutionary processes involved in an invasion, as well as for highlighting the invasion history of a species at the global scale. The Asian long‐horned beetle (ALB) Anoplophora glabripennis is a xylophagous pest native to Asia and invasive in North America and Europe. It is responsible for severe losses of urban trees, in both its native and invaded ranges. Based on historical and genetic data, several hypotheses have been formulated concerning its invasion history, including the possibility of multiple introductions from the native zone and secondary dispersal within the invaded areas, but none have been formally tested. In this study, we characterized the genetic structure of ALB in both its native and invaded ranges using microsatellites. In order to test different invasion scenarios, we used an approximate Bayesian “random forest” algorithm together with traditional population genetics approaches. The strong population differentiation observed in the native area was not geographically structured, suggesting complex migration events that were probably human‐mediated. Both native and invasive populations had low genetic diversity, but this characteristic did not prevent the success of the ALB invasions. Our results highlight the complexity of invasion pathways for insect pests. Specifically, our findings indicate that invasive species might be repeatedly introduced from their native range, and they emphasize the importance of multiple, human‐mediated introductions in successful invasions. Finally, our results demonstrate that invasive species can spread across continents following a bridgehead path, in which an invasive population may have acted as a source for another invasion.     

淡水鱼类入侵种的分布、入侵途径、机制与后果

生物入侵已经成为全球面临的三大环境问题之一。鱼类入侵现象也随全球经济一体化的进程日益严重。本文综述了全球淡水鱼类入侵的现状和研究进展, 包括鱼类入侵的定义及分布、入侵途径和机制、产生的生态和社会经济影响以及预防措施等。据统计, 目前全球外来鱼类达624种, 该数量超过30年前的两倍。外来鱼类主要通过水产养殖(51%)、观赏渔业(21%)、休闲垂钓(12%)、渔业捕捞运输(7%)等多种途径被引进。入侵鱼类对本地种产生了捕食、种内种间竞争、杂交和疾病传播等负面影响, 破坏本地生态系统, 但是其正面的生态及社会经济影响也不可忽略。近20年来全球鱼类入侵日益受到重视, 相关论文发表数量翻了8倍。值得提出的是, 近10年来全球鱼类入侵风险评价系统的研究显著增加, 一些鱼类入侵模型已应用于五大洲的多个国家。我国淡水外来鱼类共计439种。然而, 我国关于鱼类入侵的研究起步较晚, 发表文献数仅占全球的3.7%, 且主要研究方向仍集中在入侵物种的分布及生物学特性等基础研究上, 缺乏对于鱼类入侵机制及风险评价预测的研究。因此, 我们建议: (1)开展全国范围的本底调查并建立数据库, 实现数据共享, 明确鱼类入侵的历史与分布现状; (2)联合多个政府部门和机构, 对鱼类入侵进行长期观测, 从整个水生生态系统的角度出发, 深入了解其入侵机制及其产生的正面和负面生态和社会经济影响; (3)加强增殖放流的科学研究和管理; (4)构建区域性外来鱼类入侵风险评价系统, 有效预测鱼类入侵活动, 评价入侵种的危害, 并为相关政府部门的决策提供科学依据。     

Global distribution,entry routes,mechanisms and consequences of invasive freshwater fish

Biological invasion is now considered one of the three major environmental issues worldwide. Freshwater fish invasion becomes more serious with globalization of the world economy. We reviewed the current status of global freshwater fish invasions and discussed the definitions, distributions, introduction pathways, mechanisms, ecological and economic impacts, and risk assessments of freshwater fish invasions. Non-native fish are mainly introduced through food aquaculture (51%), as ornamental fish (21%), or for sport fishing (12%) and fisheries (7%). The number of introduced fish has reached 624 species, doubled the number found thirty years ago. Successful invasions may bring many negative ecological consequences, such as predation, hybridization, structure and function alteration of local freshwater ecosystems, as well as diseases transmission. However, it also brings positive biological and economic values. The number of fish invasion studies has increased eight times over the last 20 years, with studies mainly focusing on biology and the biological impact of invasive fish species. Risk assessments of freshwater fish invasions were studied over the last 10 years, and fish invasiveness screening models have been applied in countries of five continents. The number of non-native freshwater fish in China totaled 439. However, research papers on freshwater fish invasions in China was only 3.7% of the global total, and these researches were mainly on the distribution and biology of invasive fish species, and very few studies included risk assessments. Therefore, we suggest investigating the history, distribution, and mechanisms of invasive species at the national level, evaluating both the positive and negative effects of freshwater fish invasions, and also reinforcing studies of risk assessments in China.     

Recent admixture generates heterozygosity–fitness correlations during the range expansion of an invading species

Admixture, the mixing of historically isolated gene pools, can have immediate consequences for the genetic architecture of fitness traits. Admixture may be especially important for newly colonized populations, such as during range expansion and species invasions, by generating heterozygosity that can boost fitness through heterosis. Despite widespread evidence for admixture during species invasions, few studies have examined the demographic history leading to admixture, how admixture affects the heterozygosity and fitness of invasive genotypes, and whether such fitness effects are maintained through time. We address these questions using the invasive plant Silene vulgaris, which shows evidence of admixture in both its native Europe and in North America where it has invaded. Using multilocus genotype data in conjunction with approximate Bayesian computation analysis of demographic history, we showed that admixture during the invasion of North America was independent from and much younger than admixture in the native range of Europe. We tested for fitness consequences of admixture in each range and detected a significant positive heterozygosity–fitness correlation (HFC) in North America; in contrast, no HFC was present in Europe. The lack of HFC in Europe may reflect the longer time since admixture in the native range, dissipating associations between heterozygosity at markers and fitness loci. Our results support a key short‐term role for admixture during the early stages of invasion by generating HFCs that carry populations past the threat of extinction from inbreeding and demographic stochasticity.     

Molecular systematics and population genetics of biological invasions: towards a better understanding of invasive species management

The study of population genetics of invasive species offers opportunities to investigate rapid evolutionary processes at work, and while the ecology of biological invasions has enjoyed extensive attention in the past, the recentness of molecular techniques makes their application in invasion ecology a fairly new approach. Despite this, molecular biology has already proved powerful in inferring aspects not only relevant to the evolutionary biologist but also to those concerned with invasive species management. Here, we review the different molecular markers routinely used in such studies and their application(s) in addressing different questions in invasion ecology. We then review the current literature on molecular genetic studies aimed at improving management and the understanding of invasive species by resolving of taxonomic issues, elucidating geographical sources of invaders, detecting hybridisation and introgression, tracking dispersal and spread and assessing the importance of genetic diversity in invasion success. Finally, we make some suggestions for future research efforts in molecular ecology of biological invasions.     

Evolutionary history of the little fire ant Wasmannia auropunctata before global invasion: inferring dispersal patterns,niche requirements and past and present distribution within its native range

The evolutionary history of invasive species within their native range may involve key processes that allow them to colonize new habitats. Therefore, phylogeographic studies of invasive species within their native ranges are useful to understand invasion biology in an evolutionary context. Here we integrated classical and Bayesian phylogeographic methods using mitochondrial and nuclear DNA markers with a palaeodistribution modelling approach, to infer the phylogeographic history of the invasive ant Wasmannia auropunctata across its native distribution in South America. We discuss our results in the context of the recent establishment of this mostly tropical species in the Mediterranean region. Our Bayesian phylogeographic analysis suggests that the common ancestor of the two main clades of W. auropunctata occurred in central Brazil during the Pliocene. Clade A would have differentiated northward and clade B southward, followed by a secondary contact beginning about 380 000 years ago in central South America. There were differences in the most suitable habitats among clades when considering three distinct climatic periods, suggesting that genetic differentiation was accompanied by changes in niche requirements, clade A being a tropical lineage and clade B a subtropical and temperate lineage. Only clade B reached more southern latitudes, with a colder climate than that of northern South America. This is concordant with the adaptation of this originally tropical ant species to temperate climates prior to its successful establishment in the Mediterranean region. This study highlights the usefulness of exploring the evolutionary history of invasive species within their native ranges to better understand biological invasions.     

米草属植物入侵的生态后果及管理对策

生物入侵是全球变化的重要组成部分,可能对入侵地造成严重的经济和生态后果,所以评价外来种入侵的生态后果是入侵生态学研究的核心问题之一。本文以米草属(Spartina)植物为例,综述了其对入侵地区自然环境、生物种群、群落和生态系统的影响;结合国际上对米草属入侵种的管理策略,讨论了我国米草属植物管理中的一些重要问题。     

Patterns of introduction and adaptation during the invasion of Aegilops triuncialis (Poaceae) into Californian serpentine soils

Multiple introductions can play a prominent role in explaining the success of biological invasions. One often cited mechanism is that multiple introductions of invasive species prevent genetic bottlenecks by parallel introductions of several distinct genotypes that, in turn, provide heritable variation necessary for local adaptation. Here, we show that the invasion of Aegilops triuncialis into California, USA, involved multiple introductions that may have facilitated invasion into serpentine habitats. Using microsatellite markers, we compared the polymorphism and genetic structure of populations of Ae. triuncialis invading serpentine soils in California to that of accessions from its native range. In a glasshouse study, we also compared phenotypic variation in phenological and fitness traits between invasive and native populations grown on loam soil and under serpentine edaphic conditions. Molecular analysis of invasive populations revealed that Californian populations cluster into three independent introductions (i.e. invasive lineages). Our glasshouse common garden experiment found that all Californian populations exhibited higher fitness under serpentine conditions. However, the three invasive lineages appear to represent independent pathways of adaptation to serpentine soil. Our results suggest that the rapid invasion of serpentine habitats in California may have been facilitated by the existence of colonizing Eurasian genotypes pre‐adapted to serpentine soils.