生物多样性, 2012, 20(5): 581-611 doi: 10.3724/SP.J.1003.2012.31148

《生物多样性》创刊20年纪念专刊

近十年中国生物入侵研究进展

鞠瑞亭1,2, 李慧1, 石正人3, 李博,1,*

1 复旦大学生物多样性和生态工程教育部重点实验室、长江河口湿地生态系统野外科学观测研究站, 上海 200433

2 上海市园林科学研究所植物保护研究部, 上海 200232

3 台湾大学昆虫学系暨研究所, 台北 10617

Progress of biological invasions research in China over the last decade

Ruiting Ju1,2, Hui Li1, Chengjen Shih3, Bo Li,1,*

1 Ministry of Education Key Laboratory for Biodiversity Science and Ecological Engineering, Coastal Ecosystems Research Station of the Yangtze River Estuary and Institute of Biodiversity Science, Fudan University, Shanghai 200433

2 Department of Plant Protection, Shanghai Institute of Landscape Gardening Science, Shanghai 200232

3 Department and Institute of Entomology, Taiwan University, Taipei 10617

通讯作者: *E-mail:bool@fudan.edu.cn

编委: 万方浩

责任编辑: 闫文杰

收稿日期: 2012-07-26   接受日期: 2012-08-13   网络出版日期: 2012-09-20

基金资助: 国家重点基础研究发展计划项目.  2009CB119200
国家自然科学基金.  30930019
上海市科委基础研究重点项目.  10JC1414100
上海市科委基础研究重点项目.  10JC1400700
上海市优秀学科带头人计划项目.  12xd1400500

Corresponding authors: *E-mail:bool@fudan.edu.cn

Received: 2012-07-26   Accepted: 2012-08-13   Online: 2012-09-20

摘要

生物入侵已对入侵区的生态环境、社会经济和人类健康造成了严重的威胁, 成为了21世纪五大全球性环境问题之一。本文回顾了2000年以来, 中国生物入侵研究领域尤其是入侵种的多样性与格局、入侵机制及生态学效应、管理与控制等方面所取得的重要进展, 讨论了需进一步加强研究的领域, 以期为进一步拓展该领域研究的广度和深度、为我国的生物入侵预警预防和科学治理提供参考。据初步研究, 中国的入侵种数量已达529种, 其中陆生植物、陆生无脊椎动物和微生物为主要入侵类群; 原产地以北美洲和南美洲为主; 经济发达和气候温暖湿润的东部和南部省份入侵态势明显较西部和北部省份严重; 随着中国经济的进一步发展, 生物入侵问题将可能更加严峻。外来种的成功入侵是其内禀优势、资源机遇和人为干扰共同作用的结果; 其中, 表型可塑性、适应性进化、天敌释放、种间互利或偏利共生和新化感作用等因素对入侵起到了关键作用。生物入侵已对中国土著生态系统的生物多样性和生态系统服务功能造成了严重影响, 打破了生态系统的固有平衡, 危害或威胁到中国的农林牧渔业生产、交通航运、环境、人类健康和公共设施安全。针对生物入侵的管理与控制, 中国加强了包括检测监测、风险分析、生物防治、扩散阻断、根治灭除和生态恢复等技术体系的研究和实施, 并初步控制了一些重要入侵种的扩张。中国生物入侵需要在全境性科学考察、生物入侵的遗传学、基因组学、生态系统影响、全球变化和管理与控制技术创新等领域进一步加强跨领域的交叉合作和系统研究。

关键词: 生物入侵 ; 中国 ; 管理与控制 ; 多样性 ; 生态学效应 ; 入侵机制

Abstract

As one of the five major global environmental problems, invasive species have posed serious threats to native ecosystems, public health, and regional economies. Although much progress has been made in the field of biological invasions research in China over the last decade, there are still large knowledge gaps. This paper reviews progress in the field of biological invasions research since 2000 as it relates to China, covering the diversity, colonization and immigration patterns of invasive species, mechanisms and ecological effects of biological invasions, and management and control of invasive species. In China, 529 invasive alien species have been identified, which originated primarily from South and North America, and the major taxa included terrestrial plants, terrestrial invertebrates, and microorganisms. We found a higher prevalence of invasive species in the eastern and southern provinces, compared to the western and northern provinces in China. This pattern is likely due to the differences in the level of economic development and environmental suitability between the two regions. Moreover, with further economic development, China may face more serious biological invasions in the future. These invasions of alien species are largely the combined results of the interactions between the intrinsic traits of these species along with resource opportunities and disturbances by human beings. Many mechanisms are responsible for successful invasions of alien species, but phenotypic plasticity, adaptive evolution, enemy release, interspecific mutualism or commensalism, and new allelochemicals may be primary causative factors. Biological invasions in China have caused serious impacts on native ecosystems, including biodiversity and ecosystem services, alteration of biogeochemical cycles, threats to agricultural and forestry production, traffic and shipping, environmental safety, and public facilities. China has also made progress in the detection and monitoring of invasive species, risk analysis, biological control, radical elimination, and ecological restoration of degraded ecosystems. We suggest several issues that need to be addressed in invasive species research in the future, including territory-wide inventories, evolutionary ecology and genomics, direct and indirect ecosystem-level consequences, interactions between major components of global change and biological invasions, and management and control technologies.

Keywords: biological invasions ; China ; control and management ; diversity ; ecological effects ; invasion mechanisms

PDF (737KB) 元数据 多维度评价 相关文章 导出 EndNote| Ris| Bibtex  收藏本文

本文引用格式

鞠瑞亭, 李慧, 石正人, 李博. 近十年中国生物入侵研究进展. 生物多样性[J], 2012, 20(5): 581-611 doi:10.3724/SP.J.1003.2012.31148

Ruiting Ju, Hui Li, Chengjen Shih, Bo Li. Progress of biological invasions research in China over the last decade. Biodiversity Science[J], 2012, 20(5): 581-611 doi:10.3724/SP.J.1003.2012.31148

自“动物生态学之父”Charles Elton的The Ecology of Invasion by Animals and Plants一书出版以来(Elton, 1958), 全球对外来种(alien species)入侵的关注度与日俱增。一方面, 受人类干扰后的多数生态系统的抗入侵能力弱, 大量现有入侵种扩散和暴发趋势严重(Pyšek et al., 2010), 生态系统的完整性受到了极大的破坏; 另一方面, 全球化进程使得众多新的外来种传入速度进一步加快(Weber et al., 2008), 新的生物入侵风险不断加大, 社会经济发展不断面临新的威胁。在此背景下, 外来种与土著种在几乎所有生态系统中普遍混生, 我们在从事生产活动时, 已无法排除外来种的干扰(Lockwood et al., 2007)。鉴于生物入侵(biological invasions)对入侵区的生态环境、社会经济和人类健康造成了严重的威胁, 这一问题得到了各国政府、国际组织、社会公众和科学界的广泛重视, 已成为21世纪五大全球性环境问题之一(Millennium Ecosystem Assessment, 2005)。近10余年, 全球科学家在生物入侵的基础和应用研究中投入了巨大的努力, 并取得了丰硕的成果, 这不仅为我们遏制现有入侵种的进一步蔓延提供了依据, 而且也为我们将来抵御新的生物入侵提供了有效的措施。

中国是一个农业大国, 生产性人口规模巨大, 高强度的生产和经济活动给外来种的传入和散布创造了有利条件; 同时, 中国又是一个生物多样性大国, 景观、气候、生态系统类型的多样性, 造成了外来种很容易在我国领土上长期定殖; 更为重要的是, 近30年来中国经济始终处在高速发展期, 与世界各国和地区的物品和人员交流不断增多, 进一步加剧了外来种入侵中国的数量和频率(李博和马克平, 2010), 由此造成了我国当前入侵种的数量相当庞大, 影响也日趋严重。出于国家公共安全的整体需求, 我国科学家从20世纪90年代开始, 逐渐重视生物入侵研究。

我们高兴地看到, 虽然中国的生物入侵研究起步较晚, 但受到各级政府对这一问题的高度重视, 近10多年来, 中国在这一领域的研究已取得了巨大进步, 获得了诸多的重要研究进展, 在国际生物入侵舞台上也有了一定的学术地位。2008年以来, 我国科学家出版了《入侵生物学》(万方浩等, 2011a)、《生物入侵: 预警篇》(万方浩等, 2010)、《生物入侵: 检测与监测篇》(万方浩等, 2011b)、《生物入侵: 生物防治篇》(万方浩等, 2008a)、《生物入侵: 管理篇》(万方浩等, 2008b)等一系列专著。这些专著的出版, 以及近年来我国科学家在生物入侵领域内获得的诸多研究进展, 标志着中国的生物入侵研究已形成了较为全面和丰富的研究体系。当前, 具有区域特色的生物入侵研究模式和入侵生物学学科框架已逐渐形成(万方浩等, 2011c)。

同时, 我们很欣喜地发现, 在我国科技界的共同努力下, 截至2010年, 中国的生物入侵科学论文数量已在发展中国家排名第二(Nuñez & Pauchard, 2010), 并进入全球前10名的行列(万方浩等, 2011a, c), 尤其是对一些重要入侵种的研究已取得众多国际认可的喜人进展(李博和马克平, 2010), 相关成果不仅为国际入侵生物学研究提供了基础资料, 也为中国国家生物安全管理与控制策略的构建提供了重要依据。可以预见, 中国的生物入侵研究将会以更快的速度发展, 并逐渐接近世界先进水平。为总结本世纪以来中国生物入侵研究取得的主要成就, 我们对2000年以来中国科学家在入侵种的多样性与格局、入侵机制及生态学效应、生物入侵的管理与控制等方面的主要成就进行了综述, 并对该领域的进一步研究方向提出了建议, 以拓展中国生物入侵研究的广度和深度, 为生物入侵的预警预防和科学治理提供参考。

1 生物入侵的格局与趋势

1.1 入侵种的多样性与格局

采用文献调研、实地考察与专家咨询相结合的方式, 中国学者先后对境内的陆生、水生和海洋生态系统中入侵的陆生植物、陆生无脊椎动物、微生物、水生无脊椎动物、水生植物、鱼类、哺乳类、两栖爬行类和鸟类物种的种类进行了调查, 并从这些入侵种的分类地位、起源地、传入途径和时间、入侵生境和空间分布等角度分析了入侵的格局, 据此预测了中国外来种进一步入侵的趋势。

(1)数量: 查明我国外来入侵种共529种, 其中植物、动物、微生物分别为270种(Weber et al., 2008)、198种和61种, 物种数最多的类群为陆生植物、陆生无脊椎动物和微生物(图1)(万方浩等, 2009; 徐海根和强胜, 2011)。值得注意的是, 最近的研究发现, 中国的外来陆生草本植物总数就多达800种, 只是其中65%的物种其入侵性尚不明确(杨博等, 2010)。因此, 推测我国境内入侵种的实际数量可能远大于目前已列入编目的数量。

图1

图1   中国入侵物种各类群的比例(万方浩等, 2009)。1: 陆生植物; 2: 陆生无脊椎动物; 3: 微生物; 4: 水生无脊椎动物; 5: 水生植物; 6: 鱼类; 7: 哺乳类; 8: 两栖爬行类; 9: 鸟类。


(2)分类学: 中国的入侵植物来自48科177属, 其中双子叶和单子叶植物分别约占79.6%和20.4%。大陆地区最常见的入侵种来自菊科、禾本科和十字花科; 而台湾地区则多来自苋科、旋花科和柳叶菜科, 两地均未发现蕨类植物或其近缘种入侵(Wu et al., 2004; Weber et al., 2008)。在入侵动物中, 鞘翅目(29种)、同翅目(19种)、双翅目(13种)和鳞翅目(8种)的农林害虫种类最多, 其他入侵动物类群还涉及鱼类、两栖类、爬行类、哺乳类、鸟类, 以及海鞘类、螨类、虾、藤壶、蛞蝓、蜗牛、蛤、贝、螺、苔虫、线虫、海胆、盘管虫、粘体虫等(万方浩等, 2009)。在入侵微生物中, 植物病原真菌是最大的类群, 约占44.3%; 植物和养殖鱼虾类的病毒次之, 约占36.1%; 另外有19.6%的微生物为植物病原细菌(徐海根和强胜, 2011)。

(3)来源: 大陆地区绝大多数的入侵种来自北美洲(黄顶成和张润志, 2011); 而台湾、香港、澳门绝大多数的外来种多来自南美洲(Wu et al., 2004; 王发国等, 2004; 李一农等, 2007)。北美洲之所以成为中国大陆入侵种的主要输入地, 可能是因为北美同中国大陆有着较为相似的气候环境, 绝大多数来自该地区的物种传入中国后能很快适应新的生境条件, 在较短的时间内就能成功定殖; 同时, 中国作为北美的最大贸易伙伴国之一, 频繁的贸易和人员交流使外来种传入中国大陆的机会相对较多(Weber et al., 2008)。港澳台地区主要入侵种来自南美洲, 则主要是因为这些地区同南美洲的气候与生境类型具有更高的相似性。

(4)传入途径: 中国95%以上的入侵种是人为引入或带入的, 它们或作为牧草、饲料、工业材料、药用植物、蔬菜、草坪、养殖、观赏、生物防治等目的而被有意引进, 或是随引进的原木、苗木、花钵、土壤而被无意传入。其中, 大陆约有50%的外来植物和25%的外来动物是有意引入的, 有76.3%的入侵动物是由于检疫疏漏, 随贸易品或运输工具而无意传入的(徐海根等, 2004); 台湾有约60%的外来植物是无意传入的, 而另外40%的外来植物则是作为园艺品种、药用植物或畜牧饲料而引入, 但其中仍有50%的植物其实际的农业使用价值未知(Wu et al., 2004)。

(5)传入和成灾时间: 自16世纪以来, 中国外来生物入侵的时间可划分为3个阶段(万方浩等, 2009): 第一阶段为缓慢增长阶段(16-19世纪), 这一阶段由于对外交流较少, 传入的外来种数量不到100种; 第二阶段为快速增长阶段(19-20世纪中前期), 这期间由于外国列强陆续侵入中国, 导致外来种传入的数量急剧增加; 第三阶段为暴发成灾阶段(20世纪后期至今), 此阶段中国经济飞速发展, 国内产业结构不断调整, 各类商品和物品的流通十分频繁, 外来种的扩张速度和暴发机会加大, 许多检疫性有害生物也随之入侵。从外来种开始传入到暴发成灾的时间长度来看, 入侵植物传入后暴发成灾的时间相对较长, 一般在50年左右; 而入侵昆虫和微生物暴发成灾的时间相对较短, 一般在10年左右(万方浩等, 2009)。

(6)生境: 外来种在我国入侵的生态系统包括农田、森林、灌丛、草原、河流和湖泊等湿地、海洋、岛屿、城市和自然保护区等各种类型, 其中有超过80%的入侵种出现在农田、森林和植物园等人工干扰频繁的生境中(徐海根和强胜, 2011; Lu & Ma, 2005)。

(7)空间分布格局: 全国入侵种(如植物)的密度呈现出东部和南部省(市、区)高、西部和北部省(市、区)低的空间格局(吴晓雯等, 2006; Weber et al., 2008)。其中, 广东、福建、江苏、云南、台湾等5个省的重要入侵种数量在200种以上; 而宁夏、青海、甘肃等3个省(自治区)的重要入侵种数量总共不到70种(万方浩等, 2009)。该格局与我国土著物种多样性、气候适宜性、人口和交通密度、人类活动强度和经济发展水平的空间分布格局基本一致(Liu et al., 2005); 但入侵种数量和各省面积没有显著相关性(Weber et al., 2008)。最近, Huang等(2012)利用中国2008年以前发现的166种农林入侵物种为对象, 从“首发地”(first detection location)的角度进一步分析发现, 辽宁、北京、天津、河北、山东、江苏、上海、浙江、福建、广东、广西和海南等12个省(直辖市、自治区)的土地面积仅占全国的15.6%, 而入侵物种首发地所占比例却高达74.6%。这更加系统地说明了入侵种传入中国后在沿海经济发达省区首先登陆的趋势明显强于内陆省区。

(8)入侵种的传播和扩散: 入侵种的传播和扩散是入侵种产生危害和影响的最重要过程。我国对一些破坏性较大的入侵种的扩散过程开展了研究, 相关成果为我们认识外来种入侵后为什么产生巨大生态影响和入侵种的管理和控制提供了依据(Wang & Wang, 2006; Sang et al., 2010)。如对于紫茎泽兰(Eupatorium adenophorum)的研究发现, 该入侵植物的扩散路线是沿着道路和河流展开的, 由此确定了对其防控重点(Lu & Ma, 2005; Zhu et al., 2007)。同时, 入侵物种的潜在分布区预测也取得了一定进步, 并在国家入侵物种防控和管理政策的制定中得到了成功应用。

1.2 入侵趋势预测

与美国的入侵植物区系比较研究发现, 中国木本入侵种的比例明显比美国低, 未来将可能面临更多木本植物的入侵; 道路密度与入侵物种密度呈正相关; 与美国相比, 中国无论是道路密度还是入侵物种密度均较低; 而美国入侵植物的丰富度相当于中国的2倍, 因此, 中国生物入侵形势还有进一步恶化的风险(Weber & Li, 2008)。最近, Lin等(2011)通过对全球91个经济体的生物多样性水平、入侵种数量和经济发展水平的分析表明, 入侵种数量与各经济体的经济发展水平呈正相关。自20世纪70年代以来, 中国经济的迅速发展被证明是生物入侵强度加大的主要驱动力(Lin et al., 2007; Ding et al., 2008); 同时, 近年来中国快速兴起的城市化进程也推动了入侵种的区域性扩张(Wang et al., 2011; 鞠瑞亭和李博, 2012)。因此, 随着中国经济的进一步发展, 推测我国生物入侵的趋势还将更加严峻, 国家相关部门应采取更加严格和系统的管理措施, 防止新的生物入侵(Ding et al., 2008; Weber & Li, 2008)。

2 外来种入侵的机制

面对日趋严重的生物入侵问题, 科学家们一直致力于回答为什么有些外来种能在原产地以外的分布区成功入侵, 并在被入侵生态系统中“反客为主”而成为优势种(Cox, 2004)。为了解释这个问题, 近50年来学术界提出了近30种不同的假说(Catford et al., 2009)。这些假说在一定程度上确实解释了外来种入侵存在一定的规律; 但同时我们发现, 任何单一假说都很难解释普遍存在的生物入侵现象(Moles et al., 2008), 而常常需要联合多种相互关联的机制才能解释(Blumenthal, 2005), 甚至外来种入侵的机制往往就是因种而异的。近10年来, 我国学者在一些重要外来种的入侵机制研究方面取得了诸多进展, 并根据其中的一些关键机制提出了较多新的入侵假说。在此, 我们以中国农林和湿地生态系统中重要入侵种为对象, 着重探讨我国境内外来种入侵的一般机制。

2.1 外来种的成功入侵是其内禀优势、资源机遇与人类干扰共同作用的结果

尽管试图以一个统一的“理想特征”(ideal character)来归纳外来种成功入侵的机制很困难, 但无可争议的是, 成功入侵的外来种必然具备其独特的内禀优势, 这一观点得到了学术界的广泛认同, 由此形成了入侵的“内禀优势假说”(inherent superiority hypothesis)(Hufbauer & Torchin, 2007)。该假说认为, 外来种在传播和扩散的过程中, 某些内禀的优势特征(如形态、生理、生态、遗传和行为等)决定了其入侵行为的产生。这可能是因为相对于土著种, 具有内禀优势的外来种在进化中获得了更多的变异, 从而具有耐受更广泛的环境条件、利用更多资源类型并抵抗大量潜在天敌控制的能力或性状, 最终在竞争中获得了优势。

在我国成功入侵的外来种, 大都可以部分地用“内禀优势假说”来解释。例如, B型烟粉虱(Bemisia tabaci B-biotype)的成功入侵受其寄主广泛(Li et al., 2011)、繁殖能力大、抗药性强(褚栋等, 2004)、对逆境环境具有极强适应性(Cui et al., 2008; Wan et al., 2009; Yu et al., 2012a, b)等独特的内禀优势驱动。相对于土著近缘种拟松材线虫(Bursaphelenchus mucronatus), 松材线虫(B. xylophilus)具有更高的繁殖潜能, 对环境也有更强的生态和生理适应能力, 故在入侵过程中较快地替代前者(Cheng et al., 2008a)。稻水象甲(Lissorhoptrus oryzophilus)在原产地北美, 两种色型(深色和浅色型)的雌雄虫均十分常见; 但在入侵地中国, 能营孤雌生殖、具有非常显著的生殖优势的深色型雌虫占优势(Jiang et al., 2006, 2007, 2008), 这是该害虫在中国迅速扩张的原因之一。红火蚁(Solenopsis invicta)能迅速入侵的重要原因之一是其繁殖潜力极大, 一头蚁后每天可产卵1,500-5,000枚, 蚁后寿命可达6-7年(Chen et al., 2011)。入侵我国城市生态系统的悬铃木方翅网蝽(Corythucha ciliata), 具有个体发育时间短(世代发育周期20-24 d)、种群扩繁速度快(我国长江流域每年可发生5代, 比原产地多2-3代)、生殖能力强(单雌平均产卵量达280枚)等生物学特性(鞠瑞亭和李博, 2010; Ju et al., 2011a); 对新入侵区内的高温具有较强的适应性, 高温胁迫对其种群的存活和繁殖影响小(Ju et al., 2011b); 同时, 该物种虽属寒冷敏感型(chill-susceptible)昆虫(Ju et al., 2010), 但因其具有快速冷锻炼(rapid cold hardening, RCH)的生理调节功能, 可使越冬耐寒性明显提高(Ju et al., 2011c), 这些生态和生理优势是该虫能在不同温度带快速入侵的内在因素之一。大豆疫霉菌(Phytophthora sojae)对我国控制植物疫病的主要药剂甲霜灵(Metalaxyl)可产生极强的抗药性, 突变菌株(Mtr)的抗性水平可达野生型单游动孢子菌株的870倍以上(Cui et al., 2010)。致病性相关基因对寄主信号分子转录应对反应的加快是该病菌与寄主适应的主要机制之一, 在其克服寄主的抗病性中起着重要作用(Chen et al., 2007c; Wang et al., 2009, 2010)。因此, 现有药剂对大豆疫霉控害效果有限, 使其入侵能力得到了表达, 这在一定程度上促进了该病害的广泛流行(Wu et al., 2010; Wang et al., 2011)。

“内禀优势假说”虽然充分肯定了外来种本身的内禀优势在入侵中的作用, 并且在统计意义上来讲, 它可以较好地解释绝大多数外来种的入侵性; 但必须注意到, 在入侵形成的过程中, 除了外来种本身需具备入侵的特性外, 资源的可利用性亦应起到关键作用。如果不考虑资源问题, 单依靠物种的内禀特征, 还是无法解释外来种为什么只在特定地区、特定生态系统中发生入侵的现象。也就是说, 内禀优势通常只表明外来物种的入侵潜力, 但如果没有相应资源条件与之匹配, 这些特征就无法在入侵的过程中得到表现。事实上, 外来种被引入到一个新的生态系统中, 资源的可利用性决定了其在生态系统中的适合度和丰富度, 如果系统中的营养、空间和时间生态位恰好符合入侵的条件, 则外来种容易形成入侵, 也就是说, 群落的可入侵性与可利用资源的多少紧密相关。据此理论, Davis等(2000)在对大量案例的整合分析基础上, 提出了“资源机遇假说”(resource opportunity hypothesis), 该假说认为, 在大尺度的空间范围内, 可利用资源是决定生态系统可入侵性的关键因素, 群落一旦具有入侵种必需的生态资源, 包括营养、食物、水分、光照、土壤等, 而且这些生态资源大多没有被土著种有效利用, 事实上也就是必须存在空余生态位, 这种场景恰好遇上外来种的繁殖体, 便会发生生物入侵。例如, 稻水象甲在我国浙江稻区的入侵成功, 便证明了“资源机遇假说”的正确性。在浙江, 生境和食料条件是影响稻水象甲种群数量扩张的关键因子, 浙江水稻产区稻田边的山坡为稻水象甲的越冬提供了庇护, 而田埂上生长的杂草则为该虫提供了较好的早期食源和产卵环境(Jiang et al., 2004a, b; Shi et al., 2008), 杂草的存在可能起到了桥梁寄主的作用。另外一个更加鲜明的实例是椰心叶甲(Brontispa longissima), 该虫自2002年被发现传入我国以后, 华南地区的气候条件同原产地印度尼西亚和巴巴亚新几内亚极为相似, 都属于跨热带和亚热带的高温高湿地区, 而椰心叶甲恰好偏喜这种环境, 适宜的气候条件为椰心叶甲的繁殖扩散提供了先决条件(陆永跃和曾玲, 2004)。同时, 我国南方地区丰富的棕榈科植物资源, 为该虫的繁衍提供了数量巨大的食物来源(鞠瑞亭等, 2005); 而棕榈科植物上食叶害虫极少, 生态位空缺造成了该外来种一旦传入, 即会占据并充分利用充足的资源生态位(覃伟权等, 2010)。此外, 椰心叶甲主要生活在棕榈科植物心叶内, 其外围有紧密的成熟剑叶包裹, 稳定的微环境为椰心叶甲的取食、生存和种群发展提供了有利条件(陆永跃和曾玲, 2004)。以上原因共同促进了椰心叶甲在我国南方地区的成功入侵。

实际上, 我们认为, 最近有关气候变暖推动生物入侵形势严峻化的推测, 主要是因为温度这一“资源机遇”的作用。鉴于昆虫属于变温动物, 相对于其他生物, 其对温度的敏感性更强, 因此成为研究气候变暖与生物入侵相互关系较为适合的对象。Huang等(2011)发现, 1900-2005年中国大陆入侵昆虫定居速度与气温升高呈显著的正相关性, 大气温度每升高1 °C, 入侵昆虫的数量每10年就增加5种。这预示着全球变暖直接影响了外来昆虫的入侵, 在制定入侵物种控制策略和政策的过程中, 我们需要充分考虑到气候变暖的重要影响。

“资源机遇假说”还很好地解释了有些近缘外来种在入侵后为什么会相互共存或取代。譬如, 美洲斑潜蝇(Liriomyza sativae)、南美斑潜蝇(L. huidobrensis)和三叶斑潜蝇(L. trifolii)是3种先后入侵中国的近缘种害虫, 主要危害蔬菜、花卉植物。在三叶斑潜蝇传入以前, 美洲斑潜蝇和南美斑潜蝇种群已在我国共存11年, 种间竞争和取代现象并不明显。自2005年三叶斑潜蝇入侵东南沿海省份后, 原先占优势的美洲斑潜蝇种群现已明显减少, 在许多地区, 前者已取代后者成为入侵区内的优势种群(相君成等, 2012)。美洲斑潜蝇和南美斑潜蝇的长期共存, 主要是它们对环境的适应性存在差异, 相对较高的温度有利于美洲斑潜蝇种群的增长, 而相对较低的温度则有利于南美斑潜蝇的增殖(Huang & Kang, 2007), 这种差异导致两种斑潜蝇在不同的季节出现, 难以形成竞争, 因此, 热适应性分化是导致两种斑潜蝇在中国种群长期共存的主要原因(Zhao & Kang, 2000; Kang et al., 2009)。三叶斑潜蝇近年之所以取代美洲斑潜蝇, 主要是两种斑潜蝇均喜在较温暖的低海拔地区生存, 地理生态位基本重叠, 易形成竞争, 而在竞争中, 三叶斑潜蝇较美洲斑潜蝇具备更好的高温逆境耐受能力, 这导致在夏季种群竞争时, 前者比后者更占优势(相君成等, 2012)。

上面介绍了物种本身特性、资源可利用性在促进外来种入侵的过程中的作用。为了更全面地揭示入侵的成因, 人为干扰这一人类的主观作用还必须被充分考虑。譬如, 互花米草(Spartina alterniflora)的入侵, 除了自身优势以外, 很大程度上是因为我国海岸带盐沼生态系统中土著高等植物的多样性低、人类活动干扰频繁和环境的变化等有利于其生长和繁殖(Li et al., 2009); 尤其是长江河口频繁出现的咸水侵入和我国东海岸的高强度围垦更是助推了该入侵种的定居和扩张(Wang et al., 2006)。水葫芦(Eichhornia crassipes)种子萌发和个体生长与水体中磷、氮含量密切相关, 因此, 水葫芦更易入侵人类干扰后富营养化严重的水体(Chen et al., 2010b)。Ascunce等(2011)使用微卫星标记和基因比对技术, 对全球75个地区2,144个红火蚁种群的入侵源地进行判别的研究发现, 几乎所有新入侵区的红火蚁种群都源自美国南部; 其中, 最早入侵中国大陆的广东吴川种群在遗传学上与美国德克萨斯州种群最为接近, 该州是美国南部红火蚁疫区中对中国出口贸易最大的州, 因此推测贸易活动是红火蚁入侵中国的一个重要原因(Yang et al., 2012)。人类干扰还间接推动了新入侵种迅速取代先前入侵的近缘种, 如Q型烟粉虱(B. tabaci Q-biotype)自2003年入侵中国以后(Chu et al., 2006), 近年来在许多地区取代B型烟粉虱成为优势种群(Chu et al., 2010a, b; Teng et al., 2010), 此现象可能与农事活动中化学农药的频繁使用等因素有密切关系(Luo et al., 2010; Pan et al., 2011)。总的来说, 干扰的作用一方面可能为先锋入侵种的奠基提供了途径和机会; 另一方面更为外来种传入后提供了空余生态位机遇, 使外来种在与土著种的竞争中, 更有可能接触到和利用到现有的资源, 或通过干扰消灭了竞争者, 从而直接或间接地帮助了外来种的入侵。

2.2 表型可塑性和适应性进化

表型可塑性(phenotypic plasticity)是同一基因型对环境变化表现出的形态、生理和行为上的适应性响应特征, 它是生物对环境的一种适应, 反映了通过遗传同化作用于入侵种的自然选择过程, 并最终使得新的表型得到稳定表达, 从而使外来种迅速适应入侵地的环境。事实上, 这种可塑性特征属于生物有机体内禀优势的一个方面。多个入侵实例表明, 表型可塑性强的外来种通常更容易形成入侵(Richards et al., 2005)。例如, 互花米草在与土著种芦苇的竞争中, 其极强的竞争优势正是表型可塑性差异的结果(Zhao et al., 2010)。互花米草在淹水和高盐度环境中, 相比于土著植物芦苇, 对土壤氮利用表现出更强的优势(Wang et al., 2006; Zhao et al., 2010), 其光合作用能力也较芦苇更强(赵广琦等, 2005), 因此表现出明显的入侵能力。

外来种能否成功入侵不仅取决于自身的生物学特征, 同时也与其在原产地的进化历史、引种过程及其与土著生态系统的相互作用等密切相关。因此, 外来种的入侵性可能并不是天生的, 一些物种的入侵性是引入后才获得的, 即进化而来的(冯玉龙等, 2009)。这种进化主要由环境压力、奠基者效应(founder effect)、遗传漂变(genetic drift)以及杂交(hybridization)等作用所引起(Bossdorf et al., 2005), 因而属于适应性进化。外来种入侵一个新的生态系统以后, 因在入侵地缺少天敌尤其是专化性天敌, 此时进化会使外来种降低对天敌的防御能力, 并把原来用于防御的资源用于生长、繁殖等过程, 从而提高竞争能力、促进入侵(冯玉龙等, 2009)。然而, 过去我们并不清楚资源再分配是如何使入侵种提高竞争能力的, 相关文献仅关注入侵种是否降低了对天敌的防御能力、是否提高了竞争能力, 而忽略了二者之间的内在联系。以紫茎泽兰等为研究对象而得出的“氮分配的进化假说”(hypothesis of the evolution of nitrogen allocation)(Feng et al., 2009, 2011)认为, 天敌逃逸可使外来入侵植物降低叶氮向防御的分配, 同时增加氮向光合作用的分配, 这不仅提高了叶片的光合能力和光合氮利用效率, 还提高了光合能量利用效率, 缩短了叶片建成成本(construction cost)的偿还时间, 这种独特的快速偿还型能量利用策略验证了入侵植物的氮分配进化优势(Feng et al., 2007a, c, 2008, 2009; Feng, 2008a, b; Feng & Fu, 2008)。

遗传变异(genetic variation)为外来种的适应性进化提供了生物学基础; 但研究发现, 由于小种群效应, 入侵地种群的遗传多样性往往会小于原产地种群。然而, 不同来源个体的多次入侵事件能增加种群的遗传变异, 种间杂交和基因渐渗(introgres- sion)也会导致外来种基因组的快速改变, 增加入侵种群的遗传变异, 并引起一系列适应性进化特征。例如, 松材线虫在中国的入侵过程中, 由奠基者和遗传漂变引起的瓶颈效应并不明显, 中国的入侵虫源来自多个国家, 不同来源种群的多次大量入侵, 导致其入侵种群拥有了丰富的遗传变异, 入侵过程中保持较高的遗传多样性是该线虫成功入侵的遗传学机制之一(Cheng et al., 2008b; Xie et al., 2009), 并对该物种的适应性进化和入侵行为的产生具有重要的意义。

表型可塑性和遗传变异是生物适应异质生境的两种重要对策。值得注意的是, 这两种对策在外来种入侵的过程中虽然并不相互矛盾, 但二者对成功入侵的相对重要性一直是入侵生态学争论的焦点之一(施雯等, 2010)。最近有学者对紫茎泽兰的研究发现, 表型可塑性和遗传分化共同促进了其入侵, 由于受地理距离和海拔因素的影响, 入侵中国的紫茎泽兰产生了一定程度的遗传分化, 形成了不同的地理种群, 这些地理种群表现出多样化的表型可塑性性状, 丰富的遗传多样性和较高的表型可塑性使紫茎泽兰快速适应了新的环境, 从而形成了广泛的入侵(Gui et al., 2008, 2009; Wan et al., 2010)。然而, 针对喜旱莲子草(Alternanthera philoxeroides)的研究则发现, 表型可塑性而非遗传变异是其成功入侵的机制(Pan et al., 2006; Geng et al., 2007a; Jia et al., 2010; Gao et al., 2011)。围绕喜旱莲子草在不同水分异质生境下的适应对策, 采用同质园实验、分子标记和野外调查等方法研究发现, 在野外和同质园交互移植条件下, 不同生境的喜旱莲子草表现出极显著的表型趋异, 但在单独的同质园条件下, 这种差异不复存在; 分子标记分析发现, 尽管所研究的90个“个体”采自不同生境和地点, 但大多数个体具有相同的ISSR基因型(Geng et al., 2006, 2007a, b)。由此可见, 表型变异主要是对不同生长条件产生的可塑性反应的结果。对于喜旱莲子草而言, 在入侵不同生境过程中表型可塑性的作用可能比遗传变异更为重要(Li & Ye, 2006)。然而, 在许多其他入侵种的扩散过程中, 若其具有较强的表型可塑性, 则可弥补遗传多样性低所产生的不足, 减少种群对生态型分化的依赖, 从而适应异质的生境(耿宇鹏等, 2004)。例如, 入侵中国的水葫芦虽然遗传多样性较低, 几乎所有种群的基因型是相同的, 但水葫芦无性系分株呈现出较高的多样性和表型可塑性, 这可能是其成功入侵的机制之一(Li et al., 2006; Ren & Zhang, 2007)。甚至, 如前一节中的多个实例, 许多外来种因其本身固有的内禀优势很强, 它们的入侵具有环境特异性, 其生态位幅度足以维持在被入侵生境中的建群需求, 这些外来种因此既不需要表型可塑性, 也不需要遗传多样性。事实上, 从生物入侵的整体角度来看, 迄今为止, 人们还不能把所有外来种入侵的机制都用表型可塑性和遗传分化来统一解释。因此, 表型可塑性和遗传变异对入侵的相对重要性, 甚至各自间的独立重要性也是因种而异的。

2.3 天敌释放

在前面的章节中, 我们在描述“资源机遇”的作用时, 需要考虑的是, 这个假说在描述外来种入侵的机制时还存在一定的缺陷。这主要是因为该假说在提出时, 未能考虑到土著生态系统中其他物种对外来种的寄生或取食。为了更好地解释物种入侵的机制, 利用“资源机遇假说”时, 还必须联合其他假说。“天敌释放假说”(enemy release hypothesis)正是可以与之结合的最佳匹配。该假说认为, 外来种之所以能在入侵地迅速扩张, 是由于脱离了原产地自然天敌的控制作用所致。“天敌释放假说”的理论依据主要有以下3点: (1)自然天敌是外来种种群的重要调控因子; (2)本地天敌对土著种的控制作用大于对外来种的控制作用; (3)外来种可以利用在入侵地天敌调控作用降低的条件促进种群增殖(万方浩等, 2011a)。

最近, 我们在研究互花米草入侵长江口崇明东滩盐沼生态系统时有趣地发现, 互花米草的入侵正是“天敌释放假说”的最好例证。在原产地北美, 一种叫光蝉(Prokelisia marginata)的昆虫能有效控制互花米草向周边区域的扩散, 两种茎秆钻孔的飞蝇(Chaetopsis aeneaC. apicalis)幼虫对互花米草生长也有一定的限制作用; 病原微生物麦角菌(Claviceps purprea)能够感染互花米草的花部, 在种子内形成菌核, 降低种子的产生(Gray et al., 1991);互花米草在引入中国时, 这些原产地的天敌均未能一起传入。虽然我们在崇明东滩发现了一种叫素毒蛾(Laelia coenosa)的广食性土著昆虫能够取食互花米草, 但其本身就是芦苇(Phragmites australis)等多种本地植物的重要害虫, 初步观察发现, 素毒蛾取食芦苇的能力要大于互花米草, 这可能恰好证明了入侵地的广谱性天敌对土著植物芦苇的控制作用大于对外来植物互花米草的控制作用, 从而有助于互花米草的入侵。另外有一个美国白蛾(Hyphantria cunea)生物防治的实例则可以证明, 合理利用“天敌释放假说”, 可以从相反的角度挖掘本土天敌资源, 控制外来种的入侵。美国白蛾于1979年传入中国以后, 我国科学家发现并利用土著天敌昆虫白蛾周氏啮小蜂(Chouioia cunea)成功控制了该虫的进一步扩散, 在以利用该天敌控制为主, 联合其他措施的共同防控下, 目前上海、大连、青岛、烟台等几个疫点的美国白蛾已被根治(杨忠岐和张永安, 2007)。此例说明入侵地如果能发现高效率的土著天敌, 对外来种的控制效果好, 被入侵地的天敌调控作用增强可以降低入侵种种群的扩张速度。

2.4 种间互利或偏利共生

生态系统中一个物种的存在往往会依赖于与其他物种的共生。与那些不依赖于其他生物就能独立完成生活史的物种相比, 需要共生生物的物种在入侵地成功建群的难度更大。因此, 缺乏互利共生(mutualism)的生物可能会阻碍外来种的入侵; 也就是说, 如果一个外来种被引入时, 与其共生的互利生物没有被一同引入, 这个物种在入侵地就很难定殖; 相反, 一个外来种在传播的过程中, 如果一同带入了与之互利共生的物种, 则可以帮助其成功入侵。例如, 红脂大小蠹(Dendroctonus valens)与其伴生菌的共生入侵促进了其在吕梁山区的建群就验证了这一推论(Lu et al., 2007b, 2009, 2010)。研究表明, 一种叫长梗细帚霉(Leptographium procerum)的真菌是由红脂大小蠹携带从美国传入中国, 长梗细帚霉在中国形成了独特的单倍型, 相对于美国单倍型和中美共有单倍型, 这些中国独特单倍型提高了红脂大小蠹在寄主油松(Pinus tabuliformis)上与土著种的竞争能力, 还显著诱导了油松产生红脂大小蠹聚集信息素三蒈烯(3-carene)来协助该害虫的聚集。这一结果衍生出真菌独特单倍型促进虫菌共生入侵的新假说。更有趣的是, 在此基础上提出的“返入侵假说”认为, 在一些原产地的次生性害虫/伴生菌, 在新的入侵地发生变异后, 如再返回到原产地, 也会成为可以危害健康寄主树木的有害生物(Lu et al., 2011)。

然而, 大部分外来种在传播的过程中, 极少有机会携带原产地的共生生物, 但它们在新传入地区通常会遇到比原产地更好的共生体(如传粉者、共生菌等), 外来种与土著种之间新建立的共生关系亦能提高其入侵能力(Richardson et al., 2000)。例如, B型烟粉虱与土著的植物双生病毒之间由于存在互利共生关系, 它可有效获得、携带和传播双生病毒, 导致植株发病(Liu et al., 2009, 2010); 而当B型烟粉虱取食这些感病的植株以后, 与取食未感病植株的种群相比, 生殖力提高了11-17倍, 成虫寿命延长了5-6倍, 经8周后, 感病植株上种群数量提高了2-13倍, 这些增加的个体同样都是携带病毒的。显然, 媒介昆虫B型烟粉虱的大量增加促进了病毒病的流行, 而病毒的流行又更加有利于B型烟粉虱的增殖, 从而使其种群能更迅速扩张(Jiu et al., 2007)。对紫茎泽兰的研究发现, 改变土壤细菌群落是其入侵过程的一个重要组成部分。土壤微生物群落可能在该外来植物与本地植物的关系中起到了重要的“桥梁”作用, 紫茎泽兰可以通过改变重度入侵地土壤微生物群落结构阻碍本地植物的生长和更新(Yu et al., 2005)。对加拿大一枝黄花(Solidago canad- ensis)的研究也发现, 与之共生的土著菌根真菌的多样性是促进其入侵的关键(Jin et al., 2004; Zhang et al., 2009)。根据对崇明东滩不同围垦历史的被入侵生态系统的观察, 加拿大一枝黄花的入侵时间影响了菌根真菌与植物的互生关系, 即该物种根际菌根真菌多样性与其入侵时间之间呈线性正相关, 而菌根真菌多样性又反作用于入侵的过程, 进一步推动了加拿大一枝黄花的繁殖和扩散(董梅等, 2006)。这一结论既有助于我们认识该外来植物成功入侵的机制, 也回答了生态学中的一个基本问题, 即土壤中菌根真菌群落如何随地上部分植物群落的演替而变化。此外, 植物和固氮菌之间的共生关系对外来种的入侵亦十分重要, 许多“臭名昭著”的入侵植物, 如互花米草(Liao et al., 2008a; 张骁栋, 2012)、紫茎泽兰(Niu et al., 2007; Zhao et al., 2010)等都是固氮“高手”, 这种能力可能都和它们与固氮菌共生有密切关系。固氮能力使这些外来植物在与土著植物的竞争中, 获得了更多的营养资源, 从而产生了更多的生物量。

值得注意的是, 在外来种与土著种的共生过程中, 两者之间的利益的关系并非全是互利的。在有些物种共生互作的过程中, 外来种往往在从土著种那里获得有利于自身繁衍和扩张利益的同时, 却压缩了土著种繁衍和增殖的空间, 这种现象是一种典型的“偏利共生”(commensalism)表现。这方面, 一个极好的实例是B型烟粉虱与土著烟粉虱之间存在的被称为“非对称交配互作”(asymmetric mating interaction)的关系(Liu et al., 2007)。当B型烟粉虱到达新的地区与土著烟粉虱(B. tabaci)共存后, 虽然它们之间并不能完成交配, 但相互间发生的一系列求偶行为及作用, 使B型烟粉虱的交配频率迅速增加, 卵子受精率提高, 后代雌性个体比例由独处时的约60%增加到70-80%, 种群增长加快。与此同时, B型烟粉虱雄虫又频频向土著烟粉虱雌虫求偶, 干扰土著烟粉虱雌雄之间的交配, 使后者交配频率下降, 后代雌性比由独处时的约50%下降到20-40%, 降低其种群增长。由于B型烟粉虱与土著烟粉虱之间的这种求偶互作是对一方有利而对另一方有害, 故称这种现象为“非对称交配互作”(Liu et al., 2007)。这一现象揭示了动物(尤其是单-双倍体动物)入侵的一个重要的行为机制, 这种机制是入侵者的一种重要内在潜能, 当入侵者到达新地域与土著近缘生物共存产生互作, 激发这一潜能并迅速发挥作用, 驱动其入侵和对土著生物的取代过程。

2.5 化感作用

外来植物传入一个新的生态系统以后, 逃避了原产地的生物控制, 在较低的天敌压力下重新分配其资源, 导致生长和防御等方面的能量和物质发生二次重构(Wilsey & Polley, 2006)。由于与土著植物缺乏长期的协同进化关系, 外来植物的到来使得演替时间较长的土著植物群落种群间业已构建的化感平衡随之被打破(万方浩等, 2011a)。外来植物分泌的化感物质往往是土著植物无法短期适应的新毒素, 可抑制土著植物的生长, 而通常对外来植物自身或其伴生生物却无害。因此, 新化感物质会成为迅速排挤土著植物的“利器”, 从而有助于入侵植物种群的快速建成, 由此产生了入侵植物的“新武器假说”(novel weapon hypothesis)(Callaway & Ride- nour, 2004)。

尽管对“新武器假说”目前仍没有系统的认识, 但已经发现, 在一些重要外来入侵种中, 这种假说能解释某些入侵现象背后的机制。化感作用促进入侵可采用基于同质园实验的生物地理比较法, 即通过以近缘甚至同一伴生受体植物为对象, 比较入侵植物在原产地和入侵地化感作用强弱的变化, 最终推测化感在该入侵植物扩张中的贡献。这方面, 国外的经典研究案例有矢车菊(Gerbera jamesonii)和葱芥(Alliaria petiolata)等。中国从2000年之后逐渐开展了有关研究, 如发现互花米草在上海九段沙湿地, 通过分泌化感物质抑制土著植物海三棱藨草(Scirpus mariqueter)种子的萌发, 从而实现入侵(Liang et al., 2012)。以紫茎泽兰为对象的研究, 在入侵植物化感作用方面从不同入侵程度(重度、中度、轻度)证实了该外来植物化感作用对自身种群和伴生植物的两面性(Niu et al., 2007); 有关从紫茎泽兰中分离的3种主要化感物质的作用研究表明, 化感作用是紫茎泽兰与土著植物种群竞争中获得优势的重要策略(Yang et al., 2006, 2008, 2011; Niu et al., 2007)。此外, 在薇甘菊(Mikania micrantha)(Shao et al., 2005)、胜红蓟(Ageratum conyzoides)(Kong et al., 1999)、飞机草(Eupatorium odoratum)(凌冰等, 2003)、一年蓬(Erigeron annuus)(方芳等, 2005)、蟛蜞菊(Wedelia chinensis)(聂呈荣等, 2002)、三裂蟛蜞菊(W. trilobata)(张玉虎等, 2004)、豚草(Ambrosia artemisiifolia)(张风娟等, 2010)和三裂叶豚草(A. trifida)(王朋, 2008)等植物中也均发现了化感物质的存在; 但遗憾的是, 由于缺乏进一步的研究, 这些有限的例子至多能说明相关物种有化感作用, 却很难说明化感作用与入侵性的关系。

事实上, 我们推测, 几乎所有的植物都会分泌化感物质, 甚至入侵植物在原产地也有化感作用。因此, 在研究化感作用对入侵成功的相对重要性时, 一个值得考虑的问题是, 入侵植物的化感物质是否在入侵后得到了增强, 化感物质分泌的增强是否有效地推动了入侵。与本地同属植物的化感作用比较发现, 喜旱莲子草与土著近缘种莲子草(A. sessilis)的化感作用几乎相当, 而后者在我国许多地区能与其他植物共存(余柳青等, 2007a)。这说明化感作用在入侵植物扩张过程中不一定得到了增强。那么, 对于成功入侵起作用的化感究竟需要多大量的他感物质才能有效地促进入侵, 乃至化感对入侵真正的实际作用, 可能还要结合其他的机制进行综合分析。同时, 考虑入侵植物化感作用的增强与否, 有必要结合其在原产地的表现进一步比较。

严格来讲, 入侵植物化感物质分泌的增强并非是对所有土著植物都能成为有“杀伤力”的“核武器”, 如在有些人为干扰小的稳定植物群落中, 土著植物可对新到来的外来植物分泌的化感物质产生超强的抵御力, 这也从另一个方面较好地解释了为什么稳定生态系统的抗入侵能力较强; 但这种现象究竟是因为未受干扰的生境中群落间化感平衡足够强大, 从而抵抗住了新化感的“进攻”, 降低了土著种对新化感的“敏感性”; 还是因为稳定的环境削弱了外来种的化感在新生境中的“进攻性”, 这一问题还需要深入研究。甚至, 一些入侵植物分泌的化感物质还有可能促进了土著植物的生长。例如, 在美国已发现, 乌桕(Sapoum sebiferum)入侵北美以后, 土著植物裂稃草(Schizachyrium scoparium)、黑柳(Salix nigra)和落羽杉(Taxodium distichum)在乌桕的化感作用下, 能够生长得更好(Keay et al., 2000; Conway et al., 2002)。Zhu等(2011)也发现, 紫茎泽兰分泌的化感物质只有在高浓度时才抑制其他植物的生长, 在低浓度时却能促进其他植物的生长; 而且, 土壤微生物可以降低紫茎泽兰化感物质的浓度, 起到“缓冲作用”。但这种情况中是否还存在化感作用之外的其他促进因素(如土壤营养增强、天敌减少等), 还有待进一步研究。

从已有外来种化感作用增强在成功入侵中的地位来看, 不同入侵种之间化感作用不一样, 其原因来自3个方面: (1)不同生态系统中, 土著植物对外来植物分泌的化感物质敏感性不同; (2)入侵植物产生的化感物质的量不一定比土著近缘种多; (3)入侵植物产生的化感物质在不同生境中的降解速度不一样(万方浩等, 2011a, c)。总地来看, 化感物质分泌强烈的植物主要集中在菊科、禾本科和十字花科中, 而这3个科恰好是中国入侵植物种类最多的分类群。因此, 对化感作用促进植物入侵的研究对象及其作用机制还值得深入拓展。

3 外来种入侵的生态学效应

外来种入侵的生态学效应深刻地影响着我们对外来种入侵后果的认识, 是制定管理决策的重要依据。可以说, 外来种入侵的生态学影响也是生态学家在本世纪所面临的又一项挑战(Pulliam & Yu, 2005)。近10年来, 我国学者针对境内生物入侵的形势, 从个体、种群、群落、生态系统等不同水平, 对其生态学效应进行了系统考量, 并集中研究了几种新近暴发的重要入侵种对湿地、农田、林地和水域等生态系统的影响, 发现和阐明了部分外来种入侵导致土著生态系统结构崩溃及功能丧失的生态学机制。相关成果为国家外来入侵种的管理与控制决策的制定提供了有力的依据。以下我们结合这些研究进展, 对中国外来种入侵的生态学效应进行概述。

3.1 对生物多样性、群落演替格局和生态系统服务功能的影响

近年来, 外来种入侵已严重影响到我国生物多样性资源, 局部改变了物种原有的空间分布格局, 甚至造成了一些土著种濒临灭绝。在生物入侵对生物多样性影响方面, 外来植物的入侵最为明显。例如, 加拿大一枝黄花的入侵导致上海地区多种土著物种局部消失(李博等, 2001)。互花米草入侵福建等地沿海滩涂, 导致红树林湿地生态系统遭到破坏, 红树林消失, 滩涂鱼虾蟹贝类以及其他生物生存环境改变, 原有的200多种生物减少到20多种(左平等, 2009)。互花米草入侵崇明东滩盐沼湿地后, 降低了土著植物芦苇和海三棱藨草的丰度(Wang et al., 2006), 甚至造成局部海三棱藨草的灭绝(Chen et al., 2004)。在崇明东滩, 该外来种的入侵, 打破了湿地中固有的自然植物群落的演替进程, 迅速向外排挤海三棱藨草群落, 加快了芦苇群落在高潮滩的定殖, 因而严重改变了湿地土著植被的空间分布格局(Li et al., 2009)。互花米草的入侵, 还改变了被入侵生境中根际微生物功能群的多样性(Wang et al., 2007; Nie et al., 2009), 从而影响了整个生态系统的营养循环过程(Chen et al., 2007a, 2012)。此外, 它的入侵虽然对昆虫多样性指数没有显著影响, 但改变了昆虫群落结构, 互花米草单种群落中昆虫丰富度显著低于芦苇单种群落, 一些土著昆虫还发生了食性转移(Wu et al., 2009)。喜旱莲子草在池塘、湖泊、水库、沟渠、小溪、河道等富营养化的淡水生态系统中形成优势种后, 通过改变边沿带物理环境影响了伴生生物群落的组成, 降低了本地植物多样性(Pan et al., 2006; 余柳青等, 2007b)。其毯状结构还降低了水流速度, 增加泥沙沉积, 加剧植株冠层蒸腾作用, 使水生生境陆生化, 从而改变植物群落的演替方向和分布格局(潘晓云等, 2007)。紫茎泽兰通过抑制土著植物种子萌发和幼苗生长(王紫娟等, 2007; 张学文等, 2007a, b; 曹子林等, 2012), 竞争排挤和取代土著植物, 形成的单种优势群落破坏或改变了土著植物格局, 引起了一系列的生态学问题和难题(丁晖等, 2007; Niu et al., 2007; Liu et al., 2010)。豚草形成单优种群以后, 在入侵生境中土著种的丰富度指数、多样性指数均显著降低(魏守辉等, 2006; 邓旭等, 2010)。水葫芦入侵云南滇池后占据了草海湖2/3的水域面积, 导致鱼类数量锐减, 蚊蝇等病原菌媒介昆虫繁殖面积扩大, 水体中植物种类从1960年的16种减少到1990年的3种(Ding et al., 2001)。

与入侵植物相类似, 一些杂食性外来动物的入侵, 对土著生态系统动物多样性亦构成了一定威胁。例如, 红火蚁入侵我国以后, 捕食许多土著捕食性蚂蚁, 导致生态系统捕食者结构简单化, 土著蚂蚁种类和数量明显减少, 在红火蚁发生严重的深圳地区, 红火蚁正逐渐取代土著蚂蚁成为当地的优势种, 被入侵草坪和荒草地中土著蚂蚁的种类分别减少了6种和3种(吴碧球等, 2008; 宋侦东等, 2010); 被入侵的豆田中, 土著蚂蚁的种群密度降为原来的1/5(Huang et al., 2010)。克氏原螯虾(Procambarus clarkii)通过捕食本地水生动物, 对长江流域重要水产品中华绒螯蟹(Eriocheir sinensis)和青虾(Macrob- rachim nipponense)具有极大的杀伤力(武正军等, 2008); 还减少了本地其他水生动物的生存空间, 使斗鱼(Macropodus opercularis)、鳑鲏(Rhodeus sinensis)等野生鱼类消失(江舒等, 2007)。

有趣的是, 有些植食性动物入侵以后, 土著生态系统的生物多样性并不像植物入侵那样一直降低。譬如, 马尾松(Pinus massoniana)纯林和混交林被松材线虫危害后, 原有森林群落均未向灌丛方向退化, 生物多样性反而比危害前有较大幅度增加, 这说明松材线虫危害后, 随着时间的推移和植被的恢复, 整个群落能够朝更高、更稳定的方向演替和发展(吴蓉等, 2005)。松材线虫入侵加速松林演替过程中植被结构的变化也对林内鸟类多度和丰富度波动产生了影响, 波动的趋势是: 随着群落演替阶段的推进, 鸟类多样性先增加, 到演替顶极期下降, 但在随后更长时间内又会出现小的增长, 而后又下降(蒋科毅等, 2005)。

外来种入侵后, 破坏了土著生态系统的结构和功能, 使生态系统服务功能下降, 带来了一系列严重后果。例如, 长江口湿地是东亚-澳大利亚候鸟迁徙途径的重要停歇点, 互花米草入侵该湿地替代土著植被后, 改变了生态系统的食物链结构, 降低了湿地对候鸟保育的生态服务功能(Li et al., 2009), 导致鸟类栖息地丧失或破碎化(Jing et al., 2007), 目前互花米草群落中鸟的种类和数量都明显下降, 鸟类群落的结构发生了变化(Ma et al., 2009)。喜旱莲子草根系发达、地上部分繁茂, 通过竞争作用影响农作物、园艺作物和绿化植物的生长(喻大昭等, 2008), 影响局域生态系统的耕作服务功能和种植价值。水葫芦入侵后影响了生境水体质量, 改变了生态系统群落组成及河道景观, 降低了水体的生态净化功能和景观服务价值(高雷和李博, 2004)。

值得强调的是, 外来种入侵对生态系统服务功能的影响最值得关注的领域来自森林生态系统。森林在陆地生态系统中承担着如生物多样性维持、防风固沙、涵养水源、碳固定、保育土壤、营养积累、释放O2和净化大气环境等重要的生态服务功能。森林生态系统在全球碳循环中承担重要角色, 在温室气体排放中, 森林碳排放和吸收强烈影响生态系统的持续管理。我国目前森林外来种入侵形势严峻, 由此带来的生态影响巨大。以松材线虫为例, 根据国家林业局公布的资料, 该线虫可寄生57种松属植物, 目前其疫区已扩展到全国14个省份的192个县市, 累计致死松树5亿多株, 毁灭松林33万余ha。松树是我国重要的绿化造林树种, 总面积达3,300多万 ha, 每年可吸收CO2 1.2亿吨, 释放O2 7千万吨, 贮碳量2.3亿吨, 对维护国家生态安全和缓解气候变暖至关重要(贾治邦, 2009) (贾治邦 (2009) 贾治邦在全国松材线虫病防治工作会议上的讲话: 认清严峻形势, 采取有力措施, 坚决遏制松材线虫病扩散蔓延的势头. 2009年4月21日, 安徽黄山.)。如果松材线虫病继续扩散蔓延, 将对松林生态系统的生态服务功能造成毁灭性影响。

3.2 对生态系统化学性质和生物地球化学循环的影响

外来种入侵导致生境的化学性质发生改变, 又反作用于外来种与土著种的竞争, 促进生物入侵的进一步发生, 形成了入侵“正反馈”效应(Liao et al., 2008b)。如互花米草能截留海水中的硫元素, 提高单位面积植物组织的硫储量(Zhou et al., 2009), 其凋落物进入土壤增加了硫酸盐还原反应的底物, 提高了底泥中的溶氧度、氧化还原电位及硫化物的氧化度, 这些变化可能抑制了土著植物的生长, 进而促进了互花米草单物种群落的形成(陈中义等, 2005)。

外来种入侵可改变系统的生物地球化学循环过程。例如, 互花米草比海三棱藨草和芦苇具有更高的初级生产力, 其入侵后向生态系统输入更多的凋落物, 而这些凋落物的分解速率低于土著种, 因此增加了生态系统的碳库(Cheng et al., 2006; Liao et al., 2007), 该植物入侵后激活了土壤中固氮生物的活性, 显著增加了土壤氮浓度和氮库(Liao et al., 2008a), 进而影响了生态系统的碳氮循环。紫茎泽兰通过改变土壤微生物群落结构、改善土壤养分、增加入侵地土壤固氮菌的数量, 影响了生态系统的氮循环过程和速率(Xu et al., 2012)。

有些外来种入侵后, 土壤营养资源被极大消耗, 由此造成了土壤的可耕性受到破坏。例如, 豚草对土壤中水分和氮、磷消耗极大, 可造成土壤干旱贫瘠, 影响农作物的生长(常望霓等, 2011)。松材线虫病危害松林后, 土壤磷、全氮和有机质等营养元素均不同程度地减少, 影响到松林的生长和扩繁速度(黎镇湘等, 2005)。另外, 有些外来植物入侵后, 虽然改善了土壤的营养结构, 但这种改善反而更加有利于自身的生长。例如, 紫茎泽兰入侵后, 改变了入侵地土壤微生物群落结构和功能, 增加了土壤功能菌群数量、酶活力和土壤肥力, 从而更有利于自身生长, 这种“正反馈效应”形成了紫茎泽兰的一个自我促进式的入侵模式(Niu et al., 2007; 戴莲等, 2012)。

3.3 其他影响

外来种入侵对经济活动的最大危害, 在于其对农、林、牧、渔业生产、交通航运等带来的直接破坏, 并由此造成了巨额的经济损失。据测算, 我国283种主要入侵种每年带来的直接和间接经济损失达144.5亿美元(Xu et al., 2006)。例如, 紫茎泽兰入侵后, 侵占大量农田、林地和草地, 不仅影响农林生产, 造成牧草减产甚至丧失放牧利用价值; 还因该植物富含有毒物质, 牲畜误食后引起中毒或死亡, 由此导致中国畜牧业损失达9.89亿元/年(Zhu et al., 2005)。水葫芦、喜旱莲子草等水生植物的肆意扩张, 覆盖湖泊和河流的广大区域, 减缓水流, 堵塞航道, 影响水上交通、灌溉和水电设施, 造成了严重的经济损失, 在2000年中国生物入侵造成的经济损失中, 水葫芦入侵占到了14-17%(杨凤辉等, 2002), 仅每年用于其人工打捞的费用就高达1亿元(Ding & Chu, 2005)。

相对于直接危害, 外来种入侵带来的间接危害则更加巨大。例如, 一些媒介昆虫的入侵, 间接造成了农作物病害的大暴发。Q型烟粉虱最近几年在我国多数省份成为棉花、蔬菜等农作物上的优势害虫(Chu et al., 2010a, b; Teng et al., 2010), 由其引起的番茄黄化曲叶病毒(TYLCV)在我国广大地区暴发成灾(Pan et al., 2011; Pan et al., 2012), 导致农作物大面积受害。松材线虫的入侵使其他有害生物易于从伤口感染松树, 间接造成了松树的大面积死亡(Zhao et al., 2003)。克氏原螯虾携带的病原体, 造成中华绒螯蟹和青虾发生疾病的交叉感染, 引起大规模的水产损失(Wang et al., 2005)。红火蚁的入侵使藿香蓟(Ageratum conyzoides)的密度明显增大(Huang et al., 2011); 而藿香蓟也是一种入侵植物, 由此形成了外来昆虫和外来植物的协同入侵产生复合损失。

外来种入侵除了导致严重的经济损失外, 关于其对人类健康、环境和公共设施安全带来的威胁也值得重视。例如, 豚草可产生大量的花粉, 引起过敏和哮喘等并发症, 在我国部分地区引发了居民的呼吸道疾病(Liang et al., 2010)。红火蚁通过捕食、筑巢等行为, 改变了土壤生态系统的结构和功能, 使土壤环境恶化; 叮咬人、畜, 造成重大健康隐患; 啃啮植物, 造成农业减产; 破坏公共设施, 造成公共财产损失和安全事故发生(曾玲等, 2005a, b)。克氏原螯虾在淡水江湖大坝周围, 一旦种群密度过高, 其所挖掘的洞穴可引起大坝漏水, 造成溃堤风险, 带来巨大的安全隐患(江舒等, 2007)。

4 主要入侵种的控制与管理

生物入侵之所以深受全球广泛重视, 不仅仅是因为它为科学家提供了研究生态学和进化问题的理想模式, 更为重要的是, 它正侵蚀着我们赖以生存的自然和人工生态系统。事实上, 从应用层面来看, 无论我们对生物入侵的研究如何深入, 其最终的成果都要面向实际应用, 所以有关入侵种的控制与管理显得尤为重要(万方浩等, 2008b; 李博和马克平, 2010)。也正由于此, 能否将现有的生物入侵研究成果转化成公众的知识、管理行动或政府决策的依据, 并最终找到对付其的方法, 是公众对生态学家的“完美期待”。因此, 有关生物入侵管理与控制的途径与方法成为联系生态学家、政策制定者和社会公众等各类群体间的转化生态学(translational ecology)命题(Schlesinger, 2010)。

为了应对日益严重的生物入侵形势, 中国一方面从政府管理入手, 强化了检验检疫、预警监测、应急处置等管理体系的建设, 如农业部、环保部、国家质检总局和国家林业局均设立有专门管理外来种入侵的机构。农业部针对全国85种重要农业入侵物种, 制定了预警监测和应急防控指南(张国良等, 2008); 环保部分别于2003年和2010年先后分两批发布了共36种入侵物种的清单; 国家质检总局定期发布口岸外来种入侵预警警报; 国家林业局制定了林业入侵物种应急控制预案。同时, 这些部门联手牵头, 组织全国不同领域专家, 正着手起草国家层面上的生物入侵中长期管理与防控计划及政策指南(万方浩等, 2008b)。另一方面, 我国科技界针对农林和湿地生态系统中重要入侵种, 着力加强了包括数据库、检测监测、风险预警、生物防治、扩散阻断等技术体系研究。这些工作主要包括: 对70余种重要入侵物种的适生区进行了定量风险预测(万方浩等, 2010), 开展了20种重大入侵物种和潜在入侵物种的快速检测分子基础、技术和方法研究, 并开发了多种快速检测与野外监测的试剂盒; 同时, 与管理部门联手, 对多种已形成严重危害的入侵种, 开展了根除和治理的统一行动, 相关成果为我国组建生物入侵的早期预警与狙击、应急控制、阻断与扑灭、可持续综合防御与控制等四大技术体系提供了方法和依据(万方浩等, 2011b)。下面我们以案例形式, 简要回顾我国一些重大入侵种管理与控制技术的进展。

(1)互花米草: 互花米草是我国东海岸盐沼中危害最为严重的外来入侵植物, 其面积从1985年的仅260 ha迅速扩张至2000年的11.2万 ha (An et al., 2007), 长江河口盐沼是该物种入侵的重灾区(Wang et al., 2006; Li et al., 2009)。为此, 2006年起, 上海市绿化和市容管理局(上海市林业局)与复旦大学等单位一直致力于互花米草在崇明东滩湿地的入侵控制, 提出了刈割、水文调控以及生物替代组成的综合控制途径。就刈割而言, 在花期对高潮滩和低潮滩中的互花米草分别进行4次和3次刈割, 可以取得有效的控制效果(Tang et al., 2010), 尤其是扬花期刈割能有效控制互花米草的有性繁殖和扩散(肖德荣等, 2011)。整个控制工程, 实际上包括了“围、割、淹、晒、种、调”6个过程, 几年的中试取得了很好的效果; 目前正在上海崇明东滩实施互花米草控制和鸟类栖息地恢复生态工程。上海市政府计划在崇明东滩投资近10亿元, 用于互花米草的持续治理, 其中, 围堰建设是六字互花米草治理技术方案中的关键工程, 2012年, 二期项目中的围堰总长已达4,000 m, 围区面积133.3 ha。

(2)喜旱莲子草: 机械或人工打捞、拔除等手段是控制喜旱莲子草的常用方法, 但这些物理方法的使用耗费的人力太大(Pan et al., 2009)。为了控制水生生境中的喜旱莲子草, 中国于1986年从南美洲引进莲草直胸跳甲(Agasicel hygrophila)开展以虫防草获得成功, 目前, 在福州、海口、温州和湘潭4个基地每年可生产这种跳甲8,805万到9,525万头, 利用该天敌对湖南岳阳和江永、湖北武昌、江西南昌、广东韶关、广西来宾等地的喜旱莲子草控制效率达47%以上(周忠实等, 2011a); 但有学者认为, 这种甲虫对陆生生境喜旱莲子草防效甚微, 这可能与陆生型喜旱莲子草茎木质化程度高导致跳甲化蛹困难(马瑞燕和王韧, 2004)、茎组织质量密度大不利于跳甲发育等有关(Pan et al., 2011)。刈割可能是陆生型喜旱莲子草的一种有效的人工控制方法; 但使用该法时, 刈割需频繁持久, 在刈割过程中, 应避免喜旱莲子草根的破碎化, 万一出现碎根, 应及时清除出土壤并焚烧, 以减少营养繁殖体库(Jia et al., 2009)。此外, 在利用真菌感染除草的研究中, 已筛选出喜旱莲子草的高效生防菌蕉斑镰刀菌(Fusa- rium stoveri)的一些有效菌株, 但野外大规模应用效果仍有待评估(庄义庆, 2009)。

(3)紫茎泽兰: 紫茎泽兰入侵成功与否与入侵地生态系统的异质性相关, 受干扰的生境更易被紫茎泽兰入侵, 而稳定生态系统对紫茎泽兰的抵御能力较强, 因此恢复和保护土著生境的多样性是抵御紫茎泽兰入侵和扩张的有效途径之一(Wan et al., 2010)。紫茎泽兰具有明显的先锋植物特性, 极易形成单物种群落, 且土壤种子库持久, 一旦形成单优势群落, 进行植物替代的难度较大, 连根铲除后, 再利用生境植物替代和增加植物多样性的方法可防止其二次入侵(朱宏伟等, 2007; 赵林等, 2008)。在生物防治上, 从原产地引进天敌昆虫泽兰实蝇(Procecidochares utilis)和其他草食性昆虫对紫茎泽兰的生长有一定的控制作用, 但由于种种原因, 目前控草效果仍然有限(王文琪等, 2006)。在生防菌的筛选上, 我国现已分离到19个链格孢菌属(Alternaria spp.)的菌株, 其菌丝具有较强的致病力, 通过产生广谱触杀性毒素细交链孢菌酮酸(AAC-Toxin)可引起紫茎泽兰感染褐斑病, 应用前景广阔(Chen et al., 2007b)。最近, Niu等(2010)报道了紫茎泽兰的一些新天敌。他们发现, 东方行军蚁(Dorylus orientalis)能啃食紫茎泽兰根茎部表皮、皮层、韧皮部, 甚至木质部, 影响紫茎泽兰地上、地下器官物质和能量的交换, 短时间内就能导致大量植株死亡; 更为重要的是, 这种蚂蚁对紫茎泽兰还具有一定程度的选择性, 这一进展可能为我们开发紫茎泽兰新的生物防治替代品提供了重要基础。

(4)豚草: 2003年, 中国提出了“南紫北豚”的治理策略和口号, “南紫”为上面提到的紫茎泽兰, “北豚”即豚草。基于此需求, 我国农业科学家提出了豚草的生物防治、替代控制、化学防治一体化综合治理技术体系, 开创了国内入侵杂草综合治理的先河, 目前该体系已逐步在全国推广, 其中天敌昆虫的工厂化繁殖和释放已在湖南等南方省区取得成功(万方浩等, 2008a)。尤其值得强调的是, 有关豚草的生物防除技术是我国在杂草生防中获得的最好代表性成果之一, 其主要手段是从原产地引进专一性天敌和植物病原菌开展生物防治, 1987年开始, 我国先后引进了豚草条纹叶甲(Zygogramma suturalis)和豚草卷蛾(Epiblema strenuana)等天敌昆虫, 发现他们对豚草的生防效果较理想, 尤其是后者目前已在湖南、湖北和江苏等省成功定殖, 对豚草的控制起到了积极作用(万方浩等, 2008a)。有趣的是, 近年来, 我国又新发现了1种非有意引进的天敌昆虫——豚草广聚萤叶甲(Ophraella communa)也极具生防潜力(胡亚鹏和孟玲, 2007), 目前已对其控害有效性与安全性评估(Cao et al., 2011; Guo et al., 2011; Zhou et al., 2011)、生物生态学特性及扩繁技术(Zhou et al., 2010a, b, 2012a, b; Zhu et al., 2012)等进行了一系列研究。通过广聚萤叶甲和豚草卷蛾的联合控制, 在广西来宾和湖南汨罗释放区内, 豚草植株死亡率达98%以上, 且天敌迅速向邻近豚草发生区扩散, 两种天敌对扩散区豚草的控制率可达50%以上, 该技术已达到大面积推广应用的要求(周忠实等, 2011b, c)。在生防微生物的筛选上, 2005年以前, 我国就先后发现了6种真菌可感染豚草, 但其中只有少数病原菌严重发生时才能使豚草致死(万方浩等, 2008a)。最近新发现三裂叶豚草锈菌(Puccinia xanthii f. sp. ambrosiae-trifidae)侵染宿主三裂叶豚草的叶片, 可破坏细胞结构, 导致细胞内膜系统紊乱, 降低细胞器结构稳定性(曲波等, 2011), 但此病菌对普通豚草是否能构成有效的致病效果仍有待进一步研究。

(5)水葫芦: 目前全世界控制水葫芦的方法主要有通过人工或机械物理去除, 施用除草剂, 利用天敌和病原菌进行生物防治等3种方法。在中国, 物理去除和施用除草剂都不能起到良好效果(高雷和李博, 2004; Lu et al., 2007a)。考虑到生物防治可能是控制水葫芦的最有效的手段, 我国于1995年从南美引进了两种象虫(Neochetina eichhorniaeN. bruchi)用于水葫芦的治理(Ding et al., 2001), 取得了一定的成效, 但是要达到有效治理的目的不得不每年引入天敌(Lu et al., 2007a)。此外, 对入侵植物的开发与综合利用是对其管理的一个补充性措施, 研究表明, 水葫芦能有效吸收水体中的磷, 提高对铵态氮的去除效率(Chen et al., 2010b), 促进富营养化水体的硝化、反硝化过程(高岩等, 2012), 因此, 利用水葫芦来治理水体富营养化和污染可以起到一定的控制效果。不过这种利用方式只能在严格限制的条件下进行, 否则二次入侵在所难免(Pan et al., 2011)。

(6)椰心叶甲: 针对椰心叶甲在海南省严重危害的局面, 中国热带农科院于2004年从越南引进了椰甲截脉姬小蜂(Asecodes hispinarum)及椰心叶甲啮小蜂(Tetrastichus brontispae), 经过安全性评估后(et al., 2008), 目前已建立了这两种寄生蜂的生产基地, 日产蜂达190万头。此外, 还筛选并改良了椰心叶甲高效生防绿僵菌(Metarhizium spp.)2种和Bt菌株1种, 以此开发了3种植物源杀虫剂, 均已进入生产流程。在系统评价高效绿僵菌菌株对同域节肢动物类群的安全性基础上, 着重研究了Bt生防制剂、寄生蜂与绿僵菌的增效作用。在此基础上, 建立了寄生蜂与化学农药、绿僵菌及寄生蜂的协同应用技术体系, 该体系已在海南、广东、广西等地示范推广, 累计放蜂10亿头, 有效控制面积达2.6万 ha以上, 有效压缩了椰心叶甲的危害与扩张势头(彭正强等, 2008)。

(7)松材线虫: 松材线虫病的持续治理是国家林业局林业有害生物防治工作中重中之重的内容, 我国对松材线虫病的持续治理已近30年, 2002年国务院办公厅发布了《关于进一步加强松材线虫病预防和除治工作的通知》, 这是我国唯一受国务院重视的单个入侵种的治理文件。在国家的高度重视下, 我国学者在松材线虫的检测、检疫技术方面获得了重要进展, 以南京林业大学森林保护科研人员为主的团队开发了基于生物化学和分子生物学的一系列新型检测工具、技术和试剂盒, 与传统技术相比, 这些技术大大提高了检疫效率和准确性(陈凤毛等, 2007a, b)。在防治上, 我们目前应用的方法主要集中在化学防治、物理防治、生物防治、基因控制及生态控制等方面; 但总体上仍以化学防治为主。在生物防治方面虽然发现管氏肿腿蜂(Scleroderma guani)、花绒坚甲(Dastarcus longulus)和球孢白僵菌(Beauveria bassiana)等天敌昆虫及病原微生物对松材线虫传播媒介昆虫松褐天牛(Monochamus alternatus)具有一定的控制效果, 但大面积推广难度较大。在生态控制方面, 我国学者基于松材线虫的基础研究成果(Xie et al., 2009), 提出了通过生态系统的改造, 构建系统优化结构, 加强林分抚育, 增强树势, 提高系统自我控制松材线虫病的功能的思路; 据此利用乡土阔叶树种与松树的混交造林和林分改造手段达到了分散松材线虫的危害并抵御松材线虫自然传播的目的(张锴等, 2010)。总之, 我国松材线虫病的治理已取得明显的成效, 连续实现松材线虫病发生面积和病死树数量的“双下降”, 发生面积减少到6.7万 ha以内, 病死树数量控制在110万株以下(贾治邦, 2009) (贾治邦 (2009) 贾治邦在全国松材线虫病防治工作会议上的讲话: 认清严峻形势, 采取有力措施, 坚决遏制松材线虫病扩散蔓延的势头. 2009年4月21日, 安徽黄山.)。

(8)美国白蛾: 美国白蛾是我国重大检疫性入侵害虫, 对林业和园林绿化植物具有重大危害。我国林业科技人员从1984年开始, 结合前苏联等国家从美国白蛾原产地引进天敌进行生物防治没能成功的经验和教训, 立足调查和寻找本土天敌。经过21年的努力, 发现和筛选出美国白蛾的特优天敌——白蛾周氏啮小蜂, 基于对这种小蜂的行为学、生态学、生物学、解剖学及人工大量繁殖、放蜂防治技术的长期研究, 筛选出人工大量繁殖小蜂的替代寄主, 解决了小蜂大量繁殖的瓶颈问题; 通过放蜂防治试验, 取得了良好的防治效果(魏建荣等, 2004)。同时, 筛选出防治美国白蛾的高毒力核型多角体病毒(HcNPV)毒株, 并攻克了该病毒大量扩增和常年生产的技术难题, 做到了病毒的常年规模化生产, 保证大面积生产防治的需要(段彦丽等, 2009)。在这些基础上, 提出天敌昆虫和病原微生物的综合利用模式: 在美国白蛾幼虫期喷洒HcNPV病毒, 蛹期释放白蛾周氏啮小蜂, 达到既控制美国白蛾当代的危害, 又有效抑制其下代的种群数量, 取得长期持续控制美国白蛾的防治效果。以该项技术为主, 辅助监测、预测预报和适当的化学防治, 结合群防群治、联防联治的防控策略使用, 已完全控制了上海、大连、烟台和青岛的美国白蛾数量, 特别是上海利用该生物防治技术防治后, 已经连续11年没有再发现美国白蛾。由于该项生物防治技术利用的是土著天敌昆虫和病毒控害, 因而做到了既消灭害虫, 又保证了生态环境和人畜的安全, 也保护了生物多样性(杨忠岐和张永安, 2007)。

(9)马铃薯甲虫(Leptinotarsa decemlineata): 1993年5月, 马铃薯甲虫疫情在新疆突然暴发, 农业部、新疆有关部门以及中国科学院动物研究所等科研单位, 构建了“捕、诱、毒、饿、治”为一体的马铃薯甲虫封锁与控制技术体系, 实施了对马铃薯甲虫的应急封锁和长期控制。在明确了我国马铃薯甲虫发生区域以后, 科研人员人工合成了马铃薯甲虫的聚集信息素; 筛选出高引诱活性的寄主植物挥发物; 创制了马铃薯甲虫越冬地地膜覆盖技术; 提出了利用一年生天仙子(Hyoscyamus niger)作为诱集带的成虫消灭技术; 筛选并制定了有效的化学药剂封锁控制技术和大规模人工应急灭虫技术; 研发出具有自主知识产权的转抗虫基因的马铃薯(Solanum tuberosum)植株或品系。推广过程中发现, 秋翻可使马铃薯甲虫越冬死亡率达33.3-76.4%, 轮作可降低虫口基数29.4-84.3%, 推迟成虫发生期4.7 d; 筛选出的14种高效的环境友好药剂, 使用20 d后田间防效均达90.0%以上。在以上技术研究的基础上, 制定的马铃薯甲虫监测防控技术体系在我国马铃薯主要产区广泛推广, 从而有效控制了马铃薯甲虫的继续扩散(郭文超等, 2011)。法国马铃薯甲虫权威研究人士Jolivet(1991)曾预测, 该虫最迟将在2003年穿越中国, 到达日本海和北朝鲜的沿海地区。但事实上, 目前马铃薯甲虫疫情仍然被封锁和控制在新疆局部地区, 扩散前沿为新疆木垒县(张润志等, 2010)。

(10)苹果蠹蛾(Carpocapsa pomonella): 苹果蠹蛾1953年首次在中国新疆报道发现, 目前已入侵新疆、甘肃、宁夏、内蒙古、黑龙江和吉林6省(自治区)。2004年, 农业部开始在西北黄土高原、环渤海湾2个中国苹果优势产区建设总面积达133.3万 ha的苹果非疫区, 苹果蠹蛾的入侵严重威胁到这两大苹果非疫区的建设, 并对全国苹果产业的安全生产构成直接威胁(徐婧等, 2012)。中国科学院动物研究所联合各级农技推广部门在深入了解苹果蠹蛾繁殖、发育和重要行为过程的基础上, 重点研究并突破了该害虫的监测、检疫和无公害防治等关键技术, 集成了农业防治、化学防治、生物防治等技术(病毒、赤眼蜂等), 并开展了大范围推广应用, 为我国苹果、梨等重要水果的健康发展和消除以苹果蠹蛾为技术壁垒的国际贸易障碍提供了重要技术支撑(张润志等, 2012)。在监测技术方面, 我国自主研发了高效的苹果蠹蛾诱芯和板式、盆式、瓶式诱捕器(石磊等, 2009; 朱虹昱等, 2012a); 发现黑光灯极具监测与防治苹果蠹蛾的潜力(张润志等, 2012)。在防治技术方面, 提出了“消灭越冬代, 重防第1、第2代, 控制第3、第4代”的防治策略(王兰等, 2011)。发现化学防治、迷向防治等措施对苹果蠹蛾成虫发生期的控制效果较好(徐婧等, 2012a, b); 初步证实国产迷向制剂对苹果蠹蛾具有较好防治效果(朱虹昱等, 2012b); 发现生物药剂甲维盐和高抗Bt联合应用具有速效性、持久性、环境友好等特征, 可作为苹果蠹蛾防治的首选药剂(杨建强等, 2011); 发现苹果蠹蛾颗粒体病毒张掖株(CypoGV-zy)具有作为生物农药工厂化生产的潜力(郑春寒等, 2011), 苹果蠹蛾颗粒体病毒株CpGV-CJ01并可作为延缓世界范围内CpGV抗性的候选菌株(申建茹等, 2012); 利用赤眼蜂防治苹果蠹蛾杏园蛀果减退率可达64%以上(王兰等, 2011)。以上技术从2010年开始, 在陕西、内蒙古、甘肃、新疆等省区进行了规模化示范, 累计设置监测点4,746个, 覆盖面积近22万 ha, 涉及主要高速公路、果品市场、果园等重点区域; 尤其在甘肃7市23个县(区)苹果产区开展示范后, 苹果蠹蛾发生面积较2006年减少了28%, 平均蛀果率减少了95%, 平均被害株率减少了93%(张润志等, 2012)。根据这些经验, 全国农技推广服务中心建立了全国范围内的苹果蠹蛾应急反应体系及非疫区管理制度, 进一步加强了公众宣传以及提高苹果非疫区外围的检疫和防治力度。以上措施成功阻止了苹果蠹蛾这一重大入侵害虫在我国的进一步扩散蔓延(王福祥等, 2012)。

(11)桔小实蝇(Bactrocera dorsalis): 桔小实蝇是华南、华东和西南地区的重要入侵害虫, 它的大面积暴发, 严重影响了我国南方水果安全生产和出口贸易。华南农业大学在该虫的检疫监测、预报预警和种群控制等方面开展了一系列研究, 在多方面充实了桔小实蝇持续控制的基础科学理论, 解决了多项决策和防治中的关键技术问题, 创建了种群持续控制技术体系。相关成果在我国南方的大范围应用, 有效遏制了桔小实蝇的猖獗危害, 显著提升了我国对桔小实蝇的整体防控水平和对重大外来实蝇的应急防控能力, 保护了中国南方果蔬的安全生产与贸易, 为保障我国的食品安全与生态安全做出了重大贡献(林进添, 2003; 任璐等, 2008)。在生物防治方面, 福建农林大学近年陆续引进或培养出阿里山潜蝇茧蜂(Fopius arisanus)、布氏潜蝇茧蜂(Biosteres vandenboschi)、长尾潜蝇茧蜂(Diachasm- imorpha longicaudata)和切割潜蝇茧蜂(Opius incisi)等寄生蜂种群。寄主专一性检测表明, 这些寄生蜂只寄生实蝇科害虫, 与本地寄生蜂能形成生态位互补, 对桔小实蝇的种群控制效果较好, 目前其工厂化繁殖技术基本成熟, 已进入扩繁阶段(梁光红等, 2006; 李夕英等, 2012)。

(12)红火蚁: 红火蚁的防治需依靠大范围的统防统治才能达到效果。目前对该害虫的治理主要在监控的基础上以实施有效的药剂灭除为主。红火蚁传入台湾后, 为防止疫情在岛内的迅速扩展, 台湾当局集合顶尖昆虫专家, 于2004年11月成立了跨部门的红火蚁防治中心, 先后举办100余次防治宣传讲习班, 设置覆盖北台湾的近90,000个监测点, 开展了以化学防治为主的红火蚁扑灭行动。通过各界民众9年来的共同努力, 目前台湾的红火蚁疫情得到有效控制, 种群密度已下降到民众可接受范围内(石正人等, 2007)。大陆2004年发现红火蚁疫情以后, 农业部迅速启动应急预案, 组织力量开展全国性普查, 建立监测点2,553个, 制定的一系列红火蚁防控技术在发生区进行了扑灭治理示范。我国植保技术人员创制了多个监测与控制的新方法、新装置, 建立了高效的检疫、监测和除害技术体系(曾玲等, 2005b; Zhang et al., 2007), 先后截获红火蚁628批次, 检疫灭除率100%, 有效遏制了红火蚁的危害。

(13)烟粉虱: 我国农业部门对烟粉虱的入侵治理一直高度重视。基于“隔离、净苗、诱捕、寄生和调控”为核心的烟粉虱可持续控制技术体系目前已在我国设施农业蔬菜主产区广泛应用, 此举可减少70%的杀虫剂使用量(张友军等, 2011); 针对粉虱类害虫在我国暴发危害的现状, 我国农业科技人员在华南、华东、华北、华中和西南等地区开展了粉虱类害虫的监测调查和预警防控技术研究的集成与示范。发现阿维菌素和烯啶虫胺是目前华北地区防治烟粉虱较为理想的药剂, 而该虫对联苯菊酯、噻虫嗪、啶虫脒和吡虫啉等药剂均产生了不同程度的抗药性(Qiu et al., 2009)。科研人员还研究了苘麻诱集、黄板诱杀、天敌昆虫和病原微生物对粉虱类害虫的控制作用, 并在华南、华东、华北、华中和西南等地区建立了12个示范基地, 分别在不同地区开展了粉虱类害虫综合防治技术的示范和推广, 累计印刷技术宣传手册12.4万份, 培训各类农民和基层农技人员5.43万人次, 累计示范面积4,300 ha, 技术辐射累计超过40,000 ha(任顺祥等, 2011)。在生物防治中, 我国科研人员发现了多种本地天敌种类及优势寄生蜂(Li et al., 2011), 建立了以甘蓝(Brassica oleracea)为寄主的烟粉虱优势寄生蜂规模化生产技术, 但其田间大范围推广仍存在许多技术问题有待攻关。值得注意的是, 最近几年来, 由于我国农区烟粉虱存在不同入侵生物型(如B与Q), 同时优势生物型间存在更替, 这为烟粉虱的综合治理提出了新的要求, 如Q型烟粉虱往往比B型烟粉虱对许多杀虫剂(尤其是新烟碱类杀虫剂)具有更强的抗药性(Luo et al., 2010; Yuan et al., 2012)。因此, 烟粉虱的管理与控制要注意不同生物型的抗药性监测, 及时研究和调整控制技术。

5 研究展望

生物入侵已成为全球广泛关注的重大问题; 然而, 我们对其深层次的认识还处于初期, 尚存在诸多悬而未决的问题, 譬如, 如何科学地预见生物入侵(Williamson, 1999)、物种的入侵与进化的关系(Allendorf & Lundquist, 2003)、生物入侵与全球变化其他过程之间的联系(Perrings et al., 2010)等。现代科学技术的发展, 将为我们拨开这些“重重迷雾”提供技术手段和思路。可以预见, 随着入侵生物学与其他多学科的深度融合, 在不久的将来, 我们将在生物入侵的机制、影响与控制等领域取得更多令人振奋的成就, 从而进一步促进人类对生物入侵问题的更深理解, 并最终寻找到科学的预测方法以及预防和治理的对策和措施。结合目前现状, 我们建议, 近期中国的生物入侵研究应在以下几方面重点 推进:

(1)生物入侵全境性科学考察: 我国环境保护部门分别于2001-2003年、2008-2010年先后两批对全国入侵种进行了普查(丁晖等, 2011), 农业、林业部门也于2006年首先在广东、福建、浙江、重庆和海南等5省(市)启动了第一批区域性入侵种的普查工作。综合这些调查结果的物种编目表明, 目前入侵我国的外来种已多达520余种(万方浩等, 2009)。与国土面积、生境类型等相近的国家相比, 中国目前的入侵种数量似乎并不多(Weber et al., 2008)。出现这种情况的一个很大的可能是, 我们目前对中国境内究竟有多少入侵种的认识仍是模糊的, 这就给依据管理(evidence-based management)带来了困难(李博和马克平, 2010)。因此, 关于外来入侵种的全境性科学考察和安全性评估仍是当前一项迫切需要开展的工作。事实上, 入侵种的本底调查是进行其管理和控制的第一步, 也是最重要的一步。基于目前的条线管理现状, 我们建议将原先分散在环保、农业、林业、渔业、海洋等各个领域的科学家及其资源进行整合, 在已有工作基础上, 以农业生态系统、林业生态系统及自然保护区、草原生态系统、淡水及海洋生态系统为对象, 延长调查的时间尺度, 扩展调查的空间尺度, 进行一次跨部门、跨学科、跨区域的全境性入侵种综合考察, 全面、系统、科学地阐明中国34个省级行政区内的入侵种多样性及其影响现状。基于新的调查结果, 进一步对本底资料进行深度加工与分析, 为入侵种的管理与控制提供更为有力的依据。

(2)入侵种的进化生态学: 外来种的入侵过程压缩了传统生态学和进化生物学所涉及的时间和空间尺度, 入侵种中的进化现象可以发生在数十个至数百个世代(而非百万代)的时间尺度内(Cox, 2004), 所以外来入侵种成为研究生态学和进化生物学重大问题的理想材料或模式生物(李博和马克平, 2010)。外来种的进化是在自然选择和人类干扰双重作用下的进化, 它既不同于纯粹的自然进化, 也不同于人类控制下的定向选择。近20年来, 国际生态学界在外来种进化的研究中获得的研究成果迅速增长, 而我国当前对这一问题的重视程度还远远不够。有鉴于此, 我们认为, 我国生态学家应从更宏观的尺度出发, 在如下方面重点关注生物入侵的进化生态学问题: ①外来种与土著种通过天然杂交与遗传渐渗而发生的基因组混合增加新入侵种或生物型进化的风险及其机理; ②外来种在新的区域扩张和移殖过程中的进化动态, 以及与入侵相关的遗传变异式样, 即遗传多样性对外来种的短期入侵和长期进化的影响机制; ③外来种与土著种的相互利用(如取食、寄生、互利、偏利等)选择与适应过程及其机制; ④外来种在入侵区的进化所导致的生态系统对全球气候变化响应的动态及机理。

(3)入侵种的基因组学: 入侵种的基因组学是一个新兴的研究领域, 它利用基因组学方法研究与外来种入侵性相关的分子基础和表达调控机制, 甄别入侵性的基因型, 进而在基因组水平上揭示外来种入侵性产生和进化的分子机制。目前运用基因组学方法对虉草(Phalaris arundinacea) (Lavergne et al., 2010)、巨大一枝黄花(Solidago gigantean) (Schlaepfer et al., 2008)、红火蚁(Wurm et al., 2011)等外来种的入侵机制研究已取得了重要进展, 涉及的技术主要包括比较基因组学、群体基因组学和表观基因组学等。对入侵台湾的红火蚁种群利用比较基因组法分析发现, 该入侵种的同源端粒酶至少有4种剪切形式, 在两套互斥外显子中有不同的作用, 同源端粒酶和其他方面独特的基因组可能是促使该物种复杂社会功能形成和入侵行为产生的重要生理机制(Wurm et al., 2011)。通过比较3,676种被子植物的DNA 1C值和基本基本组(basic genome)大小发现, DNA C值的大小可能是衡量被子植物入侵性的一个重要指标(Chen et al., 2010a)。但整体来看, 入侵基因组学仍处于发展初期。因此, 选择理想的入侵模式种, 建立其入侵性研究的模式体系, 是当前亟待解决的问题。基于此需求, 可以选择具有合适的基因组学研究特征(基因组小、二倍体、易遗传转化、繁殖迅速、自花授粉、株型小等)并在中国入侵态势较为严重的外来种(如菊科植物)首先切入, 重点解决以下3个科学问题: ①与外来种的入侵性密切相关的基因或基因型的发现; ②这些基因或基因型增强外来种入侵性表达的机理; ③在基因组水平上外来种的入侵性(特征)实现进化的途径(尚蕾等, 2010)。

(4)生物入侵的生态系统影响: 生物入侵的影响及危害是制订入侵种管理政策的重要依据之一。过去10年来, 我们对生物入侵的影响认识, 在很大程度上还滞留在其引起的农、林、牧、渔业的产量与质量的损失和防治费用这样的阶段。随着人们对生态系统变化和稳定性维持的重视, 评价外来种入侵对生态系统的全面影响已成为当前入侵生物学领域的研究热点之一。尽管我们近年来对互花米草等极个别入侵种给予了系统研究(Li et al., 2009), 但相对于我国庞大的外来种数量, 这些研究还显得远远不够。因此, 我们建议, 在今后一段时间, 有必要从多尺度、多个生态系统出发, 针对更多的外来种(包括植物、动物和微生物)入侵对森林、草原、农田、海洋和淡水生态系统等的影响展开系统、深入的研究。在陆地生态系统中, 尤其要加强外来种入侵对生态系统的结构和组成(如生物多样性、种间关系、营养关系、群落演替等)、物质循环(如碳、氮物质循环)以及生态系统服务(如生态系统产品、气候调节、病害防治、传粉、水体的净化、文化和美学价值等)等方面的影响研究, 并探讨其中的机制。在海洋生态系统中, 尤其要加强外来种入侵对海洋生物多样性、海洋生物灾害及海洋的生态服务功能等影响的研究。在淡水生态系统中, 尤其要加强外来种入侵对淡水生物多样性、养殖业和航运等方面的影响及其机制研究。

(5)生物入侵与全球变化主要过程的相互作用: 生物入侵并非独立事件, 而是与全球变化其他问题同时袭来。可以说, 几乎所有全球环境变化的主要过程(如全球气候变化、土地利用和土地覆盖变化、环境污染、荒漠化与生物多样性丧失等)都与生物入侵相互作用(Mooney & Hobbs, 2000; Perrings et al., 2010), 这不仅正在催生出一次进化史上的革命(Cox, 2004), 而且将可能产生难以预测的生态、经济和社会后果。然而, 尽管这种将生物入侵置于全球变化整体框架内的全局观念早已存在(Elton, 1958), 但生物入侵与全球变化其他过程之间的关系才刚刚得到重视(van der Putten et al., 2010)。在这方面, 我们首先应该了解全球变化是如何影响入侵种在特定生态系统中的入侵行为、能力和动态, 以及这种影响又是如何改变被入侵生态系统的结构与功能的(邓自发等, 2010)。因此, 针对当前全球变化及外来种入侵的现状, 有必要在不同层次上, 系统研究全球环境变化对外来种的入侵性、土著生态系统的可入侵性以及生态系统面临的潜在后果(李博和马克平, 2010)。当前和今后一段时间, 需要从景观、区域和全球等不同尺度, 重点围绕外来种入侵对气候变化、大气组成变化、土地利用变化、生物地球化学循环变化等过程的响应及其机制, 揭示外来种加剧土著生物多样性丧失、生态系统崩溃及生境均质化等的机制。特别是要加强气候变暖、CO2浓度上升、氮沉降等气候变化主要过程对外来种和土著种的物候、栖息地、种间关系、种群动态等影响的研究, 从而阐明气候变化导致外来种入侵的时间和空间格局的改变及其后果, 并制定科学的预警对策; 同时, 关于城市化扩张、土地利用变化等干扰过程对外来种入侵潜力和后果及相应的控制对策调整等也值得深入探讨。通过以上研究, 最终为人类在全球变化背景下找到生物入侵管理与控制的有效对策提供科学依据。

(6)管理与控制技术创新: 生物入侵的管理与控制渠道是否畅通, 技术是否有效, 将直接决定我国生物入侵进一步发生的趋势。中国生物入侵的管理与预防控制技术创新虽然已取得了重要成就, 但相对于生物入侵的严重态势, 今后这方面的研究仍然值得进一步强化。建议重点从以下几方面推进: 一是借鉴发达国家生物入侵的管理方法, 构建跨行业、跨部门的生物入侵管理机构, 制定生物入侵管理的法律法规, 加强生物入侵的行政和法律干预。二是进一步强化更加高效的生物入侵预防和控制技术体系建设, 如针对生物入侵的预防预警, 重点研究不同管理和气候背景下外来种的定量风险评估技术, 针对不同生态系统和使用目的, 构建针对性风险评估系统, 如城市绿地外来种入侵风险评估体系(鞠瑞亭和李博, 2012), 构建基于Web GIS的外来种远程风险评估模型和风险决策支撑平台, 为管理部门更加便捷、科学地认识外来种的风险性提供依据(万方浩等, 2011c)。三是要针对检疫性入侵种的口岸检测监测, 进一步研究恶性外来种的快速分子检测技术、高通量基因芯片诊断技术和DNA条形码鉴定技术, 构建远程专家识别、诊断平台, 强化PDA等智能系统与网格化管理监测体系的结合应用和二次开发, 提高监测的效率和精度(罗卿权等, 2011)。四是针对重大入侵种的阻截扑灭, 发展这些物种的热处理、灯光处理、辐射不育等物理灭除技术, 环境友好药剂治理等化学处理技术, 以及生物防治、转基因致死及遗传防治等生物技术的研究, 尤其在生物防治上, 需以土著天敌资源挖掘和原产地引进相结合的途径, 重点发展天敌规模化工厂生产技术, 建立大型的高标准天敌生产工厂, 并开展天敌释放研究, 探索生物防治的最佳控害模式。最后, 针对已成灾的入侵种的生态修复, 重点开发新型替代种的筛选及其修复功能评估, 构建以替代更新和生态恢复为核心的受损生态系统修复技术体系(万方浩等, 2011c)。

致谢:

中国农业科学院植物保护研究所万方浩研究员对本文提出许多修改意见; 中国科学院动物研究所张润志研究员提供了马铃薯甲虫和苹果蠹蛾资料, 并对论文初稿进行了修改; 沈阳农业大学生物科学技术学院冯玉龙教授、中国科学院植物研究所桑卫国研究员、扬州大学园艺与植物保护学院杜予州教授、杨国庆副教授对本文初稿进行了完善; 青岛农业大学褚栋教授提供了烟粉虱资料; 华南农业大学许益隽博士提供了红火蚁资料; 上海市园林科学研究所高磊博士、王凤工程师和复旦大学卫书娟同学协助资料收集。在此, 一并敬致谢意。

参考文献

Allendorf FW, Lundquist LL (2003)

Introduction: population biology, evolution, and control of invasive species

Conservation Biology, 17, 24-30.

DOI:10.1046/j.1523-1739.2003.02365.x      URL     [本文引用: 1]

An SQ, Gu BH, Zhou CF, Wang ZS, Deng ZF, Zhi YB, Li HL, Chen L, Yu DH, Liu YH (2007)

Spartina invasion in China: implications for invasive species management and future research

Weed Research, 47, 183-191.

DOI:10.1111/wre.2007.47.issue-3      URL     [本文引用: 1]

Ascunce MS, Yang CC, Oakey J, Calcaterra L, Wu WJ, Shih CJ, Goudet J, Ross KG, Shoemaker D (2011)

Global invasion history of the fire ant Solenopsis invicta

Science, 331, 1066-1068.

DOI:10.1126/science.1198734      URL     [本文引用: 1]

The fire ant Solenopsis invicta is a significant pest that was inadvertently introduced into the southern United States almost a century ago and more recently into California and other regions of the world. An assessment of genetic variation at a diverse set of molecular markers in 2144 fire ant colonies from 75 geographic sites worldwide revealed that at least nine separate introductions of S. invicta have occurred into newly invaded areas and that the main southern U.S. population is probably the source of all but one of these introductions. The sole exception involves a putative serial invasion from the southern United States to California to Taiwan. These results illustrate in stark fashion a severe negative consequence of an increasingly massive and interconnected global trade and travel system.

Blumenthal D (2005)

Ecology-interrelated causes of plant invasion

Science, 310, 243-244.

DOI:10.1126/science.1114851      URL     PMID:16224008      [本文引用: 1]

In his or her Perspective, Blumenthal discusses how plants from high-resource habitats are often poorly defended, nutritious, and strongly regulated by enemies. Consequently, these species may benefit the most by entering new habits to escape their natural enemies. This hypothesis predicts that high-resource invasive species may be particularly susceptible to biological control and that increases in resource availability will favor exotic plants.

Bossdorf O, Auge H, Lafuma L, Rogers WE, Siemann E, Prati D (2005)

Phenotypic and genetic differentiation between native and introduced plant populations

Oecologia, 144, 1-11.

DOI:10.1007/s00442-005-0070-z      URL     [本文引用: 1]

Plant invasions often involve rapid evolutionary change. Founder effects, hybridization, and adaptation to novel environments cause genetic differentiation between native and introduced populations and may contribute to the success of invaders. An influential idea in this context has been the Evolution of Increased Competitive Ability (EICA) hypothesis. It proposes that after enemy release plants rapidly evolve to be less defended but more competitive, thereby increasing plant vigour in introduced populations. To detect evolutionary change in invaders, comparative studies of native versus introduced populations are needed. Here, we review the current empirical evidence from: (1) comparisons of phenotypic variation in natural populations; (2) comparisons of molecular variation with neutral genetic markers; (3) comparisons of quantitative genetic variation in a common environment; and (4) comparisons of phenotypic plasticity across different environments. Field data suggest that increased vigour and reduced herbivory are common in introduced plant populations. In molecular studies, the genetic diversity of introduced populations was not consistently different from that of native populations. Multiple introductions of invasive plants appear to be the rule rather than the exception. In tests of the EICA hypothesis in a common environment, several found increased growth or decreased resistance in introduced populations. However, few provided a full test of the EICA hypothesis by addressing growth and defence in the same species. Overall, there is reasonable empirical evidence to suggest that genetic differentiation through rapid evolutionary change is important in plant invasions. We discuss conceptual and methodological issues associated with cross-continental comparisons and make recommendations for future research. When testing for EICA, greater emphasis should be put on competitive ability and plant tolerance. Moreover, it is important to address evolutionary change in characteristics other than defence and growth that could play a role in plant invasions.]]>

Callaway RM, Ridenour WM (2004)

Novel weapons: invasive success and the evolution of increased competitive ability

Frontiers in Ecology and Environment, 2, 436-443.

DOI:10.1890/1540-9295(2004)002[0436:NWISAT]2.0.CO;2      URL     [本文引用: 1]

Cao Z, Wang H, Meng L, Li B (2011)

Risk to nontarget plants from Ophraella communa (Coleoptera: Chrysomelidae), a potential biological control agent of alien invasive weed Ambrosia artemisiifolia (Asteraceae) in China

Ambrosia artemisiifolia, 46, 375-381.

[本文引用: 1]

Cao ZL (曹子林), Wang XL (王晓丽), Yang GY (杨桂英) (2012)

Allelopathic effect of volatiles of Eupatorium adenophorum on seed germination and seedling growth of Pinus yunanensis

Acta Agriculturae Universitis Jiangxiensis (江西农业大学学报), 34, 77-81. (in Chinese with English abstract)

[本文引用: 1]

Catford JA, Jansson R, Nilsson C (2009)

Reducing redundancy in invasion ecology by integrating hypotheses into a single theoretical framework

Diversity and Distributions, 15, 22-40.

[本文引用: 1]

Chang WN (常望霓), Huang JL (黄建利), Zhang Y (张亚), Bai LY (柏连阳) (2011)

Present situation and prospects in biological control of Ambrosia artemisiifolia

Weed Science (杂草科学), 29, 7-10. (in Chinese)

[本文引用: 1]

Chen FM (陈凤毛), Ye JR (叶建仁), Wu XQ (吴小芹) (2007a)

Detection technique of Bursaphelenchus xylophilus using real time PCR

Journal of Nanjing Forestry University (Natural Science Edition) (南京林业大学学报(自然科学版)), 31, 121-124. (in Chinese with English abstract)

[本文引用: 1]

Chen FM (陈凤毛), Ye JR (叶建仁), Wu XQ (吴小芹), Tang J (汤坚) (2007b)

Development of quick detection kit for Bursaphelenchus xylophilus

Journal of Nanjing Forestry University (Natural Science Edition) (南京林业大学学报(自然科学版)), 31, 133-135. (in Chinese with English abstract)

[本文引用: 1]

Chen GQ, Guo SL, Yin LP (2010b)

Applying DNA C-values to evaluate invasiveness of angiosperms: validity and limitation

Biological Invasions, 12, 1335-1348.

[本文引用: 2]

Chen HL, Li B, Hu JB, Chen JK, Wu JH (2007a)

Benthic nematode communities in the Yangtze River estuary as influenced by Spartina alterniflora invasions

Marine Ecology Progress Series, 336, 99-110.

[本文引用: 1]

Chen JH, Wang L, Li YL, Zhang WQ, Fu XH, Le YQ (2012)

Effect of Spartina alterniflora invasion and its controlling technologies on soil microbial respiration of a tidal wetland in Chongming Dongtan, China

Ecological Engineering, 41, 52-59.

[本文引用: 1]

Chen SG, Xu XM, Dai XB, Yang CL, Qiang S (2007b)

Identification of tenuazonic acid as a novel type of natural photosystem II inhibitor binding in QB-site of Chlamy- domonas reinhardtii

Biochimica et Biophysica Acta, 1767, 306-318.

DOI:10.1016/j.bbabio.2007.02.007      URL     PMID:17379181      [本文引用: 1]

Tenuazonic acid (TeA) is a natural phytotoxin produced by Alternaria alternata, the causal agent of brown leaf spot disease of Eupatorium adenophorum. Results from chlorophyll fluorescence revealed TeA can block electron flow from Q(A) to Q(B) at photosystem II acceptor side. Based on studies with D1-mutants of Chlamydomonas reinhardtii, the No. 256 amino acid plays a key role in TeA binding to the Q(B)-niche. The results of competitive replacement with [(14)C]atrazine combined with JIP-test and D1-mutant showed that TeA should be considered as a new type of photosystem II inhibitor because it has a different binding behavior within Q(B)-niche from other known photosystem II inhibitors. Bioassay of TeA and its analogues indicated 3-acyl-5-alkyltetramic and even tetramic acid compounds may represent a new structural framework for photosynthetic inhibitors.

Chen X, Chen XX, Wan XW, Weng BQ, Huang Q (2010a)

Water hyacinth (Eichhornia crassipes) waste as an adsorbent for phosphorus removal from swine wastewater

Bioresource Technology, 101, 9025-9030.

DOI:10.1016/j.biortech.2010.07.013      URL     PMID:20674342      [本文引用: 1]

Both live plants and dried straw of water hyacinth were applied to a sequential treatment of swine wastewater for nitrogen and phosphorus reduction. In the facultative tank, the straw behaved as a kind of adsorbent toward phosphorus. Its phosphorus removal rate varied considerably with contact time between the straw and the influent. In the laboratory, the straw displayed a rapid total phosphorus reduction on a KH(2)PO(4) solution. The adsorption efficiency was about 36% upon saturation. At the same time, the water hyacinth straw in the facultative tank enhanced NH(3)-N removal efficiency as well. However, no adsorption was evident. This study demonstrated an economically feasible means to apply water hyacinth phosphorus straw for the swine wastewater treatment. The sequential system employed significantly reduced the land use, as compared to the wastewater stabilization pond treatment, for pollution amelioration of swine waste.

Chen XR, Shen G, Wang YL, Zheng XB, Wang YC (2007c)

Identification of Phytophthora sojae genes upregulated during the early stage of soybean infection

FEMS Microbiology Letters, 269, 280-288.

URL     PMID:17263843      [本文引用: 1]

Chen YC, Kafle L, Shih CJ (2011)

Interspecific competition between Solenopsis invicta and two native ant species Pheidole fervens and Monomorium chinensis

Journal of Economic Entomology, 104, 614-621.

DOI:10.1603/ec10240      URL     PMID:21510213      [本文引用: 1]

This study was designed to understand the effects of the interspecific competition between red imported fire ant, Solenopsis invicta Buren and two native ant species, Pheidole fervens Smith and Monomorium chinense Santschi, by conducting colony interference and individual confrontation tests under laboratory conditions. The colony interference test showed that both native ant species, owing to their numerical advantage, killed the Solenopsis invicta virus-1 (SINV-1)-infected or healthy queens of S. invicta. Significantly less time was required for M. chinense to kill all SINV-1-infected S. invicta compared with the time required to kill the healthy S. invicta. Compared with healthy S. invicta, SINV-1-infected S. invicta spent a longer time eliminating the P. fervens colonies. In confrontation tests, M. chinense killed a significantly higher number of infected S. invicta minors than they did healthy minors, but the number of S. invicta majors (either infected or healthy) killed was substantially less. This study found that the viral infection weakened the competitive ability of S. invicta and made them prone to be eliminated by M. chinense but not by P. fervens.

Chen ZY, Li B, Zhong Y, Chen JK (2004)

Local competitive effects of introduced Spartina alterniflora on Scirpus mariqueter at Dongtan of Chongming Island, the Yangtze River estuary and their potential ecological consequences

Hydrobiologia, 528, 99-106.

DOI:10.1007/s10750-004-1888-9      URL     [本文引用: 1]

Chen ZY (陈中义), Fu CZ (傅萃长), Wang HY (王海毅), Li B (李博), Wu JH (吴纪华), Chen JK (陈家宽) (2005)

Effects of Spartina alterniflora invasions on the benthic macro-onvertebrates community at Dongtan of Chongming salt marsh, the Yangtze River Estuary

Wetland Science (湿地科学), 3, 1-7. (in Chinese with English abstract)

URL     [本文引用: 1]

Cheng XL, Luo YQ, Chen JQ, Lin GH, Chen JK, Li B (2006)

Short-term C4 plant

Spartina alterniflora invasions change the soil carbon in C3 plant-dominated tidal wetlands on a growing estuarine island

Soil Biology and Biochemistry, 38, 3380-3386.

DOI:10.1016/j.soilbio.2006.05.016      URL     [本文引用: 1]

Cheng XY, Xie PZ, Cheng FX, Xu RM, Xie BY (2008a)

Competitive displacement of a native species ompetitive displacement of a native species Bursap- helenchus mucronatus by an alien species Bursaphelenchus xylophilus (Aphelenchida: Aphelenchoididae): a case of successful invasion

Biological Invasions, 11, 205-213.

DOI:10.1007/s10530-008-9225-2      URL     [本文引用: 1]

Cheng XY, Cheng FX, Xu RM, Xie BY (2008b)

Genetic variation in the invasive process of Bursaphelenchus xylophilus (Aphelenchida: Aphelenchoididae) and its possible spread routes in China

Heredity, 100, 356-365.

DOI:10.1038/sj.hdy.6801082      URL     PMID:18091770      [本文引用: 1]

Pinewood nematode (Bursaphelenchus xylophilus) is an invasive species that causes a destructive forest disease-pine wilt disease. This disease has been prevalent in some countries in Asia since the 1970s. An amplified fragment length polymorphism survey was used to compare the genetic variation of native and invasive nematode populations in China and to examine the changes in genetic diversity during the invasion process. The genetic diversity of Chinese populations was slightly higher than that of American populations. Analysis of groups sampled from different invasive stages in China, showed that no obvious change in genetic diversity. Hence, genetic drift and founder effects are not obvious in the invasion process. Phylogenetic analysis showed that Chinese pinewood nematode populations were closer to Japanese populations than to American populations. On the basis of the genetic relationships among samples, two major invasion pathways in China are suggested. One is from Guangdong to Anhui and Zhejiang, and the other is from Guangdong to Jiangsu and then from Jiangsu to Hubei, Guizhong and Congqing. The results imply that it is important to reinforce both domestic and international quarantine to control the spread of pinewood nematode.

Chu D (褚栋), Zhang YJ (张友军), Cong B (丛斌), Xu BY (徐宝云), Wu QJ (吴青君) (2004)

The invasive mechanism of a worldwide important pest, Bemisia tabaci (Gennadius) biotype B

Acta Entomologica Sinica (昆虫学报), 47, 400-406. (in Chinese with English abstract)

URL     [本文引用: 1]

Bemisia tabaci (Gennadius) biotype B is an important pest that have outbreaked and caused severe damage worldwide through the last 20 years. Recent progress in studying mechanisms of its invasion indicates the greater ability of competitive displacement which involves the competition of ecological niche, reproductive interference, the interactions of whitefly and virus with indigenous B. tabaci and other competing herbivores, is the most important factor. The successful invasion and rapid spread is affected by the pesticide resistance. In the meantime, the ecological environment (such as host, climatic factor, overwintering sites) also affected the invasion of B. tabaci biotype B. Finally the importance of the invasive mechanism of B. tabaci biotype B and potential directions for future research are discussed.]]>

Chu D, Zhang YJ, Brown JK, Cong B, Xu BY, Wu QJ, Zhu GR (2006)

The introduction of the exotic Q biotype of Bemisia tabaci (Gennadius) from the Mediterranean region into China on ornamental crops

Florida Entomologist, 89, 168-174.

DOI:10.1653/0015-4040(2006)89[168:TIOTEQ]2.0.CO;2      URL     [本文引用: 1]

Chu D, Wan FH, Zhang YJ, Brown JK (2010a)

Change in the biotype composition of Bemisia tabaci in Shandong Province of China from 2005 to 2008

Environmental Entomology, 39, 1028-1036.

DOI:10.1603/EN09161      URL     PMID:20550819      [本文引用: 2]

Certain biotypes of the Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) complex cause extensive damage and are important pests and virus vectors in agricultural crops throughout the world. Among the most invasive and well studied are the B and Q biotypes. Recent reports in Shandong Province, China, have indicated that the Q biotype was introduced there in approximately 2005, whereas the B biotype has been established there for approximately 10 yr. Even so, the present distribution of the two biotypes in Shandong has not been examined. The results of this study showed that the B and Q biotypes are both present in Shandong Province based on bar-coding using a approximately 450-base fragment of the mitochondrial cytochrome oxidase I (mtCOI) gene. In addition, a B biotype-specific polymerase chain reaction primer pair that amplifies a approximately 300 bp mtCOI fragment was designed and used to examine the biotype composition of B. tabaci in selected crops from six provincial locations, using the general mtCOI primers as an internal positive control for DNA quality. The results of this study indicated that the Q biotype was the predominant B. tabaci colonizing all of the crops in the study sites examined. This suggests that the Q biotype has displaced the B biotype in Shandong Province of China, which until now was the predominant biotype. This is the first report of the displacement of the B by the Q biotype in field grown crops in China, and in a locale where neither the B nor the Q biotype is native. We hypothesize that this phenomenon may have been exacerbated by the widespread use of neonicotinoid insecticides for whitefly control, given the sustained efficacy thus far of neonicotinoids against the B biotype, and their failure at times to effectively control the Q biotype.

Chu D, Zhang YJ, Wan FH (2010b)

Cryptic invasion of the exotic Bemisia tabaci biotype Q occurred widespread in Shandong Province of China

Florida Entomologist, 93, 203-207.

DOI:10.1653/024.093.0209      URL     [本文引用: 2]

Conway WC, Smith LM, Bergan JF (2002)

Potential allelopathic interference by the exotic Chinese tallow tree (Sapium sebiferum)

The American Midland Naturalist, 148, 43-53.

DOI:10.1674/0003-0031(2002)148[0043:PAIBTE]2.0.CO;2      URL     [本文引用: 1]

Cox GW (2004) Alien Species and Evolution: The Evolutionary Ecology of Exotic Plants, Animals, Microbes, and Interacting Native Species. Island Press, Washington.

[本文引用: 3]

Cui L, Yin W, Tang Q, Dong S, Zheng X, Zhang Z, Wang Y (2010)

Distribution, pathotypes, and metalaxyl sensitivity of Phytophthora sojae from Heilongjiang and Fujian provinces in China

Plant Disease, 94, 881-884.

URL     PMID:30743553      [本文引用: 1]

Cui X, Wan F, Xie M, Liu T (2008)

Effects of heat shock on survival and reproduction of two white fly species, Trialeurodes vaporariorum and Bemisia tabaci biotype B

Journal of Insect Science, 8, 24.

[本文引用: 1]

Dai L (戴莲), Li HN (李会娜), Jiang ZL (蒋智林), Wan FH (万方浩), Liu WX (刘万学) (2012)

Invasive effects of Ageratina adenophora (Asteraceae) on the changes of effective functional bacteria, enzyme activity and fertility in rhizosphere soil ecosystem

Ecology and Environmental Sciences (生态环境学报), 21, 237-242. (in Chinese with English abstract)

[本文引用: 1]

Davis MA, Grime JP, Thompson K (2000)

Fluctuating resources in plant communities: a general theory of invisibility

Journal of Ecology, 88, 528-534.

DOI:10.1046/j.1365-2745.2000.00473.x      URL     [本文引用: 1]

Deng X (邓旭), Wang J (王娟), Tan JC (谭济才) (2010)

Effect of ragweed (Ambrosia artemisiifolia L.), an invasive weed, on plant biodiversity in Hunan Province

Chinese Agricultural Science Bulletin (中国农学通报), 26, 309-314. (in Chinese with English abstract)

URL     [本文引用: 1]

To find out the nutrition characteristics of Brazilian Banana between rhizosphere and bulk soil, we selected three parent materials,determined the nitrogen status of various forms, soil organic carbon content and changes of s oil pH both in the rhizosphere and bulk soil , analyzed their significant difference and relevance. The result is that: Total nitrogen, ammonium, nitrate and alkaline in rhizosphere were 50.7 %、35.9%、36.7 % and 19.3 % higher than that the bulk soil . Soil organic in the rhizosphere was 36.8 % higher and s oil pH was lower by 0.06 units than the average. The nutrition of rhizosphere was higher than bulk soil in three parent materials. And there were significant differences in content of soil total nitrogen, nitrate and organic content between rhizosphere and bulk soil among three parent materials, however, the ammonium and alkaline were significant differences in different parent materials. Ammonium in the rhizosphere and bulk soil was not correlated with other nutrients.

Deng ZF (邓自发), Ouyang Y (欧阳琰), Xie XL (谢晓玲), Qing H (清华), Xiao Y (肖燕), An SQ (安树青) (2010)

The effects of primary process of global change on biological invasion in coastal ecosystem

Biodiversity Science (生物多样性), 18, 605-614. (in Chinese with English abstract)

URL     [本文引用: 1]

2, sea level rise and eutrophication on the process of biological invasion have been analyzed based on the invasive status of coastal ecosystem. The results indicate that elevated CO2 disturbs the competitive balance between invasive and native species. Sea level rise adjusts the spatial pattern of invasive species and eutrophication of coastal ecosystem promotes further spread of alien species. At last, the research hotspots about interactions between global change and biological invasion in coastal ecosystem are discussed.]]>

Ding H (丁晖), Xu HG (徐海根), Liu ZL (刘志磊) (2007)

Impacts of invasion of Eupatorium adenophorum on vegetation diversity

Journal of Ecology and Rural Environment (生态与农村环境学报), 23, 29-32, 75. (in Chinese with English abstract)

URL     [本文引用: 1]

Based on 48 years of summer rainfall data,Morlet wavelet analysis was done of the sequence of summer rainfall in Nenjiang and Harbin,on a multi-time scale.Results show that in recent years,the summer rainfall in Harbin displayed the most apparent cyclical variation on a 29-38-year scale,the largest modulus,and the strongest energy;and the summer rainfall in Nenjiang did an obvious cyclical variation on a 19-30-year scale,and its energy value is quite large.Both the two cyclical variations showed a trend of being stronger in energy with the time.Presently,both the summer rainfalls in Nenjiang and Harbin are in a long cyclical period of low-water.It is predicted that in a relatively long period the summer rainfall will be lower than the average of the past 48 years.

Ding H (丁晖), Xu HG (徐海根), Qiang S (强胜), Meng L (孟玲), Han ZM (韩正敏), Liao JL (缪锦来), Hu BS (胡白石), Sun HY (孙红英), Huang C (黄成), Lei JC (雷军成), Le ZF (乐志芳) (2011)

Status quo and trends of biological invasion into China

Journal of Ecology and Rural Environment (生态与农村环境学报), 27, 35-41. (in Chinese with English abstract)

URL     [本文引用: 1]

Combining literature review, field survey and expert consultation, investigations were carried out of invasive alien species (IAS) in forest, wetland, grassland, desert, inland water and ocean (sea) ecosystems of China. A total of 488 IAS were identified with detailed information collected including name, taxonomy, distribution, origin, location and time first detected or introduced, pathway, hazard, nutrient circumstance, and habits. Of the 488 IAS,265 were plants, 171 animals, 26 fungi, 12 vira, 11 procaryotes and 3 protists. Analysis of the invasion years of the 392 IAS that have specific records reveals that only 31 IAS came in before 1850 and the number of IAS has been increasing steadily since 1850, especially since 1950. During the 60 years after 1950, 209 IAS appeared. Their debuts occurred mainly in the coastal regions and border provinces, like Yunnan Province and Xinjiang Uyghur Autonomous Region, but the stage tends to be moving northward. The percentage of intentional introduction almost equaled to that of unintentional introduction, but the latter has become the primary in recent years. America, Europe and other parts of Asia are the main origins of IAS of China. The number of IAS presents a decreasing trend from coastal towards inland.

Ding J, Mack RN, Lu L, Ren M, Huang H (2008)

China’s booming economy is sparking and accelerating biological invasions

BioScience, 58, 317-324.

DOI:10.1641/B580407      URL     [本文引用: 2]

Ding JR, Wang WF, Zhang G (2001)

Water hyacinth in China: its distribution, problems and control status

In: Biological and Integrated Control of Water Hyacinth, Eichhornia crassipes (eds Julien MH, Hill MP, Center TD, Ding JQ), pp. 29-32. Australian Centre for International Agricultural Research, Canberra.

[本文引用: 2]

Ding Y, Chu JJ (2005)

Biological control of Eichhornia crassipes (Mart.) Solms

Weed Science, 3, 1-5.

URL     [本文引用: 1]

Dong M (董梅), Lu JZ (陆建忠), Zhang WJ (张文驹), Chen JK (陈家宽), Li B (李博) (2006)

Canada goldenrod (Solidago canadensis): an invasive alien weed rapidly spreading in China

Acta Phytotaxonomica Sinica (植物分类学报), 44, 72-85. (in Chinese with English abstract)

DOI:10.1360/aps050068      URL     [本文引用: 1]

Duan YL (段彦丽), Qu LJ (曲良建), Wang YZ (王玉珠), Zhang YA (张永安), Tao WQ (陶万强), Guan L (关玲), Yang ZQ (杨忠岐) (2009)

Sustainable effects for Hyphantria cunea nuclear polyhedrosis virus on the host insects and the diffusion approach of the virus

Scientia Silvae Sinicae (林业科学), 45, 83-86. (in Chinese with English abstract)

DOI:10.11707/j.1001-7488.20090713      URL     [本文引用: 1]

A binomial logistic model was established based on RS, GIS and GPS techniques to simulate and predict the distribution of nest-site of Red-crowned Crane (Grus japonensis) in Zhalong National Nature Reserve (ZLNNR), Heilongjiang Province, China. By using the data of present and absent points, a spatial auto-correlation test was conducted to screen the variables for the model. The result showed the factors affected the nest-site selection by Red-crowned Cranes included natural environment factors such as reed height and coverage, and elevation, and disturbance factors such as distance to residential area, density of dykes and roads around the nest. The model was tested through calculating the match coefficients of observed and predicted present/absent data. The correctly classified rate reached to 80.4%. Meanwhile, the model predicted that the suitable habitat for nesting by Red-crowned Crane reached to 223.46 km2, which mainly distributed in the middle and north of ZLNNR. The model prediction basically reflected the suitable nesting habitat distribution, and might provide scientific basis for efficient conservation and management of the nesting habitats, and restoration of key nesting habitats for Red-crowned Crane in ZLNNR.]]>

Elton CS (1958) The Ecology of Invasions by Animals and Plants. Chapman and Hall, London.

[本文引用: 2]

Fang F (方芳), Mao W (茅玮), Guo SL (郭水良) (2005)

Study on allelopathic effects of the invasive plant annual fleabane

Bulletin of Botanical Research (植物研究), 25, 449-452. (in Chinese with English abstract)

URL     [本文引用: 1]

The extract from the organs above ground of Annual Fleabane(Erigeron annuus) was used to treat the seeds of some crop seeds, we determined the seeds germination, the length of root and seeding height of four crops.The results were as follows:(1) the allelopathic effects of Annual Fleabane does exist.(2) Higher concentration of the water extract of Annual Fleabane could restrain the seed germination and infancy developing, while lower concentration of the water extract could accelerate developing seed of Brassica chinensis var.communis and Lycoperiscum esculetum.

Feng YL, Auge H, Ebeling SK (2007a)

Invasive Buddleja davidii allocates more nitrogen to its photosynthetic machinery than five native woody species

Oecologia, 153, 501-510.

URL     PMID:17534664      [本文引用: 1]

Feng YL, Wang JF, Sang WG (2007b)

Biomass allocation, morphology and photosynthesis of invasive and noninvasive exotic species grown at four irradiance levels

Acta Oecologica, 31, 40-47.

Feng YL, Wang JF, Sang WG (2007

c) Irradiance acclimation, capture ability, and efficiency in invasive and non-invasive alien plant species

Photosynthetica, 45, 245-253.

Feng YL (2008a)

Nitrogen allocation and partitioning in invasive and native Eupatorium species

Physiologia Plantarum, 132, 350-358.

DOI:10.1111/j.1399-3054.2007.01019.x      URL     PMID:18275466      [本文引用: 1]

There is a trade-off between nitrogen (N) allocation to photosynthesis and to defence. Invasive species may reduce N allocation to defence because of the absence of natural enemies. Thus, I hypothesised that invasive species may allocate a higher fraction of total leaf N to photosynthesis and have higher light-saturated photosynthetic rate (P(max)) and photosynthetic N-use efficiency (PNUE) than closely related native species. To test these hypotheses, invasive Eupatorium adenophorum and native E. chinense and E. heterophyllum were compared in a limestone shrub. Unlike expectation, the invader did not allocate a higher fraction of leaf N to photosynthesis than the natives. However, it was more efficient in photosynthetic N partitioning than the natives. It partitioned a higher fraction of the photosynthetic N to carboxylation and showed higher use efficiency of the photosynthetic N, while the natives partitioned a higher fraction of the photosynthetic N to light-harvesting components. Total leaf N content was not significantly different among the three studied invasive and native species. For the invader, the higher fraction of leaf N allocated to carboxylation resulted in the higher N content in carboxylation and in both carboxylation and bioenergetics, which led to higher P(max), and therefore to higher PNUE, water-use efficiency, respiration efficiency and apparent quantum yield. These physiological advantages of the invader and its higher leaf area ratio may contribute to its invasiveness.

Feng YL (2008b)

Photosynthesis, nitrogen allocation and specific leaf area in invasive Eupatorium adenophorum and native Eupatorium japonicum grown at different irradiances

Physiologia Plantarum, 133, 318-326.

DOI:10.1111/j.1399-3054.2008.01072.x      URL     PMID:18312498      [本文引用: 1]

The mechanisms underlying biological invasions are still not well elucidated. In this study, ecophysiological traits of invasive Eupatorium adenophorum and native E. japonicum were compared at 10 irradiances in field. I hypothesized that the invader may allocate a higher fraction of leaf nitrogen (N) to photosynthesis and have higher light-saturated photosynthetic rate (P(max)) and specific leaf area (SLA) than E. japonicum. The invader had a significantly higher ability to acclimate to high irradiance than E. japonicum, while it showed a similar shade-tolerant ability. The invader indeed allocated a higher fraction of leaf N to photosynthesis than E. japonicum, which, with its high leaf N content (N(A)), resulted in a higher N content in photosynthesis (N(P)), contributing to its higher biochemical capacity for photosynthesis and P(max). However, the invader had a significantly lower SLA than E. japonicum, contributing to its higher P(max) but increasing its area-based leaf construction cost. The abilities to acclimate to a wider range of irradiance and to allocate a higher fraction of leaf N to photosynthesis, and the higher P(max), N(A), N(P) and leaf area ratio may contribute to the invasion of the invader. High SLA is not always necessary for invasive species.

Feng YL, Fu GL (2008)

Nitrogen allocation, partitioning and use efficiency in three invasive plant species in comparison with their native congeners

Biological Invasions, 10, 891-902.

Feng YL, Fu GL, Zheng YL (2008)

Specific leaf area relates to the differences in leaf construction cost, photosynthesis, nitrogen allocation and use efficiencies between invasive and noninvasive alien congeners

Planta, 228, 383-390.

URL     PMID:18392694      [本文引用: 1]

Feng YL, Lei YB, Wang RF, Callawy RM, Valiente-Banuet A, Inderjit, Li YP, Zheng YL (2009)

Evolutionary tradeoffs for nitrogen allocation to photosynthesis versus cell walls in an invasive plant

Proceedings of the National Academy of Sciences, USA, 106, 1853-1856.

[本文引用: 2]

Feng YL (冯玉龙), Liao ZY (廖志勇), Zhang R (张茹), Zheng YL (郑玉龙), Li YP (李扬苹), Lei YB (类延宝) (2009)

Adaptive evolution in response to environmental gradients and enemy release in invasive alien plant species

Biodiversity Science (生物多样性), 17, 340-352. (in Chinese with English abstract)

DOI:10.3724/SP.J.1003.2009.09115      URL     [本文引用: 2]

Feng YL, Li YP, Wang RF, Callawy RM, Valiente-Banuet A, Inderjit (2011)

A quicker return energy-use strategy by populations of a subtropical invader in the non-native range: a potential mechanism for the evolution of increased competitive ability

Journal of Ecology, 99, 1116-1123.

DOI:10.1111/j.1365-2745.2011.01843.x      URL     [本文引用: 1]

2. Previously, we found that invasive Ageratina adenophora plants increase nitrogen allocation to photosynthesis and reduce allocation to cell walls compared with native Ageratina plants, suggesting a shift from defence to growth in invasive populations. Here, carrying this work forward, we measured construction costs and benefits associated with photosynthesis at light saturation to leaves. We hypothesized that invasive Ageratina populations might employ a quicker return energy-use strategy by increasing light-saturated photosynthetic rates and photosynthetic energy-use efficiency (PEUE) and by decreasing leaf construction costs.3. Faster-growing plants from invasive populations (China and India) had significantly higher leaf nitrogen concentrations and specific leaf areas than plants from native populations (Mexico). Inconsistent with our prediction, leaf construction costs were not significantly different between plants from invasive and native populations, but higher light-saturated photosynthetic rates and in turn higher PEUE resulted in a significantly shorter payback time of construction costs, which allowed plants from invasive populations to grow faster.4. Synthesis. Our results indicate that Ageratina plants from populations in non-native ranges have a distinct quick return energy-use strategy, a high PEUE and a short payback time but not lower construction costs, which might provide a mechanistic explanation for the commonly observed increase in growth when plants are introduced to new parts of the world. To our knowledge, this is the first study to compare energy-use strategy for plants from invasive and native populations of a noxious invasive species. We cannot exclude some alternative hypotheses for these patterns, such as founder effects, but these ecophysiological differences might provide mechanistic insight for how the evolution of decreased allocation to defence may increase growth and competitive ability.]]>

Gao L (高雷), Li B (李博) (2004)

The study of a specious invasive plant, water hyacinth (Eichhornia crassipes): achievements and challenges

Acta Phytoecologica Sinica (植物生态学报), 28, 735-752. (in Chinese with English abstract)

[本文引用: 2]

Gao LX, Geng YP, Li B, Chen JK, Yang J (2011)

Genome- wide DNA methylation alterations of Alternanthera philoxeroides in natural and manipulated habitats: implications for epigenetic regulation of rapid responses to environmental fluctuation and phenotypic variation

Plant, Cell and Environment, 33, 1820-1827.

DOI:10.1111/j.1365-3040.2010.02186.x      URL     PMID:20545886      [本文引用: 1]

Alternanthera philoxeroides (alligator weed) is an invasive weed that can colonize both aquatic and terrestrial habitats. Individuals growing in different habitats exhibit extensive phenotypic variation but little genetic differentiation in its introduced range. The mechanisms underpinning the wide range of phenotypic variation and rapid adaptation to novel and changing environments remain uncharacterized. In this study, we examined the epigenetic variation and its correlation with phenotypic variation in plants exposed to natural and manipulated environmental variability. Genome-wide methylation profiling using methylation-sensitive amplified fragment length polymorphism (MSAP) revealed considerable DNA methylation polymorphisms within and between natural populations. Plants of different source populations not only underwent significant morphological changes in common garden environments, but also underwent a genome-wide epigenetic reprogramming in response to different treatments. Methylation alterations associated with response to different water availability were detected in 78.2% (169/216) of common garden induced polymorphic sites, demonstrating the environmental sensitivity and flexibility of the epigenetic regulatory system. These data provide evidence of the correlation between epigenetic reprogramming and the reversible phenotypic response of alligator weed to particular environmental factors.

Gao Y (高岩), Yi N (易能), Zhang ZY (张志勇), Liu HQ (刘海琴), Zou L (邹乐), Zhu HB (朱华兵), Yan SH (严少华) (2012)

Effect of water hyacinth on N2O emission through nitrification and denitrification reactions in eutrophic water

Acta Scinetiae Circumstantiae (环境科学学报), 32, 349-359. (in Chinese with English abstract)

[本文引用: 1]

Geng YP (耿宇鹏), Zhang WJ (张文驹), Li B (李博), Chen JK (陈家宽) (2004)

Phenotypic plasticity and invasiveness of alien plants

Biodiversity Science (生物多样性), 12, 447-455. (in Chinese with English abstract)

[本文引用: 1]

Geng YP, Pan XY, Xu CY, Zhang WJ, Li B, Chen JK (2006)

Phenotypic plasticity of invasive Alternanthera philoxe- roides in relation to different water availability, compared to its native congener

Acta Oecologica, 30, 380-385.

[本文引用: 1]

Geng YP, Pan XY, Xu CY, Zhang WJ, Li B, Chen JK, Lu BR, Song ZP (2007a)

Phenotypic plasticity rather than locally adapted ecotypes allows invasive alligator weed to colonize a wide range of habitats

Biological Invasions, 9, 245-256.

[本文引用: 2]

Geng YP, Pan XY, Xu CY, Zhang WJ, Li B, Chen JK (2007b)

Plasticity and ontogenetic drift of biomass allocation in response to above- and below-ground resource availabilities in perennial herbs: a case study of Alternanthera philoxeroides

Ecological Research, 22, 255-260.

[本文引用: 1]

Gray AJ, Marshall DF, Raybould AF (1991)

A century of evolution in Spartina anglica

Advances in Ecological Research, 21, 1-62.

[本文引用: 1]

Gui FR, Wan FH, Guo JY (2008)

Population genetics of Ageratina adenophora using inter-simple sequence repeat (ISSR) molecular markers in China

Plant Biosystems, 142, 255-263.

DOI:10.1080/11263500802150399      URL     [本文引用: 1]

Gui FR, Wan FH, Guo JY (2009)

Population structure of the invasive species, Ageratina adenophora as revealed by ISSR-PCR markers

Russian Journal of Plant Physiology, 56, 410-416.

DOI:10.1134/S1021443709030157      URL     [本文引用: 1]

A genetic analysis of nuclear DNA was performed by inter-simple sequence repeat-polymerase chain reaction (ISSR-PCR) technique on 32 populations of Ageratina adenophora, an invasive triploid weed in China. Among the 100 ISSR markers detected, 12 showed genetic variation both within and among the populations. Among the 446 amplified bands, 93.5% were found polymorphic. Most individuals (99%) displayed a unique ISSR fingerprint pattern, which yielded a high level of polymorphism (P o = 93.5%) and genetic diversity (Nei’s H T = 0.2354). The estimates of population variation, based on ISSR-PCR, were high, as measured by the analysis of molecular variance (AMOVA, F ST = 0.3140), the Wright’s F-statistics (G ST = 0.3453), and the Shannon’s information index (H sp = 0.3716). AMOVA revealed 68.6% genetic variation within the populations and 91.2% within the provinces. The Mantel test showed that genetic distance was significantly correlated with geographic distance.]]>

Guo JY, Zhou ZS, Zheng XW, Chen HS, Wan FH, Luo YH (2011)

Control efficiency of leaf beetle,

Ophraella com- muna, on the invasive common ragweed, Ambrosia artemis- iifolia, at different growing stages. Biocontrol Science and Technology, 21, 1049-1063.

[本文引用: 1]

Guo WC (郭文超), Tuerxun (吐尔逊), Xu YM (许咏梅), Liu J (刘建), Xu JJ (许建军), Wang PL (王佩玲), He J (何江), Xia ZH (夏正汉), Fu WJ (付文君), Jing XY (景新跃), Zhang DM (张冬梅) (2011)

Study and application on sustained and integrated control techniques of Colorado potato beetle

Xinjiang Agricultural Sciences (新疆农业科学), 48, 197-203. (in Chinese with English abstract)

[本文引用: 1]

Hu YP (胡亚鹏), Meng L (孟玲) (2007)

Potential impacts of alien herbivorous insect Ophraella communa (Coleoptera: Chrysomelidae) on non-target plants in mainland China

Chinese Journal of Ecology (生态学杂志), 26, 56-60. (in Chinese with English abstract)

[本文引用: 1]

Huang DC (黄顶成), Zhang RZ (张润志) (2011)

Taxonomic categorisation, origins and trends of invasive alien species in China

Journal of Biosafety (生物安全学报), 20, 113-118. (in Chinese with English abstract)

[本文引用: 2]

Huang D, Haack RA, Zhang R (2011)

Does global warming increase establishment rates of invasive alien species? a centurial time series analysis

PLoS ONE, 6, e24733.

DOI:10.1371/journal.pone.0024733      URL     PMID:21931837      [本文引用: 1]

BACKGROUND: The establishment rate of invasive alien insect species has been increasing worldwide during the past century. This trend has been widely attributed to increased rates of international trade and associated species introductions, but rarely linked to environmental change. To better understand and manage the bioinvasion process, it is crucial to understand the relationship between global warming and establishment rate of invasive alien species, especially for poikilothermic invaders such as insects. METHODOLOGY/PRINCIPAL FINDINGS: We present data that demonstrate a significant positive relationship between the change in average annual surface air temperature and the establishment rate of invasive alien insects in mainland China during 1900-2005. This relationship was modeled by regression analysis, and indicated that a 1 degrees C increase in average annual surface temperature in mainland China was associated with an increase in the establishment rate of invasive alien insects of about 0.5 species year(-)(1). The relationship between rising surface air temperature and increasing establishment rate remained significant even after accounting for increases in international trade during the period 1950-2005. Moreover, similar relationships were detected using additional data from the United Kingdom and the contiguous United States. CONCLUSIONS/SIGNIFICANCE: These findings suggest that the perceived increase in establishments of invasive alien insects can be explained only in part by an increase in introduction rate or propagule pressure. Besides increasing propagule pressure, global warming is another driver that could favor worldwide bioinvasions. Our study highlights the need to consider global warming when designing strategies and policies to deal with bioinvasions.

Huang D, Zhang R, Kim KC, Suarez AV (2012)

Spatial pattern and determinants of the first detection locations of invasive alien species in mainland China

PLoS ONE, 7, e31734.

DOI:10.1371/journal.pone.0031734      URL     PMID:22363715      [本文引用: 1]

BACKGROUND: The unintentional transport of species as a result of human activities has reached unprecedented rates. Once established, introduced species can be nearly impossible to eradicate. It is therefore essential to identify and monitor locations where invaders are most likely to establish new populations. Despite the obvious value of early detection, how does an agency identify areas that are most vulnerable to new invaders? Here we propose a novel approach by using the

Huang J, Xu YJ, Lu YY, Zeng L, Liang GW (2010)

Effects of red imported fire ants, Solenopsis invicta (Hymenoptera: Formicidae) on the relationship between native ants and Aphis craccivora (Hemiptera: Aphididae) in mungbean fields

Sociobiology, 55, 415-425.

[本文引用: 1]

Huang J, Xu YJ, Zeng L, Lu YY, Liang GW (2011)

Changes to the spatial distribution of Ageratum conyzoides (Asterales: Asteraceae) due to red imported fire ants Solenopsis invicta (Hymenoptera: Formicidae) in China

Journal of Insect Behavior, 24, 307-316.

Huang LH, Kang L (2007)

Cloning and interspecific altered expression of heat shock protein’s genes in the two leafminer species in response to thermal stress

Insect Molecular Biology, 16, 491-500.

DOI:10.1111/j.1365-2583.2007.00744.x      URL     PMID:17651238      [本文引用: 1]

Studies have demonstrated differences in temperature tolerance between two Liriomyza species, L. huidobrensis and L. sativae. To investigate whether the heat shock proteins (Hsps) in the two species have different expression profiles during temperature stress, we cloned hsp90, 70, 60, 40 and 20, and analysed their expression profiles across temperature gradients by real-time quantitative PCR and Western blotting. The results revealed that the number of TATA-box-like elements and A/T-rich insertion/deletions within the 5' UTRs of the hsps are different in the two species. The temperatures for onset (T(on)) or maximal (T(max)) induction of hsp expression in L. huidobrensis were generally 2.5-10 degrees C lower than those in L. sativae, and the T(on) were highly consistent with the temperature limits of the northern boundary of the range of these two leafminer species. These studies confirmed, in terms of gene expression levels, that L. huidobrensis is more cold tolerant than L. sativae, which is more heat tolerant, and suggest that the T(on) (or T(max)) of hsps can represent the differences in temperature tolerance of these two leafminer species, and may be used to determine their natural geographical distribution limits.

Hufbauer RA, Torchin ME (2007)

Integrating ecological and evolutionary theory of biological invasions

In: Biological Invasions (ed. Nentwig W), pp. 79-96. Springer Berlin Heidelberg, Berlin.

[本文引用: 1]

Jia X, Pan XY, Li B, Chen JK, Yang XZ (2009)

Allometric growth, disturbance regime, and dilemmas of controlling invasive plants: a model analysis

Biological Invasions, 11, 743-752.

[本文引用: 1]

Jia X, Pan XY, Sosa A, Li B, Chen JK (2010)

Differentiation in growth and biomass allocation among three native Alternanthera philoxeroides varieties from Argentina

Plant Species Biology, 25, 85-92.

[本文引用: 1]

Jiang K, Ma ZJ, Li B, Li J, Chen JK (2007)

Foraging strategies involved in habitat use of shorebirds at the intertidal area of Chongming Dongtan, China

Ecological Research, 22, 559-570.

[本文引用: 1]

Jiang KY (蒋科毅), Yu MJ (于明坚), Ding P (丁平), Xu XH (徐学红), Jiang P (蒋平), Zhou CM (周成枚), Lu G (陆高) (2005)

Avian community response to vegetation succession caused by the pine wood nematode in Zhejiang, China

Biodiversity Science (生物多样性), 13, 496-506. (in Chinese with English abstract)

[本文引用: 1]

Jiang MX, Way MO, Yoder R, Zhang WJ, Cheng JA (2006)

Elytral color dimorphism in rice water weevil: occurrence in spring populations and relation to female reproductive development

Annals of the Entomological Society of Ame- rica, 99, 1127-1132.

[本文引用: 1]

Jiang MX, Way MO, Zhang WJ, Cheng JA (2007)

Rice water weevil females of different elytral color morphs: comparisons of locomotor activity, mating success, and reproductive capacity

Environmental Entomology, 36, 1040-1047.

DOI:10.1603/0046-225x(2007)36[1040:rwwfod]2.0.co;2      URL     PMID:18284726     

Females and males of the rice water weevil, Lissorhoptrus oryzophilus Kuschel (Coleoptera: Erirhinidae), have two elytral color morphs in Texas: the central pattern of their elytra is black (dark morph) or gray (light morph). In southeast Texas, the dark and light females exhibited similar proportions (28.6 and 32.0%, respectively) in populations collected during the spring. In this study, females of the two morphs were collected near rice fields in southeast Texas during April and May 2005. In the laboratory, light females were more active than dark ones. They mated equally successfully with males, irrespective of morph. For females supplied with males for 2 d or kept solitary, and then reared on rice seedlings for 48 d, no significant differences were found between the two morphs in oviposition period, number of eggs deposited, and survival rate. In both morphs, a proportion of mated females did not oviposit throughout the rearing period, implying that a mating experience might be necessary before reproductive development can be initiated. However, oviposition occurred in a proportion of females in which no mating experience could be detected, and their eggs produced larvae, which suggests the probability of existing parthenogenetic females in southeast Texas.

Jiang MX, Way MO, Du XK, Ji XH, He Y (2008)

Reproductive biology of summer/fall populations of rice water weevil (Lissorhoptrus oryzophilus Kuschel) in southeastern Texas

Southwestern Entomologist, 33, 129-137.

[本文引用: 1]

Jiang MX, Zhang WJ, Cheng JA (2004a)

Reproductive capacity of first-generation adults of the rice water weevil Lissorhoptrus oryzophilus Kuschel (Coleoptera: Curculionidae) in Zhejiang, China

Journal of Pest Science, 77, 145-150.

[本文引用: 1]

Jiang MX, Zhang WJ, Cheng JA (2004b)

Termination of reproductive diapause in the rice water weevil with particular reference to the effects of temperature

Applied Entomology and Zoology, 39, 683-689.

[本文引用: 1]

Jiang S (江舒), Pang L (庞璐), Huang C (黄成) (2007)

Damage and control of an alien species, Procambarus clarkia

Bulletin of Biology (生物学通报), 42(5), 15-16. (in Chinese)

[本文引用: 2]

Jin L, Gu YJ, Xiao M, Chen JK, Li B (2004)

The history of Solidago canadensis invasion and the development of its mycorrhizal associations in newly-reclaimed land

Func- tional Plant Biology, 31, 979-986.

[本文引用: 1]

Jiu M, Zhou XP, Tong L, Xu J, Yang X, Wan FH, Liu SS (2007)

Vector-virus mutualism accelerates population increase of an invasive whitefly

PLoS ONE, 2, e182.

DOI:10.1371/journal.pone.0000182      URL     PMID:17264884      [本文引用: 1]

The relationships between plant viruses, their herbivore vectors and host plants can be beneficial, neutral, or antagonistic, depending on the species involved. This variation in relationships may affect the process of biological invasion and the displacement of indigenous species by invaders when the invasive and indigenous organisms occur with niche overlap but differ in the interactions. The notorious invasive B biotype of the whitefly complex Bemisia tabaci entered China in the late 1990s and is now the predominant or only biotype in many regions of the country. Tobacco curly shoot virus (TbCSV) and Tomato yellow leaf curl China virus (TYLCCNV) are two whitefly-transmitted begomoviruses that have become widespread recently in south China. We compared the performance of the invasive B and indigenous ZHJ1 whitefly biotypes on healthy, TbCSV-infected and TYLCCNV-infected tobacco plants. Compared to its performance on healthy plants, the invasive B biotype increased its fecundity and longevity by 12 and 6 fold when feeding on TbCSV-infected plants, and by 18 and 7 fold when feeding on TYLCCNV-infected plants. Population density of the B biotype on TbCSV- and TYLCCNV-infected plants reached 2 and 13 times that on healthy plants respectively in 56 days. In contrast, the indigenous ZHJ1 performed similarly on healthy and virus-infected plants. Virus-infection status of the whiteflies per se of both biotypes showed limited effects on performance of vectors on cotton, a nonhost plant of the viruses. The indirect mutualism between the B biotype whitefly and these viruses via their host plants, and the apparent lack of such mutualism for the indigenous whitefly, may contribute to the ability of the B whitefly biotype to invade, the displacement of indigenous whiteflies, and the disease pandemics of the viruses associated with this vector.

Jolivet P (1991)

The Colorado potato beetle threatens Asia. Leptinotarsa decemlineata (Say) 1824 (Coleoptera: Chrysomelidae)

Entomologiste, 47, 29-48.

[本文引用: 1]

Ju RT (鞠瑞亭), Li B (李博) (2010)

Sycamore lace bug, Corythucha ciliata, an invasive alien pest rapidly spreading in urban China

Biodiversity Science (生物多样性), 18, 638-646. (in Chinese with English abstract)

DOI:10.3724/SP.J.1003.2010.638      URL     [本文引用: 2]

Ju RT (鞠瑞亭), Li B (李博) (2012)

A risk analysis system for alien species in urban green spaces and application to the 2010 Expo, Shanghai

Biodiversity Science (生物多样性), 20, 12-23. (in Chinese with English abstract)

DOI:10.3724/SP.J.1003.2012.12139      URL     [本文引用: 2]

Invasive species pose risks to regional economies and public health as well as imposing serious damage to natural ecosystems. Urban ecosystems are more susceptible to biological invasions than other ecosystems because they are subjected to more frequent human disturbances. Green space acts as important channels for the spread of alien species into urban ecosystems. Risk analysis can be an effective tool used for preventing invasion of alien species. Although various risk analysis systems have been developed for agricultural and forest ecosystems or nature reserves, the domestic effort to manage risks associated with invasive species in urban green spaces has been limited in China. Therefore, there is a pressing need for formulating scientifically sound methods and approaches for risk analysis in this emerging field. Here, we presented a risk analysis system including risk identification, assessment and management for biological invasions in urban green spaces. Based on risk identification for alien species, we established a risk assessment index that consisted of 4 different layers, i.e. object, item, factor, index layers. A total of 26 index parameters were proposed to assess the risk level, which originated from the risks involved in introduction, colonization, spread and damage of alien species in urban green spaces. In addition, methods for index quantification, weight setting, modeling and risk grading were presented. Applying this system, we analyzed the risks of alien pest species potentially attached to trees brought from Japan to China to green landscapes for the 2010 Expo in Shanghai. Our results showed that 7 species posed a high risk (4 insects, 2 plant pathogens, and 1 plant nematode), 10 species a moderate risk (3 insects, 4 plant pathogens and 3 plant nematodes), and other 2 species (1 insect and 1 nematode) a low or extremely low risk. Based on this analysis, risk management measures were suggested to help local policy-makers set priorities for the management of these alien pests. Monitoring did not reveal the presense of any invasive pests during or after the Expo, suggesting that the risk analysis system may have provided an effective tool for preventing the introduction of invasive alien species and effectively ensuring the ecological security of the 2010 Expo, Shanghai, China.

Ju RT (鞠瑞亭), Peng ZQ (彭正强), Yin LP (印丽萍), Yan W (严巍), Du YZ (杜予州) (2005)

Risk analysis of alien invasive pest, Brontispa longissima (Gestro), in China

Acta Phytophylacica Sinica (植物保护学报), 32, 246-250. (in Chinese with English abstract)

[本文引用: 1]

Ju RT, Wang F, Li B (2011a)

Effects of temperature on the development and population growth of the sycamore lace bug, Corythucha ciliata

Journal of Insect Science, 11, 16.

DOI:10.1673/031.011.0116      URL     PMID:21526932      [本文引用: 1]

The sycamore lace bug, Corythucha ciliata (Say) (Hemiptera: Tingidae), is an important invasive exotic pest of Platanus (Proteales: Platanaceae) trees in China. The objective of this study was to determine the effects of temperature on C. ciliata in the laboratory so that forecasting models based on heat accumulation units could be developed for the pest. Development and fecundity of C. ciliata reared on leaves of London plane tree (Platanus x acerifolia) were investigated at seven constant temperatures (16, 19, 22, 26, 30, 33, and 36 degrees C) and at a relative humidity of 80% with a photoperiod of 14:10 (L:D). The developmental time was found to significantly decrease with increasing temperature. The developmental time from egg hatching to adult emergence was respectively 47.6, 35.0, 24.1, 20.0, and 17.1 days at the temperatures of 19, 22, 26, 30, and 33 degrees C. C. ciliata could not complete full development at 16 degrees and 36 degrees C. The developmental threshold temperature (C) estimated for egg-to-adult was 11.17 degrees C, with a thermal constant of (K) 370.57 degree-days. Longevity of females was found to be the shortest, 17.7 days at 33 degrees C and the longest, 58.9 days at 16 degrees C, and that of males was the shortest, 19.7 days at 33 degrees C and the longest, 59.7 days at 16 degrees C. Fecundity was the highest at 30 degrees C, being 286.8 eggs per female over an oviposition period of 8.9 days. Female lifetime fecundity was reduced at other temperatures, being the lowest (87.7 eggs per female) at 19 degrees C. The population trend index (I) of C. ciliata was the highest (130.1) at 30 degrees C and the lowest (24.9) at 19 degrees C. Therefore, the optimal developmental temperature for C. ciliata was determined to be 30 degrees C.

Ju RT, Chen GB, Wang F, Li B (2011b)

Effects of heat shock, heat exposure pattern, and heat hardening on survival of the sycamore lace bug, Corythucha ciliata (Say) (Hemiptera: Tingidae)

Entomologia Experimentalis et Applicata, 41, 168-177.

Ju RT, Xiao YY, Li B (2011c)

Rapid cold hardening increases cold and chilling tolerances more than acclimation in the adults of the sycamore lace bug, Corythucha ciliata (Say) (Hemiptera: Tingidae)

Journal of Insect Physiology, 57, 1577-1582.

DOI:10.1016/j.jinsphys.2011.08.012      URL     PMID:21872604      [本文引用: 1]

The sycamore lace bug, Corythucha ciliata is a new, invasive pest of Platanus trees in China. Although C. ciliata is often subjected to acute low temperatures in early winter and spring in northern and eastern China, the cold tolerance of C. ciliata has not been well studied. The objectives of this study were to determine whether adults of C. ciliata are capable of rapid cold hardening (RCH), and to compare the benefits of RCH vs. cold acclimation (ACC) in the laboratory. When the adult females incubated at 26 degrees C were transferred directly to the discriminating temperature (-12 degrees C) for 2 h, survival was only 22%. However, exposure to 0 degrees C for 4 h before transfer to -12 degrees C for 2 h induced RCH, i.e., increased survival to 68%. RCH could also be induced by gradual cooling of the insects at rates between 0.1 and 0.25 degrees C min(-1). The protection against cold shock obtained through RCH at 0 degrees C for 4 h was lost within 1h if the adults were returned to 26 degrees C before exposure to -12 degrees C. Survival at both -12 and -5 degrees C was greater for RCH-treated than for ACC-treated adults (for ACC, adults were kept at 15 degrees C for 5 days), and the lethal temperature (2 h exposure) was lower for RCH-treated than for ACC-treated adults. The results suggest that RCH may help C. ciliata survive the acute low temperatures that often occur in early winter and early spring in northern and eastern China.

Ju RT, Xiao YY, Wang F, Li B (2010)

Supercooling capacity and cold hardiness of the adults of the sycamore lace bug, Corythucha ciliata (Hemiptera: Tingidae)

CryoLetters, 31, 445-453.

URL     PMID:21410013     

Supercooling point (SCP) of female adults of Corythucha ciliata was significantly lower than that of male adults, with an average being -11.49 degrees C and -9.54 degrees C, respectively. Low temperature survival of adults of different ages indicated that there were differences in cold survival ability among age groups of adults. Nonlinear regression analysis found that the response of C. ciliata adults to exposure time under different low temperature regimes (above -5 degrees C) was best fitted by a logistic equation. Both low temperature and exposure time had significant effects on mortality of adults. Temperatures above 5 degrees C did not prevent C. ciliata adults from surviving. C. ciliata was shown to be a freeze-intolerant but chill-tolerant insect. C. ciliata could tolerate subzero temperatures by supercooling. Temperature around -8 degres C is a critical point for successful overwintering of C. ciliata adults, which can establish in the whole areas where Platanus trees are planted in China.

Kang L, Chen B, Wei JN, Liu TX (2009)

Roles of thermal adaptation and chemical ecology in Liriomyza distribution and control

Annual Review of Entomology, 54, 127-145.

DOI:10.1146/annurev.ento.54.110807.090507      URL     PMID:18710304      [本文引用: 1]

Many Liriomyza species are pests of agricultural and ornamental plants. In the past two decades, the occurrence and distribution of certain Liriomyza species have changed dramatically, leading to an extensive body of research papers. First, we review the association of thermal tolerance with population dynamics, geographic distribution, and species displacement. Differences in thermal tolerances between species result in their differential geographic locations and overwintering ranges. Displacements among Liriomyza species are associated with their temperature adaptation. We examine the chemical linkage of plants, Liriomyza, and their parasitoids. Chemical compounds from host and nonhost plants mediate the behavior of Liriomyza and their parasitoids. Liriomyza and their parasitoids use chemical cues to locate their hosts. Induced compounds can be used as attractants of parasitoids or repellents of Liriomyza. Thus, understanding the thermal tolerances and chemical ecology of Liriomyza may enable researchers to predict geographic distribution and to develop novel control strategies.

Keay J, Rogers WE, Lankau R, Siemann E (2000)

The role of allelopathy in the invasion of the Chinese tallow tree (Sapium sebiferum)

Texas Journal of Science, 52, 57-64.

[本文引用: 1]

Kong CH, Hu F, Xu T, Lu YH (1999)

Allelopathic potential and chemical constituents of volatile oil from Ageratum conyzoides

Journal of Chemical Ecology, 25, 2347-2356.

[本文引用: 1]

Lavergne S, Muenke NJ, Molofsky J (2010)

Genome size reduction can trigger rapid phenotypic evolution in invasive plants

Annals of Botany, 105, 109-116.

URL     PMID:19887472      [本文引用: 1]

Li B (李博), Hsu PS (徐炳声), Chen JK (陈家宽) (2001)

Perspectives on general trends of plant invasions with special reference to alien weed flora of Shanghai

Biodiversity Science (生物多样性), 9, 446-457. (in Chinese with English abstract)

[本文引用: 1]

Li B (李博), Ma KP (马克平) (2010)

Biological invasions: opportunities and challenges facing Chinese ecologists in the era of translational ecology

Biodiversity Science (生物多样性), 18, 529-532. (in Chinese)

[本文引用: 6]

Li B, Liao CZ, Zhang XD, Chen HL, Wang Q, Chen ZY, Gan XJ, Wu JH, Zhao B, Ma ZJ, Cheng XL, Jiang LF, Chen JK (2009)

Spartina alterniflora invasions in the Yangtze River estuary, China: an overview of current status and ecosystem effects

Ecological Engineering, 35, 511-520.

[本文引用: 5]

Li J, Ye WH (2006)

Genetic diversity of alligator weed ecotypes is not the reason for their different responses to biological control

Aquatic Botany, 85, 155-158.

[本文引用: 2]

Li SJ, Xue X, Ahmed MZ, Ren SX, Du YZ, Wu JH, Cuthbertson AGS, Qiu BL (2011)

Host plants and natural enemies of Bemisia tabaci (Hemiptera: Aleyrodidae) in China

Insect Science, 18, 101-120.

[本文引用: 2]

Li WG, Wang BR, Wang JB (2006)

Lack of genetic variation of an invasive local plant Eichhornia crassipes in China revealed by RAPD and ISSR markers

Aquatic Botany, 84, 176-180.

Li XY (李夕英), Tan JC (谭济才), Song DB (宋东宝), You LS (游兰韶) (2012)

Parasitoids of Bactrocera dorsalis and their application to bio-control

Biological Disaster Science (生物灾害科学), 35, 12-17. (in Chinese with English abstract)

[本文引用: 1]

Li YN (李一农), Li FR (李芳荣), Lou DF (娄定风), Zhang QH (张青汉) (2007)

Biological invasions in Shenzhen and Hong Kong

Plant Quarantine (植物检疫), 21, 29-31. (in Chinese)

[本文引用: 1]

Li ZX (黎镇湘), An SJ (安树杰), Zhang XL (张晓丽), Luo YQ (骆友庆), Yan XS (严晓素), Jiang P (蒋平) (2005)

Principal component analyses of soil in a forest ecological system invaded by Bursaphelenchus xylophilus disease

Chinese Journal of Soil Science (土壤通报), 36, 938-942. (in Chinese with English abstract)

[本文引用: 1]

Liang GH (梁光红), Wu Y (武英), Chen JH (陈家骅) (2006)

Seasonal incidence of Bactrocera dorsalis and its parasitoids in the field

Journal of Southwest Forestry College (西南林学院学报), 26, 72-74. (in Chinese with English abstract)

[本文引用: 1]

Liang KL, Su MC, Shiao JY, Wu SH, Li YH, Jiang RS (2010)

Role of pollen allergy in Taiwanese patients with allergic rhinitis

Journal of the Formosan Medical Association, 109, 879-885.

URL     PMID:21195885      [本文引用: 1]

Liang X, Zheng H, He CQ, Xu QY, Zhan YW, Lei YR, Du W, Yang JN (2012)

Allelopathic effects of invasive Spartina alterniflora root exudates in soil on the offspring (seeds) of Scirpus mariqueter

Allelopathy Journal, 29, 251-261.

[本文引用: 1]

Liao CZ, Luo YQ, Jiang LF, Zhou XH, Wu XW, Chen JK, Li B (2007)

Invasion of Spartina alterniflora enhanced ecosystem carbon and nitrogen stocks in the Yangtze estuary, China

Ecosystems, 10, 1351-1361.

[本文引用: 1]

Liao CZ, Luo YQ, Fang CM, Chen JK, Li B (2008a)

Litter pool sizes, decomposition, and nitrogen dynamics in Spartina alterniflora-invaded and native coastal marshlands of the Yangtze estuary

Oecologia, 156, 589-600.

DOI:10.1007/s00442-008-1007-0      URL     PMID:18327617      [本文引用: 2]

Past studies have focused primarily on the effects of invasive plants on litter decomposition at soil surfaces. In natural ecosystems, however, considerable amounts of litter may be at aerial and belowground positions. This study was designed to examine the effects of Spartina alterniflora invasion on the pool sizes and decomposition of aerial, surficial, and belowground litter in coastal marshlands, the Yangtze Estuary, which were originally occupied by two native species, Scirpus mariqueter and Phragmites australis. We collected aerial and surficial litter of the three species once a month and belowground litter once every 2 months. We used the litterbag method to quantify litter decomposition at the aerial, surficial and belowground positions for the three species. Yearly averaged litter mass in the Spartina stands was 1.99 kg m(-2); this was 250 and 22.8% higher than that in the Scirpus (0.57 kg m(-2)) and Phragmites (1.62 kg m(-2)) stands, respectively. The litter in the Spartina stands was primarily distributed in the air (45%) and belowground (48%), while Scirpus and Phragmites litter was mainly allocated to belowground positions (85 and 59%, respectively). The averaged decomposition rates of aerial, surficial, and belowground litter were 0.82, 1.83, and 1.27 year(-1) for Spartina, respectively; these were 52, 62 and 69% of those for Scirpus litter at corresponding positions and 158, 144 and 78% of those for Phragmites litter, respectively. The differences in decomposition rates between Spartina and the two native species were largely due to differences in litter quality among the three species, particularly for the belowground litter. The absolute amount of nitrogen increased during the decomposition of Spartina stem, sheath and root litter, while the amount of nitrogen in Scirpus and Phragmites litter declined during decomposition for all tissue types. Our results suggest that Spartina invasion altered the carbon and nitrogen cycling in the coastal marshlands of China.

Liao CZ, Peng RH, Luo YQ, Zhou XH, Wu XW, Fang CM, Chen JK, Li B (2008b)

Altered ecosystem carbon and nitrogen cycles by plant invasion: a meta-analysis

New Phytologist, 177, 706-714.

[本文引用: 1]

Lin W, Cheng X, Xu R (2011)

Impact of different economic factors on biological invasions on the global scale

PLoS ONE, 6, e18797.

DOI:10.1371/journal.pone.0018797      URL     PMID:21533203      [本文引用: 1]

Social-economic factors are considered as the key to understand processes contributing to biological invasions. However, there has been few quantified, statistical evidence on the relationship between economic development and biological invasion on a worldwide scale. Herein, using principal factor analysis, we investigated the relationship between biological invasion and economic development together with biodiversity for 91 economies throughout the world. Our result indicates that the prevalence of invasive species in the economies can be well predicted by economic factors (R(2) = 0.733). The impact of economic factors on the occurrence of invasive species for low, lower-middle, upper-middle and high income economies are 0%, 34.3%, 46.3% and 80.8% respectively. Greenhouse gas emissions (CO(2), Nitrous oxide, Methane and Other greenhouse gases) and also biodiversity have positive relationships with the global occurrence of invasive species in the economies on the global scale. The major social-economic factors that are correlated to biological invasions are different for various economies, and therefore the strategies for biological invasion prevention and control should be different.

Lin W, Zhou G, Cheng X, Xu R (2007)

Fast economic development accelerates biological invasions in China

PLoS ONE, 2, e1208.

DOI:10.1371/journal.pone.0001208      URL     PMID:18030342      [本文引用: 1]

Increasing levels of global trade and intercontinental travel have been cited as the major causes of biological invasion. However, indirect factors such as economic development that affect the intensity of invasion have not been quantitatively explored. Herein, using principal factor analysis, we investigated the relationship between biological invasion and economic development together with climatic information for China from the 1970s to present. We demonstrate that the increase in biological invasion is coincident with the rapid economic development that has occurred in China over the past three decades. The results indicate that the geographic prevalence of invasive species varies substantially on the provincial scale, but can be surprisingly well predicted using the combination of economic development (R(2) = 0.378) and climatic factors (R(2) = 0.347). Economic factors are proven to be at least equal to if not more determinant of the occurrence of invasive species than climatic factors. International travel and trade are shown to have played a less significant role in accounting for the intensity of biological invasion in China. Our results demonstrate that more attention should be paid to economic factors to improve the understanding, prediction and management of biological invasions.

Lin JT (林进添) (2003) Biology, Ecology and Control Technologies of Bactrocera dorsalis (桔小实蝇生物学生态学及控制技术研究). PhD dissertation, South China Agricultural University, Guangzhou. (in Chinese)

[本文引用: 1]

Ling B (凌冰), Zhang MX (张茂新), Kong CH (孔垂华), Pang XF (庞雄飞), Liang GW (梁广文) (2003)

Chemical composition of volatile oil from Chromolaena odorata and its effect on plant, fungi and insect growth

Chinese Journal of Applied Ecology (应用生态学报), 14, 744-746. (in Chinese with English abstract)

URL     PMID:12924132      [本文引用: 1]

The volatile oil from C. odorata had a significant effect on the growth of plants, fungi and insects. The result showed that the seedling growth of five test plants was obviously inhibited, in the order of ryegrass > Chinese cabbage > radish > snap been > rice. The inhibitory effect of the oil at its middle concentration (800 mg.L-1) for Pyricularia grisea was the strongest, the next was to Phytophthora nicotianae, and the weakest was to Fusarium axysporum. The inhibitory percentage was 61.40%, 29.27% and 14.44%, respectively. The volatile oil from C. odorata at the dose of 10-20 microliters.plant-1 had a significant oviposition deterrent effect on Phyllotreta striolata and Plutella xylostella. The volatile oil from C. odorata was analyzed by GC/MS and thirty components were identified. Terpenoids compounds were major components of the volatile oil, such as trans-caryophyllene (16.58%), delta-cadinene (15.85%), alpha-copaene (11.58%), caryophyllene oxide (9.63%), germacrene-D (4.96%), and delta-humulene (4.32%).

Liu J, Li M, Li JM, Huang CJ, Zhou XP, Xu FC, Liu SS (2010)

Viral infection of tobacco plants improves the performance of insect viral vector Bemisia tabaci more in its invasive biotype than in an indigenous one

Journal of Zhejiang University (Science B) (Biomedicine & Biotechnology), 11, 30-40.

DOI:10.1631/jzus.B0900213      URL     PMID:20043350      [本文引用: 2]

The ecological effects of plant-virus-vector interactions on invasion of alien plant viral vectors have been rarely investigated. We examined the transmission of Tomato yellow leaf curl China virus (TYLCCNV) by the invasive Q biotype and the indigenous ZHJ2 biotype of the whitefly Bemisia tabaci, a plant viral vector, as well as the influence of TYLCCNV-infection of plants on the performance of the two whitefly biotypes. Both whitefly biotypes were able to acquire viruses from infected plants and retained them in their bodies, but were unable to transmit them to either tobacco or tomato plants. However, when the Q biotype fed on tobacco plants infected with TYLCCNV, its fecundity and longevity were increased by 7- and 1-fold, respectively, compared to those of the Q biotype fed on uninfected tobacco plants. When the ZHJ2 biotype fed on virus-infected plants, its fecundity and longevity were increased by only 2- and 0.5-fold, respectively. These data show that the Q biotype acquired higher beneficial effects from TYLCCNV-infection of tobacco plants than the ZHJ2 biotype. Thus, the Q biotype whitefly may have advantages in its invasion and displacement of the indigenous ZHJ2 biotype.

Liu J, Liang SC, Liu FH, Wang RQ, Dong M (2005)

Invasive alien plant species in China: regional distribution patterns

Diversity and Distributions, 11, 341-347.

[本文引用: 1]

Liu J, Zhao H, Jiang K, Zhou XP, Liu SS (2009)

Differential indirect effects of two plant viruses on an invasive and an indigenous whitefly vector: implications for competitive displacement

Annals of Applied Biology, 155, 439-448.

[本文引用: 1]

Liu SS, De Barro PJ, Xu J, Luan JB, Zang LS, Ruan YM, Wan FH (2007)

Asymmetric mating interactions drive widespread invasion and displacement in a whitefly

Science, 318, 1769-1772.

DOI:10.1126/science.1149887      URL     PMID:17991828      [本文引用: 2]

The role of behavioral mechanisms in animal invasions is poorly understood. We show that asymmetric mating interactions between closely related but previously allopatric genetic groups of the whitefly Bemisia tabaci, a haplodiploid species, have been a driving force contributing to widespread invasion and displacement by alien populations. We conducted long-term field surveys, caged population experiments, and detailed behavioral observations in Zhejiang, China, and Queensland, Australia, to investigate the invasion process and its underlying behavioral mechanisms. During invasion and displacement, we found increased frequency of copulation leading to increased production of female progeny among the invader, as well as reduced copulation and female production in the indigenous genetic groups. Such asymmetric mating interactions may be critical to determining the capacity of a haplodiploid invader and the consequences for its closely related indigenous organisms.

Liu WX, Niu HB, Wan FH, Liu B (2010)

Effects of leachates of the invasive plant, Ageratina adenophora (Sprengel) on soil microbial community

Acta Ecologica Sinica, 30, 196-200.

Lockwood JL, Hoopes MF, Marchetti MP (2007) Invasion Ecology. Blackwell Publishing, Malden, MA, USA.

[本文引用: 1]

Lu JB, Wu JG, Fu ZH, Zhu L (2007a)

Water hyacinth in China: a sustainability science-based management framework

Environmental Management, 40, 823-830.

DOI:10.1007/s00267-007-9003-4      URL     PMID:17768654      [本文引用: 2]

The invasion of water hyacinth (Eichhornia crassipes) has resulted in enormous ecological and economic consequences worldwide. Although the spread of this weed in Africa, Australia, and North America has been well documented, its invasion in China is yet to be fully documented. Here we report that since its introduction about seven decades ago, water hyacinth has infested many water bodies across almost half of China's territory, causing a decline of native biodiversity, alteration of ecosystem services, deterioration of aquatic environments, and spread of diseases affecting human health. Water hyacinth infestations have also led to enormous economic losses in China by impeding water flows, paralyzing navigation, and damaging irrigation and hydroelectricity facilities. To effectively control the rampage of water hyacinth in China, we propose a sustainability science-based management framework that explicitly incorporates principles from landscape ecology and Integrated Pest Management. This framework emphasizes multiple-scale long-term monitoring and research, integration among different control techniques, combination of control with utilization, and landscape-level adaptive management. Sustainability science represents a new, transdisciplinary paradigm that integrates scientific research, technological innovation, and socioeconomic development of particular regions. Our proposed management framework is aimed to broaden the currently dominant biological control-centered view in China and to illustrate how sustainability science can be used to guide the research and management of water hyacinth.

Lu JB, Zhu L, Hu GL, Wu JG (2010)

Integrating animal manure-based bioenergy production with invasive species control: a case study at Tongren Pig Farm in China

Biomass and Bioenergy, 34, 821-827.

Lu M, Miller D, Sun JH (2007b)

Cross-attraction between an exotic and a native pine bark beetle: a novel invasion mechanism?

PLoS ONE, 12, e1302.

[本文引用: 1]

Lu M, Wingfield MJ, Gillette NE, Mori SR, Sun JH (2010)

Complex interactions among host pines and fungi vectored by an invasive bark beetle

New Phytologist, 187, 859-866.

[本文引用: 1]

Lu M, Wingfield MJ, Gillette NE, Sun JH (2011)

Do novel genotypes drive the success of an invasive bark beetle-fungus complex? Implications for potential reinvasion

Ecology, 92, 2013-2019.

DOI:10.1890/11-0687.1      URL     PMID:22164824      [本文引用: 1]

Novel genotypes often arise during biological invasions, but their role in invasion success has rarely been elucidated. Here we examined the population genetics and behavior of the fungus, Leptographium procerum, vectored by a highly invasive bark beetle, Dendroctonus valens, to determine whether genetic changes in the fungus contributed to the invasive success of the beetle-fungal complex in China. The fungus was introduced by the beetle from the United States to China, where we identified several novel genotypes using microsatellite markers. These novel genotypes were more pathogenic to Chinese host seedlings than were other genotypes and they also induced the release of higher amounts of 3-carene, the primary host attractant for the beetle vector, from inoculated seedlings. This evidence suggests a possible mechanism, based on the evolution of a novel genotype during the two or three decades since its introduction, for the success of the beetle-fungal complex in its introduced region.

Lu M, Zhou XD, De Beer ZW, Wingfield MJ, Sun JH (2009)

Ophiostomatoid fungi associated with the invasive pine-infesting bark beetle, Dendroctonus valens, in China

Fungal Diversity, 38, 133-145.

[本文引用: 1]

Lu YY (陆永跃), Zeng L (曾玲) (2004)

The invasion pathway of Brontispa longissima (Gestro) to China and its causes

Forest Pest and Disease (中国森林病虫), 23, 12-15. (in Chinese with English abstract)

[本文引用: 2]

Lu Z, Ma K (2005)

Scale dependent relationships between native plant diversity and the invasion of croftonweed ( Eupatorium adenophorum) in southwest China

Weed Science, 53, 600-604.

[本文引用: 2]

Luo C, Jones CM, Devine G, Zhang F, Denholm I, Gorman K (2010)

Insecticide resistance in Bemisia tabaci biotype Q (Hemiptera: Aleyrodidae) from China

Crop Protection, 29, 429-434.

[本文引用: 2]

Luo QQ (罗卿权), Ju RT (鞠瑞亭), Xu Y (徐颖), Wang F (王凤) (2011)

Development and application of PDA pest real-time monitoring system of urban green space based on Web Service

Acta Phytophylacica Sinica (植物保护学报), 38, 9-14. (in Chinese with English abstract)

[本文引用: 1]

BQ, Tang C, Peng ZQ, Salle JL, Wan FH (2008)

Biological assessment in quarantine of Asecodes hispinarum Bouček (Hymenoptera: Eulophidae) as an imported biological control agent of Brontispa longissima (Gestro) (Coleoptera: Hispidae) in Hainan, China

Biological Control, 45, 29-35.

[本文引用: 1]

Ma RY (马瑞燕), Wang R (王韧) (2004)

Effect of morphological and physiological variations in the ecotypes of alligator weed, Alternanthera philoxerodes on the pupation rate of its biocontrol agent Agasicles hygrophila

Chinese Journal of Plant Ecology (植物生态学报), 28, 24-30. (in Chinese with English abstract)

[本文引用: 1]

Ma ZJ, Wang Y, Gan XJ, Li B, Jing K, Tang SM, Chen JK (2009)

Waterbird population changes in the wetlands at Chongming Dongtan in the Yangtze River estuary, China

Environmental Management, 43, 1187-1200.

DOI:10.1007/s00267-008-9247-7      URL     PMID:19139954      [本文引用: 1]

We studied the changes in wetland habitats and waterbird communities between the 1980s and the 2000s at Chongming Dongtan, a Ramsar site in the Yangtze River estuary, an ecologically important region. This region is an important stopover site for shorebirds along the East Asian-Australasian flyway and is extensively used by waterfowl. A net loss of 11% of the wetland area was estimated during study periods at Chongming Dongtan. The change was dependent on wetland types: while the area of artificial habitats such as paddy fields and aquacultural ponds more than doubled, more than 65% of natural habitats including sea bulrush (Scirpus mariqueter) and common reed (Phragmites australis) marshes were lost over the two decades. An exotic plant species introduced from North America, smooth cordgrass (Spartina alterniflora), occupied 30% of the vegetated intertidal zone by the 2000s. Although waterbird species richness did not change between the 1980s (110) and the 2000s (111), 13 species found in 1980s were replaced by 14 newly recorded species. Moreover, there were more species with declining trends (58) than with increasing trends (19). The population trends of species were affected by residential status and habitat types. Transients, wintering migrants, and habitat specialists were more likely to show declining trends compared to those breeding at Dongtan (including year-round and summer residents) and habitat generalists. Furthermore, species associated mainly with natural wetlands were more likely to decline than those associated mainly with artificial wetlands. These patterns suggest that the loss and change of wetland habitats at Chongming Dongtan adversely affected local population dynamics and might have contributed to the global decline of some waterbird species. Because Chongming Dongtan provides stopover and wintering habitats for many migratory waterbirds, protection and restoration of natural wetlands at Chongming Dongtan are urgently needed.

Millennium Ecosystem Assessment (2005) Ecosystems and Human Well-being: Synthesis, Island Press, Washington, DC.

[本文引用: 1]

Moles AT, Gruber MAM, Bonser SP (2008)

A new frame work for predicting invasive plant species

Journal of Ecology, 96, 13-17.

[本文引用: 1]

Mooney HA, Hobbs RJ (2000) Invasive Species in a Changing World. Island Press, Washington, DC.

[本文引用: 1]

Nie CR (聂呈荣), Li HS (黎华寿), Huang JH (黄京华), Chen LQ (陈莉琼), Hong MQ (洪明祺) (2002)

The allelopanhic effects of Wedelia chinensis Merr on peanut (Arachis hyogaea L) and other crops

Journal of Peanut Science (花生学报), 31, 30-32. (in Chinese with English abstract)

[本文引用: 1]

Nie M, Wang M, Li B (2009)

Effects of saltmarsh invasion by Spartina alterniflora on sulfate-reducing bacteria in the Yangtze River estuary

Ecological Engineering, 35, 1804-1808.

[本文引用: 1]

Niu HB, Liu WX, Wan FH, Liu B (2007)

An invasive aster ( Ageratina adenophora) invaders and dominates forest understories in China: altered soil microbial communities facilitate the invader and inhibit natives

Plant and Soil, 294, 73-85.

[本文引用: 5]

Niu YF, Feng YL, Xie JL, Luo FC (2010)

Noxious invasive Eupatorium adenophorum may be a moving target: implications of the finding of a native natural enemy, Dorylus orientalis

Chinese Science Bulletin, 55, 3743-3745.

[本文引用: 1]

Nuñex MA, Pauchard A (2010)

Biological invasions in developing and developed countries: Does on model fit all?

Biological Invasions, 12, 707-714.

[本文引用: 1]

Pan HP, Chu D, Ge DQ, Wang SL, Wu QJ, Xie W, Jiao XG, Liu BM, Yang X, Yang NN, Su Q, Xu BY, Zhang YJ (2011)

Further spread of and domination by Bemisia tabaci biotype Q on field crops in China

Journal of Economic Entomology, 104, 978-985.

DOI:10.1603/ec11009      URL     PMID:21735919      [本文引用: 4]

The sweetpotato whitefly, Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae), causes severe crop losses to many crops. The worst of these losses are often associated with the invasion and establishment of biotypes B and Q of this pest. Previous research in 2007 showed that biotype Q occurred with other biotypes in most field populations in China. To determine the current status of the biotype composition in the field, an extensive survey covering mainly eastern parts of China was conducted in 2009. Using polymerase chain reaction primers specific for the mitochondrial cytochrome oxidase I of biotypes B and Q and gene sequencing, we determined the biotypes composition in 61 whitefly populations and their distribution across 19 provinces in China. Our research revealed that only biotypes B and Q have been found in the field in 2009 in China. Among them, biotype Q was dominant in 44 locations (100.0%) and biotype B was dominant in 17 locations (100.0%). The current survey indicates that biotype Q has rapidly displaced biotype B in most locations in China.

Pan HP, Chu D, Yan WQ, Su J, Liu BM, Wang SL, Wu QJ, Xie W, Jiao XG, Li RM, Yang NA, Yang X, Xu BY, Brown JK, Zhou XG, Zhang YJ (2012)

Rapid spread of Tomato yellow leaf curl virus in China is aided differentially by two invasive whiteflies

PLoS ONE, 7, e34817.

DOI:10.1371/journal.pone.0034817      URL     PMID:22514670      [本文引用: 1]

BACKGROUND: Tomato yellow leaf curl virus (TYLCV) was introduced into China in 2006, approximately 10 years after the introduction of an invasive whitefly, Bemisia tabaci (Genn.) B biotype. Even so the distribution and prevalence of TYLCV remained limited, and the economic damage was minimal. Following the introduction of Q biotype into China in 2003, the prevalence and spread of TYLCV started to accelerate. This has lead to the hypothesis that the two biotypes might not be equally competent vectors of TYLCV. METHODOLOGY/PRINCIPAL FINDINGS: The infection frequency of TYLCV in the field-collected B. tabaci populations was investigated, the acquisition and transmission capability of TYLCV by B and Q biotypes were compared under the laboratory conditions. Analysis of B. tabaci populations from 55 field sites revealed the existence of 12 B and 43 Q biotypes across 18 provinces in China. The acquisition and transmission experiments showed that both B and Q biotypes can acquire and transmit the virus, however, Q biotype demonstrated superior acquisition and transmission capability than its B counterparts. Specifically, Q biotype acquired significantly more viral DNA than the B biotype, and reached the maximum viral load in a substantially shorter period of time. Although TYLCV was shown to be transmitted horizontally by both biotypes, Q biotype exhibited significantly higher viral transmission frequency than B biotype. Vertical transmission result, on the other hand, indicated that TYLCV DNA can be detected in eggs and nymphs, but not in pupae and adults of the first generation progeny. CONCLUSIONS/SIGNIFICANCE: These combined results suggested that the epidemiology of TYLCV was aided differentially by the two invasive whiteflies (B and Q biotypes) through horizontal but not vertical transmission of the virus. This is consistent with the concomitant eruption of TYLCV in tomato fields following the recent rapid invasion of Q biotype whitefly in China.

Pan XY (潘晓云), Geng YP (耿宇鹏), Sosa A, Zhang WJ (张文驹), Li B (李博), Chen JK (陈家宽) (2007)

Invasive Alternanthera philoxeroides: biology, ecology and management

Acta Phytotaxonomica Sinica (植物分类学报), 45, 884-900. (in Chinese with English abstract)

[本文引用: 1]

Pan XY, Geng YP, Zhang WJ, Li B, Chen JK (2006)

The influence of abiotic stress and phenotypic plasticity on the distribution of invasive Alternanthera philoxeroides along a riparian zone

Acta Oecologica, 30, 333-341.

[本文引用: 2]

Pan XY, Geng YP, Li B, Chen JK (2009)

Effect of root fragment length and planting depth on clonal establishment of alligator weed

Journal of Aquatic Plant Management, 47, 96-100.

[本文引用: 1]

Pan XY, Jia X, Zeng J, Sosa A, Li B, Chen JK (2011)

Stem tissue mass density is linked to growth and resistance to a stem-boring insect in Alternanthera philoxeroides

Plant Species Biology, 26, 58-65.

Pan XY, Villamagna AM, Li B (2011)

Eichhornia crassipes Mart. [Solms-Laubach] (water hyacinth)

In: A Handbook of Global Freshwater Invasive Species (ed. Francis RA), pp. 47-56. Earthscan, Oxon.

Peng ZQ (彭正强), Tang C (唐超), Qin WQ (覃伟权), Ma ZL (马子龙) (2008)

Biological control of Brontispa longissima (Gestro) (Coleoptera: Hispidae)

In: Biological Invasions: Biological Control Theory and Practice (生物入侵: 生物防治篇) (eds Wan FH (万方浩), Li BP (李保平), Guo JY (郭建英)), pp. 478-503. Science Press, Beijing. (in Chinese)

[本文引用: 1]

Perrings C, Mooney HA, Williamson M (2010) Bioinvasions and Globalization: Ecology, Economics, Management, and Policy. Oxford University Press, Oxford.

[本文引用: 2]

Pulliam R, Yu H (余华) (2005)

Challenges and strategies for the development of ecology in 21st century

In: Challenges Facing Ecologists: Questions and Approaches (生态学家面临的挑战: 问题与途径) (eds Chen JQ (陈吉泉), Li B (李博), Ma ZJ (马志军), Zhao B (赵斌)), pp. 9-15. Higher Education Press, Beijing. (in Chinese)

[本文引用: 1]

Pyšek P, Hulme PE (2010)

Biological Invasions in Europe 50 Years after Elton: Time to Sound the Alarm

In: Fifty Years of Invasion Ecology: the Legacy of Charles Elton (ed. Richardson DM). Wiley-Blackwell, Oxford, UK. doi: 10.1002/9781444329988.ch7.

[本文引用: 1]

Qin WQ (覃伟权), CJ (吕朝军), Li CX (李朝绪), Huang SC (黄山春), Peng ZQ (彭正强) (2010)

Research of pests of coconut in China

Chinese Agricultural Science Bulletin (中国农学通报), 26, 200-204. (in Chinese with English abstract)

[本文引用: 1]

Qiu BL, Liu L, Li XX, Mathur V, Qin ZQ, Ren SX (2009)

Genetic mutations associated with chemical resistance in the cytochrome P450 genes of invasive and native Bemisia tabaci (Hemiptera: Aleyrodidae) populations in China

Insect Science, 16, 237-245.

[本文引用: 1]

Qu B (曲波), Wang XZ (王学治), GZ (吕国忠), Zhai Q (翟强), Chen XH (陈旭辉), Yang H (杨红) (2011)

Microscopic and ultramicroscopic investigation into the infection process of giant ragweed rust (Puccinia xanthii f. sp. ambrosiae-trifidae)

Journal of Biosafety (生态安全学报), 20, 317-320. (in Chinese with English abstract)

[本文引用: 1]

Ren L (任璐), Zeng L (曾玲), Lin JT (林进添), Lu YY (陆永跃), Liu CY (刘春燕) (2008)

Biological control of Bactrocera dorsalis (Hendel) (Diptera: Trypetidae)

In: Biological Invasions: Biological Control Theory and Practice (生物入侵: 生物防治篇) (eds Wan FH (万方浩), Li BP (李保平), Guo JY (郭建英)), pp. 413-440. Science Press, Beijing. (in Chinese)

[本文引用: 1]

Ren MX, Zhang QG (2007)

Clonal diversity and structure of the invasive aquatic plant Eichhornia crassipes in China

Aquatic Botany, 87, 242-246.

[本文引用: 1]

Ren SX (任顺祥), Qiu BL (邱宝利), Ge F (戈峰), Zhang YJ (张友军), Du YZ (杜予州), Chen XX (陈学新), Guo JY (郭建英) (2011)

Research progress of the monitoring, forecast and sustainable management of whitefly pests in China

Chinese Journal of Applied Entomology, 48, 7-15.

[本文引用: 1]

Richards CL, Pennings SC, Donovan LA (2005)

Habitat range and phenotypic variation in salt marsh plants

Plant Ecology, 176, 263-273.

DOI:10.1007/s11258-004-0841-3      URL     [本文引用: 1]

Ecologists have long speculated that species with wider environmental ranges would have broader ranges in phenotype; however, most tests of this hypothesis have involved small numbers of species and/or closely related taxa. We related phenotypic variation in twelve salt marsh plant species from six families to variation in four environmental variables using multiple regression. Within species, plant phenotype was predictably related to environmental variation. Salinity was the most common predictor of plant traits, followed by organic content, water content and elevation. Across species, regressions of single plant trait CVs on range (2 × SD) of single environmental variables were not significant and did not support the hypothesis that species occupying broad environmental ranges would have broad ranges in phenotypes. However, regression of a composite phenotypic PCA1 on a composite environmental PCA1 showed a marginally significant (P = 0.054). linear relationship for 10 species. Considering the different patterns of response across species, the lack of a relationship between variation in single phenotypic traits and single environmental variables is likely because the distantly-related taxa employed fundamentally different morphological and physiological strategies to respond to environmental stress gradients. The significant relationship between composite environmental and phenotypic variables reflects the complex nature of species phenotypic response to multivariate environmental gradients. Specifically, in this system, species increase variation in the number of leaves, but decrease variation in leaf size in response to an increase in range of salinity and decrease in range of water and organic content.]]>

Richardson DM, Allsopp N, D’Antonio CM, Milton SJ, Rejamánek M (2000)

Plant invasions―the role of mutual- lisms

Biological Reviews, 75, 65-93.

DOI:10.1017/s0006323199005435      URL     PMID:10740893      [本文引用: 1]

Many introduced plant species rely on mutualisms in their new habitats to overcome barriers to establishment and to become naturalized and, in some cases, invasive. Mutualisms involving animal-mediated pollination and seed dispersal, and symbioses between plant roots and microbiota often facilitate invasions. The spread of many alien plants, particularly woody ones, depends on pollinator mutualisms. Most alien plants are well served by generalist pollinators (insects and birds), and pollinator limitation does not appear to be a major barrier for the spread of introduced plants (special conditions relating to Ficus and orchids are described). Seeds of many of the most notorious plant invaders are dispersed by animals, mainly birds and mammals. Our review supports the view that tightly coevolved, plant-vertebrate seed dispersal systems are extremely rare. Vertebrate-dispersed plants are generally not limited reproductively by the lack of dispersers. Most mycorrhizal plants form associations with arbuscular mycorrhizal fungi which, because of their low specificity, do not seem to play a major role in facilitating or hindering plant invasions (except possibly on remote islands such as the Galapagos which are poor in arbuscular mycorrhizal fungi). The lack of symbionts has, however, been a major barrier for many ectomycorrhizal plants, notably for Pinus spp. in parts of the southern hemisphere. The roles of nitrogen-fixing associations between legumes and rhizobia and between actinorhizal plants and Frankia spp. in promoting or hindering invasions have been virtually ignored in the invasions literature. Symbionts required to induce nitrogen fixation in many plants are extremely widespread, but intentional introductions of symbionts have altered the invasibility of many, if not most, systems. Some of the world's worst invasive alien species only invaded after the introduction of symbionts. Mutualisms in the new environment sometimes re-unite the same species that form partnerships in the native range of the plant. Very often, however, different species are involved, emphasizing the diffuse nature of many (most) mutualisms. Mutualisms in new habitats usually duplicate functions or strategies that exist in the natural range of the plant. Occasionally, mutualisms forge totally novel combinations, with profound implications for the behaviour of the introduced plant in the new environment (examples are seed dispersal mutualisms involving wind-dispersed pines and cockatoos in Australia; and mycorrhizal associations involving plant roots and fungi). Many ecosystems are becoming more susceptible to invasion by introduced plants because: (a) they contain an increasing array of potential mutualistic partners (e.g. generalist frugivores and pollinators, mycorrhizal fungi with wide host ranges, rhizobia strains with infectivity across genera); and (b) conditions conductive for the establishment of various alien/alien synergisms are becoming more abundant. Incorporating perspectives on mutualisms in screening protocols will improve (but not perfect) our ability to predict whether a given plant species could invade a particular habitat.

Sang W, Zhu L, Jan C, Axmacher (2010)

Invasion pattern of Eupatorium adenophorum Spreng in southern China

Biological Invasions, 12, 1721-1730.

DOI:10.1007/s10530-009-9584-3      URL     [本文引用: 1]

Eupatorium adenophorum Spreng (Crofton weed) from Burma and Vietnam into Southern China since the 1940s. Currently, E. adenophorum’s main colonisation area is located in the Yunnan–Guizhou Plateau of China, where it has caused prominent economic and ecological problems. Sixty-three years ago, Crofton weed appeared in Menghai county, Yunnan Province, from where it dispersed northwards and eastwards at an average speed of 20 km year−1. The invasion of E. adenophorum nonetheless showed pronounced variations in both time and space. Spread was relatively slow in the initial invasion period between 1940 and 1950, while the most rapid range expansions occurred in the 1980s. Environmental conditions at native and invaded sites were significantly different, reflecting a great adaptability of the species during colonization. These changes were greater than habitat differences between colonized and many adjacent un-colonized sites in Southern China. Therefore, immediate measures are required to stop a further northward and eastward expansion of Crofton weed.]]>

Schlaepfer DR, Edwards PJ, Semple JC, Billeter R (2008)

Cytogeography of Solidago gigantea (Asteraceae) and its invasive ploidy level

Journal of Biogeography, 35, 2119-2127.

[本文引用: 1]

Schlesinger WH (2010)

Translational ecology

Science, 329, 609.

DOI:10.1126/science.1195624      URL     PMID:20688985      [本文引用: 1]

Shang L (尚蕾), Yangjin Z (央金卓嘎), Yang J (杨继), Li B (李博) (2010)

Genomics: an important tool for understand- ing plant invasiveness

Biodiversity Science (生物多样性), 18, 533-546. (in Chinese with English abstract)

[本文引用: 1]

Shao H, Peng SL, Wei XY, Zhang DQ, Zhang C (2005)

Potential allelochemicals from an invasive weed Mikania micrantha H. B. K

Journal of Chemical Ecology, 31, 1657-1668.

DOI:10.1007/s10886-005-5805-0      URL     PMID:16222800      [本文引用: 1]

Phytotoxicity-directed extraction and fractionation of the aerial parts of Mikania micrantha H.B.K. led to the isolation and identification of three sesquiterpenoids: dihydromikanolide, deoxymikanolide, and 2,3-epoxy-1-hydroxy-4,9-germacradiene-12,8:15,6-diolide. These sesquiterpenoids inhibited both germination and seedling growth of tested species with deoxymikanolide possessing the strongest phytotoxicity. In a bioassay against lettuce (Lectuca sativa L.), deoxymikanolide reduced radicle elongation at low concentration (IC50 = 47 microg/ml); dihydromikanolide showed a weaker effect (IC50 = 96 microg/ml), and 2,3-epoxy-1-hydroxy-4,9-germacradiene-12,8:15,6-diolide exhibited the least effect (IC50 = 242 microg/ml). Deoxymikanolide caused yellowish lesions at the root tips of lettuce at a concentration of 50 microg/ml, and a 250 microg/ml solution killed lettuce seedlings. A bioassay against the monocot ryegrass (Lolium multiforum) revealed similar results on radicle elongation, which implied that the growth inhibition by these compounds was not selective. To evaluate their phytotoxicity to plants in natural habitats, three common companion tree species in south China, Acacia mangium, Eucalyptus robusta, and Pinus massoniana, were also tested and similar results were obtained. This is the first report on the isolation of 2,3-epoxy-1-hydroxy-4,9-germacradiene-12,8:15,6-diolide as a naturally occurring product.

Shen JR (申建茹), Liu WX (刘万学), Wan FH (万方浩), Zhang FQ (张芬琴) (2012)

Characterization of the Cydia pomonella granulovirus CpGV-CJO1 from northwest China

Chinese Journal of Applied Entomology (应用昆虫学报), 49, 96-103. (in Chinese with English abstract)

[本文引用: 1]

Shi L (石磊), Chen M (陈明), Luo JC (罗进仓) (2009)

Efficiency of three types of pheromone traps in capturing adult codling moth Laspeyresia pomonella, and its development

Journal of Gansu Agricultural University (甘肃农业大学学报), 44, 115-117. (in Chinese with English abstract)

[本文引用: 1]

Shi SW, He Y, Ji XH, Jiang MX, Cheng JA (2008)

Do rice water weevils and rice stem borers compete when sharing a host plant?

Journal of Zhejiang University: Science B, 9, 572-577.

DOI:10.1631/jzus.B0820009      URL     PMID:18600788      [本文引用: 1]

The rice water weevil (RWW) Lissorhoptrus oryzophilus Kuschel (Coleoptera: Curculionidae) is an invasive insect pest of rice Oryza sativa L. in China. Little is known about the interactions of this weevil with indigenous herbivores. In the present study, adult feeding and population density of the weevil, injury level of striped stem borer Chilo suppressalis (Walker) (Lepidoptera: Pyralidae) and pink stem borer Sesamia inferens (Walker) (Lepidoptera: Noctuidae) to rice, as well as growth status of their host plants were surveyed in a rice field located in Southeastern Zhejiang, China, in 2004 with the objective to discover interspecific interactions on the rice. At tillering stage, both adult feeding of the weevil and injury of the stem borers tended to occur on larger tillers (bearing 5 leaves) compared with small tillers (bearing 2~4 leaves), but the insects showed no evident competition with each other. At booting stage, the stem borers caused more withering/dead hearts and the weevil reached a higher density on the plants which had more productive tillers and larger root system; the number of weevils per tiller correlated negatively with the percentage of withering/dead hearts of plants in a hill. These observations indicate that interspecific interactions exist between the rice water weevil and the rice stem borers with negative relations occurring at booting or earlier developmental stages of rice.

Shi W (施雯), Geng YP (耿宇鹏), Ou XK (欧晓昆) (2010)

Genetic diversity and invasion success of alien species: where are we and where should we go?

Biodiversity Science (生物多样性), 18, 590-597. (in Chinese with English abstract)

[本文引用: 1]

Shih CJ (石正人), Yang CC (杨景程), Wu WJ (吴文哲) (2007)

Control of Solenopsis invicta in Taiwan

Taiwan Entomology (台湾昆虫), 9, 43-54. (in Chinese with English abstract)

[本文引用: 1]

Shoemaker DD, Wu JC, Lin YK, Lin CC, Wu WJ, Shih CJ (2009)

Successful establishment of the invasive fire ant Solenopsis invicta in Taiwan: insights into interactions of alternate social forms

Diversity and Distributions, 15, 709-719.

Song ZD (宋侦东), Lu YY (陆永跃), Xu YJ (许益镌), Huang J (黄俊), Liang GW (梁广文), Zeng L (曾玲) (2010)

Dynamic of native ants on the lawn with the invasion of Solenopsis invicta Buren

Acta Ecologica Sinica (生态学报), 30, 1287-1295. (in Chinese with English abstract)

[本文引用: 1]

Tang L, Gao Y, Wang CH, Wang JQ, Li B, Chen JK, Zhao B (2010)

How tidal regime and treatment timing influence the clipping frequency for controlling invasive Spartina alterniflora: implications for reducing management costs

Biological Invasions, 12, 593-601.

[本文引用: 1]

Teng X, Wan FH, Chu D (2010)

Bemisia tabaci biotype Q dominates other biotypes across China

Florida Entomo- logist, 93, 363-368.

[本文引用: 2]

van der Putten WH, Macel M, Visser ME (2010)

Predicting species distribution and abundance responses to climate change: why it is essential to include biotic interactions across trophic levels

Philosophical Transactions of the Royal Society B: Biological Sciences, 365, 2025-2034.

[本文引用: 1]

Wan FH (万方浩), Li BP (李保平), Guo JY (郭建英) (2008a) Biological Invasions: Biological Control Theory and Practice (生物入侵: 生物防治篇). Science Press, Beijing. (in Chinese)

[本文引用: 4]

Wan FH (万方浩), Xie BY (谢丙炎), Chu D (褚栋) (2008b) Biological Invasions: Legislations and Management Strategies (生物入侵: 管理篇). Science Press, Beijing. (in Chinese)

[本文引用: 3]

Wan FH (万方浩), Guo JY (郭建英), Zhang F (张峰) (2009) Research on Biological Invasions in China (中国生物入侵研究). Science Press, Beijing. (in Chinese)

[本文引用: 8]

Wan FH, Zhang GF, Liu SS, Luo C, Chu D, Zhang YJ, Zang LS, Jiu M, ZC, Cui XH, Zhang LP, Zhang F, Zhang QW, Liu WX, Liang P, Lei ZR, Zhang YJ (2009)

Invasive mechanism and management strategy of Bemisia tabaci (Gennadius) biotype B: progress report of 973 Program on invasive alien species in China

Science in China, Series C: Life Sciences, 52, 88-95.

DOI:10.1007/s11427-008-0135-4      URL     PMID:19152088     

Bemisia tabaci (Gennadius) biotype B, called a

Wan FH (万方浩), Peng DL (彭德良), Wang R (王瑞) (2010) Biological Invasions: Risk Analysis and Early Prevention (生物入侵: 预警篇). Science Press, Beijing. (in Chinese)

[本文引用: 4]

Wan FH, Liu WX, Guo JY, Qiang S, Li BP, Wang JJ, Yang GQ, Niu HB, Gui FR, Huang WK, Jiang ZL, Wang WQ (2010)

Invasive mechanism and control strategy of Ageratina adenophora (Sprengel)

Science in China, Series C: Life Sciences, 53, 1291-1298.

Wan FH (万方浩), Xie BY (谢丙炎), Yang GQ (杨国庆) (2011a) Invasion Biology (入侵生物学). Science Press, Beijing. (in Chinese)

[本文引用: 5]

Wan FH (万方浩), Feng J (冯洁), Xu J (徐进) (2011b) Biological Invasions: Detection, Surveillance and Monitoring (生物入侵: 检测与监测篇). Science Press, Beijing. (in Chinese)

[本文引用: 2]

Wan FH (万方浩), Yan Y (严盈), Wang R (王瑞), Yang GQ (杨国庆) (2011

c) Invasion biology: development and perspective in China

Journal of Biosafety (生物安全学报), 20, 1-19. (in Chinese with English abstract)

[本文引用: 5]

Wang FG (王发国), Xing FW (邢福武), Ye HG (叶华谷), Chen XY (陈孝永), Tan GG (谭国光), Mai BL (麦保林) (2004)

Preliminarily study on invasive alien species in Macau

Acta Scientiarum Naturalium Universitatis Sunyatseni (中山大学学报), 43, 105-110. (in Chinese with English abstract)

[本文引用: 1]

Wang FX (王福祥), Liu H (刘慧), Yang H (杨桦), Xu J (徐婧), Zhang RZ (张润志) (2012)

The monitoring and control of codling moth in pest-free apple producing areas of China

Chinese Journal of Applied Entomology (应用昆虫学报), 49, 275-280. (in Chinese with English abstract)

[本文引用: 1]

Wang HF, López-Pujol J, Meyerson LA, Qiu JX, Wang XK, Ouyang ZY (2011)

Biological invasions in rapidly urbanizing areas: a case study of Beijing, China

Biodive- rsity and Conservation, 20, 2483-2509.

[本文引用: 2]

Wang L (王兰), Feng HZ (冯宏祖), Guo WC (郭文超), Zhang H (张辉), Yang L (杨力), Xu JJ (许建军) (2011)

Dynamic state of Laspeyresia pomonella and release techniques of Trichogramma in orchards

Xinjiang Agricultural Sciences (新疆农业科学), 48, 261-265. (in Chinese with English abstract)

[本文引用: 2]

Wang M, Chen JK, Li B (2007)

Characterization of bacterial community structure and diversity in rhizosphere soils of three plants in rapidly changing salt marshes using 16S rDNA

Pedosphere, 17, 545-556.

[本文引用: 1]

Wang P (王朋), Wang Y (王莹), Kong CH (孔垂华) (2008)

Allelopathy of plant volatile monoterpenes mediated by soil: a case study of Ambrosia trifida L

Acta Ecologica Sinica (生态学报), 28, 62-68. (in Chinese with English abstract)

[本文引用: 1]

Wang Q, Han C, Ferreira AO, Yu X, Ye W, Tripathy S, Kale SD, Gu B, Sheng Y, Sui Y, Wang X, Zhang Z, Cheng B, Dong S, Shan W, Zheng X, Dou D, Tyler BM, Wang Y (2011)

Transcriptional programming and functional interac- tions within the Phytophthora sojae RXLR effector repert- oire

Plant Cell, 23, 2064-2086.

DOI:10.1105/tpc.111.086082      URL     PMID:21653195     

The genome of the soybean pathogen Phytophthora sojae contains nearly 400 genes encoding candidate effector proteins carrying the host cell entry motif RXLR-dEER. Here, we report a broad survey of the transcription, variation, and functions of a large sample of the P. sojae candidate effectors. Forty-five (12%) effector genes showed high levels of polymorphism among P. sojae isolates and significant evidence for positive selection. Of 169 effectors tested, most could suppress programmed cell death triggered by BAX, effectors, and/or the PAMP INF1, while several triggered cell death themselves. Among the most strongly expressed effectors, one immediate-early class was highly expressed even prior to infection and was further induced 2- to 10-fold following infection. A second early class, including several that triggered cell death, was weakly expressed prior to infection but induced 20- to 120-fold during the first 12 h of infection. The most strongly expressed immediate-early effectors could suppress the cell death triggered by several early effectors, and most early effectors could suppress INF1-triggered cell death, suggesting the two classes of effectors may target different functional branches of the defense response. In support of this hypothesis, misexpression of key immediate-early and early effectors severely reduced the virulence of P. sojae transformants.

Wang Q, Wang CH, Zhao B, Ma ZJ, Luo YQ, Chen JK, Li B (2006)

Effects of growing conditions on the growth of and interactions between salt marsh plants: implications for invasibility of habitats

Biological Invasions, 8, 1547-1560.

[本文引用: 4]

Wang R, Wang Y (2006)

Invasion dynamics and potential spread of the invasive alien plant species Ageratina adenophora (Asteraceae) in China

Diversity and Distrib- utions, 12, 397-408.

Wang W, Gu W, Ding ZF, Ren YL, Chen JX, Hou YY (2005)

A novel Spiroplasma pathogen causing systemic infecting in the crayfish Procambarus clarkii (Crustacea: Decapod), in China

FEMS Microbiology Letters, 249, 131-138.

DOI:10.1016/j.femsle.2005.06.005      URL     PMID:16000238      [本文引用: 1]

A novel disease of crayfish Procambarus clarkii appeared in the summer of 2004 in freshwater aquaculture in Jiangsu province of China. Light and transmission electron microscopy (TEM), molecular biological methods and in vitro culture were used to identify the pathogen. The agent was unique in having a helical morphology and rotary motility as observed by phase-contrast light microscopy and was found in haemolymph, muscles, nerves and connective tissues by smear method and TEM. Ultra-thin sections under TEM revealed the wall-free membrane of the microbe. The agent could pass through membrane filters with pores 220 nm in diameter and was cultivated in vitro in M1D medium. 16S rDNA of the crayfish pathogen was amplified by PCR using primers specific for Spiroplasma-specific 16S rDNA. The resultant 271bp PCR product showed 99% identity with Spiroplasma mirum 16S rDNA, having a close relationship with the spiroplasma from the Chinese mitten crab Eriocheir sinensis. This is the second time a spiroplasma has been found in a freshwater crustacean. The 271bp PCR product was also amplified from the bottom mud in the ponds associated with the disease. The PCR molecular method is an effective way to detect spiroplasma in freshwater environment. The results from this study are significant in expanding the host range of spiroplasma and freshwater ecology.

Wang WQ (王文琪), Wang JJ (王进军), Zhao ZM (赵志模) (2006)

Effects of parasitizing on the sexual reproduction of Eupatorium adenophorum Sprengel by Procecidochares utilis Stone at different microhabitats

Acta Phytophylacica Sinica (植物保护学报), 33, 391-395. (in Chinese with English abstract)

[本文引用: 2]

Wang Y, Dou A, Wang X, Li A, Sheng Y, Hua C, Cheng B, Chen X, Zheng X, Wang Y (2009)

The PsCZF1 gene encoding a C2H2 zinc finger protein is required for growth, development and pathogenesis in Phytophthora sojae

Microbial Pathogenesis, 47, 4778-4786.

[本文引用: 1]

Wang Y, Li A, Wang X, Zhang X, Zhao W, Dou A, Zheng X, Wang Y (2010)

GPR11, a putative seven-transmembrane G protein coupled receptor, controls zoospore development and virulence of Phytophthora sojae

Eukaryotic Cell, 9, 242-250.

DOI:10.1128/EC.00265-09      URL     PMID:20008081      [本文引用: 1]

G protein-coupled receptors (GPCRs) represent a large receptor family involved in a broad spectrum of cell signaling. To understand signaling mechanisms mediated by GPCRs in Phytophthora sojae, we identified and characterized the PsGPR11 gene, which encodes a putative seven-transmembrane GPCR. An expression analysis revealed that PsGPR11 was differentially expressed during asexual development. The highest expression level occurred in zoospores and was upregulated during early infection. PsGPR11-deficienct transformants were obtained by gene silencing strategies. Silenced transformants exhibited no differences in hyphal growth or morphology, sporangium production or size, or mating behavior. However, the release of zoospores from sporangia was severely impaired in the silenced transformants, and about 50% of the sporangia did not completely release their zoospores. Zoospore encystment and germination were also impaired, and zoospores of the transformants lost their pathogenicity to soybean. In addition, no interaction was observed between PsGPR11 and PsGPA1 with a conventional yeast two-hybrid assay, and the transcriptional levels of some genes which were identified as being negatively regulated by PsGPA1 were not clearly altered in PsGPR11-silenced mutants. These results suggest that PsGPR11-mediated signaling controls P. sojae zoospore development and virulence through the pathways independent of G protein.

Wang ZJ (王紫娟), Liu WX (刘万学), Wan FH (万方浩), GZ (吕国忠) (2007)

The allelopathic potential of root exudates from Ageratina adenophora Spreng on different conditions

Chinese Agricultural Science Bulletin (中国农学通报), 23, 351-357. (in Chinese with English abstract)

[本文引用: 1]

Weber E, Li B (2008)

Plant invasions in China: what is to be expected in the wake of economic development?

Bio- Science, 58, 437-444.

[本文引用: 8]

Weber E, Sun SG, Li B (2008)

Invasive alien plants in China: diversity and ecological insights

Biological Invasions, 10, 1411-1429.

Wei SH (魏守辉), Qu Z (曲哲), Zhang CX (张朝贤), Li YJ (李咏军), Li XJ (李香菊) (2006)

Invasive alien species giant ragweed (Ambrosia trifida L.) and its risk assessment

Plant Protection (植物保护), 32, 14-19. (in Chinese with English abstract)

[本文引用: 1]

Wei JR (魏建荣), Yang ZQ (杨忠岐), Wang CZ (王传珍), Qu HR (曲花荣), Shao LS (邵凌松) (2004)

Study on biological control of fall webworm by insect natural enemy

Scientia Silvae Sinicae (林业科学), 40, 90-95. (in Chinese with English abstract)

[本文引用: 1]

Williamson M (1999)

Invasions

Ecography, 22, 5-12.

[本文引用: 1]

Wilsey BJ, Polley HW (2006)

Aboveground productivity and root-shoot allocation differ between native and introduced grass species

Oecologia, 150, 300-309.

DOI:10.1007/s00442-006-0515-z      URL     [本文引用: 1]

Plant species in grasslands are often separated into groups (C4 and C3 grasses, and forbs) with presumed links to ecosystem functioning. Each of these in turn can be separated into native and introduced (i.e., exotic) species. Although numerous studies have compared plant traits between the traditional groups of grasses and forbs, fewer have compared native versus introduced species. Introduced grass species, which were often introduced to prevent erosion or to improve grazing opportunities, have become common or even dominant species in grasslands. By virtue of their abundances, introduced species may alter ecosystems if they differ from natives in growth and allocation patterns. Introduced grasses were probably selected nonrandomly from the source population for forage (aboveground) productivity. Based on this expectation, aboveground production is predicted to be greater and root mass fraction to be smaller in introduced than native species. We compared root and shoot distribution and tissue quality between introduced and native C4 grass species in the Blackland Prairie region of Central Texas, USA, and then compared differences to the more well-studied divergence between C4 grasses and forbs. Comparisons were made in experimental monocultures planted with equal-sized transplants on a common soil type and at the same density. Aboveground productivity and C:N ratios were higher, on average, in native grasses than in native forbs, as expected. Native and introduced grasses had comparable amounts of shallow root biomass and tissue C:N ratios. However, aboveground productivity and total N were lower and deep root biomass and root mass fraction were greater in native than introduced grasses. These differences in average biomass distribution and N could be important to ecosystems in cases where native and introduced grasses have been exchanged. Our results indicate that native–introduced status may be important when interpreting species effects on grassland processes like productivity and plant N accumulation.]]>

Wu BQ (吴碧球), Lu YY (陆永跃), Zeng L (曾玲), Liang GW (梁广文) (2008)

Influences of Solenopsis invicta Buren invasion on the native ant communities in different habitats in Guangdong

Chinese Journal of Applied Ecology (应用生态学报), 19, 151-156. (in Chinese with English abstract)

URL     PMID:18419088      [本文引用: 1]

By using pitfall and bait traps, an investigation was made on the diversity and similarity of ant communities in the areas infested and un-infested with Solenopsis invicta Buren in Shenzhen of Guangdong. The results showed that under the invasion of S. invicta, the ant species number in lawn and wasteland reduced obviously, with a decrease of 6 in lawn and 3 in wasteland, and the native dominant ant species in lichee orchard, especially in wasteland and lawn, were replaced by S. invicta. With the infestation of S. invicta, the diversity and evenness of ant communities in wasteland and lawn decreased but the predominance increased obviously, while it was in adverse in lichee orchard. The similarity coefficients of the ant communities between S. invicta infested and un-infested lichee orchard, wasteland and lawn were 0.6316, 0.5882 and 0.2941, respectively.

Wu R (吴蓉), Chen YW (陈友吾), Chen ZM (陈卓梅), Lin XJ (林晓佳), Liang DD (梁定东) (2005)

Effect of Bursaphelenchus xylophilus disease on vegetation succession in different types of pine forest

Journal of Southwest Forestry College (西南林学院学报), 25, 40-44. (in Chinese with English abstract)

[本文引用: 1]

Wu SH, Hsieh CF, Chaw SM, Rejmánek M (2004)

Plant invasions in Taiwan: insights from the flora of casual and naturalized alien species

Diversity and Distributions, 10, 349-362.

[本文引用: 3]

Wu XL, Zhao JM, Sun S, Yang F, Wang YC, Gai JY, Xing H (2010)

A survey of soybean germplasm for resistance to Phytophthora sojae

Euphytica, 176, 261-268.

[本文引用: 1]

Wu XW (吴晓雯), Luo J (罗晶), Chen JK (陈家宽), Li B (李博) (2006)

Spatial patterns of invasive alien plants in China and its relationship with environmental and anthropological factors

Chinese Journal of Plant Ecology (植物生态学报), 30, 576-584. (in Chinese with English abstract)

[本文引用: 1]

Wu YT, Wang CH, Zhang XD, Zhao B, Jiang LF, Chen JK, Li B (2009)

Effects of saltmarsh invasion by Spartina alterniflora on arthropod community structure and diets

Biological Invasions, 11, 635-649.

[本文引用: 1]

Wu ZJ (武正军), Cai FJ (蔡凤金), Jia YF (贾运锋), Lu JX (鲁建鑫), Jiang YF (蒋勇福), Huang CM (黄乘明) (2008)

Predation impact of Procambarus clarkii on Rana limnocharis tadpoles in Guilin area

Biodiversity Science (生物多样性), 16, 150-155. (in Chinese with English abstract)

[本文引用: 1]

Wurm YJ, Wang O, Riba-Grognuz M, Corona S, Nygaard B, Hunt B, Ingram K, Falquet L, Nipitwattanaphon M, Gotzek D, Dijkstra M, Oettler J, Shih CJ, Wu WJ, Yang CC, Thomas J, Beaudoing E, Pradervand S, Flegel V, Fabbretti R, Stockinger H, Long L, Oakey J, Harkins T, Farmerie W, Boomsma J, Pamilo P, Yi S, Heinze J, Goodisman M, Farinelli L, Harshman K, Hulo N, Cerutti L, Xenarios I, Shoemaker D, Keller L (2011)

Genome of the fire ant Solenopsis invicta

Proceedings of the National Academy of Sciences, USA, 108, 5679-5684.

[本文引用: 2]

Xiang JC (相君成), Lei ZR (雷仲仁), Wang HH (王海鸿), Gao YL (高玉林) (2012)

Interspecific competition among three invasive Liriomyza species

Acta Ecologica Sinica (生态学报), 32, 1616-1622. (in Chinese with English abstract)

[本文引用: 2]

Xiao DR (肖德荣), Zhang LQ (张利权), Zhu ZC (祝振昌), Tian K (田昆) (2011)

Response of seed production and viability of Spartina alterniflora to cutting at Shanghai Chongming Dongtan, China

Ecology and Environmental Sciences (生态环境学报), 20, 1681-1686. (in Chinese with English abstract)

[本文引用: 1]

Xie BY, Chen XY, Shi J, Zhang QW, Dai SM, Cheng FX, Luo YQ (2009)

Mechanisms of invasive population establish- ment and spread of pinewood nematodes in China

Science in China: Series C, Life Sciences, 52, 587-594.

DOI:10.1007/s11427-009-0071-y      URL     PMID:19557337      [本文引用: 2]

This paper summarizes the results of our study of the pinewood nematode (Bursaphelenchus xylophilus). By population genetic analysis, it was determined that there was no genetic bottle caused by the founder effect and genetic drift in the Chinese invasive population. Multiple invasions with large amounts of nematodes from different sources led to rich genetic diversity in the invasive population. Keeping high genetic diversity in the invasive process may be one of the genetic mechanisms in its successful invasion. By testing interspecies competition, it was shown that, with high fecundity and a strong competitive ability, B. xylophilus outcompeted the native species B. mucronatus in the natural ecosystem during the invasion process. Competitive displacement may be one of the ecological mechanisms of B. xylophilus's invasion. In addition, an unequal interspecific hybridization with introgress was in favor of the invasive species which also accelerated the replacement of B. mucronatus by B. xylophilus. The structures, functions and evolutions of a few important genes that are closely related to the ecological adaptation of pinewood nematodes were studied to explore the molecular mechanism of its ecological adaptations. Further, the resistance and resilience mechanism of the pine ecosystem invaded by pinewood nematodes was also investigated. The results of these studies uncovered a portion of the genetic and ecological mechanisms of PWN's successful invasion and laid a foundation for further study to obtain a comprehensive interpretation of the mechanisms of the nematode invasion. The results also provided a scientific basis for effectively controlling the occurrence and spread of pine wilt disease which is caused by nematodes. Various aspects requiring further investigation are considered.

Xu CW, Yang MZ, Chen YJ, Chen LM, Zhang DZ, Mei L, Shi YT, Zhang HB (2012)

Changes in non-symbiotic nitrogen-fixing bacteria inhabiting rhizosphere soils of an invasive plant Ageratina adenophora

Applied Soil Ecology, 54, 32-38.

[本文引用: 1]

Xu HG (徐海根), Qiang S (强胜), Han ZM (韩正敏), Guo JY (郭建英), Huang ZG (黄宗国), Sun HY (孙红英), He SP (何舜平), Ding H (丁晖), Wu HR (吴海荣), Wan FH (万方浩) (2004)

The distribution and introduction pathway of alien invasive species in China

Biodiversity Science (生物多样性), 12, 626-638. (in Chinese with English abstract)

[本文引用: 1]

Xu HG (徐海根), Qiang S (强胜) (2011) China’s Invasive Alien Species (中国外来入侵生物). Science Press, Beijing. (in Chinese)

[本文引用: 3]

Xu HG, Ding H, Li MY, Qiang S, Guo JY, Han ZM, Huang ZG, Sun HY, He SP, Wu HR, Wan FH (2006)

The distribution and economic losses of alien species invasion in China

Biological Invasions, 8, 1495-1500.

[本文引用: 1]

Xu J (徐婧), Jiang HX (姜红霞), A LY (阿丽亚), Guo JM (郭静敏), Zhang RZ (张润志) (2012a)

Growth and patterns of population decline in Cydia pomonella adults in Gansu, Xinjiang and Inner Mongolia

Chinese Journal of Applied Entomology (应用昆虫学报), 49, 89-95. (in Chinese with English abstract)

[本文引用: 2]

Xu J (徐婧), Du L (杜磊), Qin TJ (秦天军), Zhang JC (张建超), Zhang RZ (张润志) (2012b)

Dynamic distribution of Cydia pomonella adults in apple orchards

Chinese Journal of Applied Entomology (应用昆虫学报), 49, 80-88. (in Chinese with English abstract)

[本文引用: 1]

Yang B (杨博), Yangjin Z (央金卓嘎), Pan XY (潘晓云), Xu HG (徐海根), Li B (李博) (2010)

Alien terrestrial herbs in China: diversity and ecological insights

Biodiversity Science (生物多样性), 18, 660-666. (in Chinese with English abstract)

[本文引用: 1]

Yang CC, Ascunce MS, Luo LZ, Shao JG, Shih CJ, Shoemaker D (2012)

Propagule pressure and colony social organization are associated with the successful invasion and rapid range expansion of fire ants in China

Molecular Ecology, 21, 817-833.

DOI:10.1111/j.1365-294X.2011.05393.x      URL     PMID:22181975      [本文引用: 1]

We characterized patterns of genetic variation in populations of the fire ant Solenopsis invicta in China using mitochondrial DNA sequences and nuclear microsatellite loci to test predictions as to how propagule pressure and subsequent dispersal following establishment jointly shape the invasion success of this ant in this recently invaded area. Fire ants in Wuchuan (Guangdong Province) are genetically differentiated from those found in other large infested areas of China. The immediate source of ants in Wuchuan appears to be somewhere near Texas, which ranks first among the southern USA infested states in the exportation of goods to China. Most colonies from spatially distant, outlying areas in China are genetically similar to one another and appear to share a common source (Wuchuan, Guangdong Province), suggesting that long-distance jump dispersal has been a prevalent means of recent spread of fire ants in China. Furthermore, most colonies at outlier sites are of the polygyne social form (featuring multiple egg-laying queens per nest), reinforcing the important role of this social form in the successful invasion of new areas and subsequent range expansion following invasion. Several analyses consistently revealed characteristic signatures of genetic bottlenecks for S. invicta populations in China. The results of this study highlight the invasive potential of this pest ant, suggest that the magnitude of international trade may serve as a predictor of propagule pressure and indicate that rates and patterns of subsequent range expansion are partly determined by the interplay between species traits and the trade and transportation networks.

Yang FH (杨凤辉), Ma T (马涛), Chen JK (陈家宽), Li B (李博) (2002)

Eichhornia crassipes disaster in Huangpu River in Shanghai: cause, consequences and control strategies

Journal of Fudan University (复旦学报), 41, 599-603. (in Chinese with English abstract)

[本文引用: 1]

Yang GQ, Wan FH, Liu WX, Zhang XW (2006)

Physiological effects of allelochemicals from leachates of Ageratina adenophora (Spreng.) on rice seedlings

Allelopathy Journal, 18, 237-246.

[本文引用: 1]

Yang GQ, Wan FH, Liu WX, Guo JY (2008)

Influence of two allelochemicals from Ageratina adenophora Sprengel on ABA, IAA and ZR contents in roots of upland rice seedlings

Allelopathy Journal, 21, 253-262.

[本文引用: 1]

Yang GQ, Wan FH, Guo JY, Liu WX (2011)

Cellular and ultrastructural changes in the seedling roots of upland rice (Oryza sativa) under the stress of two allelochemicals from Ageratina adenophora

Weed Biology and Management, 11, 152-159.

[本文引用: 1]

Yang JQ (杨建强), Zhao X (赵骁), Yan YG (严勇敢), Zhang YL (张雅林), Feng JN (冯纪年) (2011)

Efficacy of seven kinds of pesticides for controlling codling moth Cydia pomonella

Acta Agriculturae Boreali-occidentalis Sinica (西北农业学报), 20, 194-196. (in Chinese with English abstract)

[本文引用: 1]

Yang ZQ (杨忠岐), Zhang YA (张永安) (2007)

Researches on techniques for biocontrol of the fall webworm, Hyphantria cunea, a severe invasive insect pest to China

Chinese Bulletin of Entomology (昆虫知识), 44, 465-471. (in Chinese with English abstract)

[本文引用: 2]

Yuan L, Wang S, Zhou J, Du Y, Zhang Y, Wang J (2012)

Status of insecticide resistance and associated mutations in Q-biotype of whitefly, Bemisia tabaci, from eastern China

Crop Protection, 31, 67-71.

[本文引用: 1]

Yu DZ (喻大昭), Wei SH (魏守辉), Zhu WD (朱文达), Cao AC (曹坳程), Zhang CX (张朝贤), Song ZX (宋兆辛) (2008)

Influence of Alternanthera philoxeroides on the growth of paddy rice and its economic threshold

Acta Phytophylacica Sinica (植物保护学报), 35, 69-73. (in Chinese with English abstract)

[本文引用: 1]

Yu LQ (余柳青), Fujii Y, Zhou YJ (周勇军), Zhang JP (张建萍), Lu YL (陆永良), Xuan SN (玄松南) (2007a)

Comparison of allelopathy potential between an exotic invasive weed Alternanthera philoxeroides and a local weed A. sessilis

Chinese Journal of Rice Science (中国水稻科学), 21, 84-89. (in Chinese with English abstract)

[本文引用: 1]

Yu LQ (余柳青), Fujii Y, Zhou YJ (周勇军), Zhang JP (张建萍), Lu YL (陆永良), Xuan SN (玄松南) (2007b)

Response of aquatic Alternanthera philoxeroides to environmental factors and its competition with rice

Chinese Journal of Rice Science (中国水稻科学), 21, 209-214. (in Chinese with English abstract)

[本文引用: 1]

Yu H, Wan FH, Guo JY (2012a)

cDNA cloning of heat shock protein genes and their expression in an indigenous cryptic species of the whitefly Bemisia tabaci complex from China

Journal of Integrative Agriculture, 11, 293-302.

[本文引用: 1]

Yu H, Wan FH, Guo JY (2012b)

Different thermal tolerance and hsp gene expression in invasive and indigenous sibling species of Bemisia tabaci

Biological Invasions, 14, 1587-1595.

[本文引用: 1]

Yu X, Yu D, Lu Z, Ma K (2005)

A new mechanism of invader sucesss: extotic plant inhibits natural vegetation restoration by change soil microbe community

Chinese Science Bulletin, 50, 1105-1112.

[本文引用: 1]

Zeng L (曾玲), Lu YY (陆永跃), He XF (何晓芳), Zhang WQ (张维球), Liang GW (梁广文) (2005a)

Identification of red imported fire ant Solenopsis invicta to invade mainland China and infestation in Wuchuan, Guangdong

Chinese Bulletin of Entomology (昆虫知识), 42, 144-148. (in Chinese with English abstract)

[本文引用: 1]

Zeng L (曾玲), Lu YY (陆永跃), Chen ZN (陈忠男) (2005b) Monitoring and Control of Solenopsis invicta (红火蚁监测与防治). Guangdong Science and Technology Press, Guangzhou. (in Chinese)

[本文引用: 2]

Zhang FJ (张风娟), Guo JY (郭建英), Long R (龙茹), Wan FH (万方浩) (2010)

Allelopathy of different treated residues of Ambrosia artemisiifolia L. on wheat growth

Chinese Journal of Ecology (生态学杂志), 29, 669-673. (in Chinese with English abstract)

[本文引用: 1]

Zhang GL (张国良), Fu WD (付卫东), Liu K (刘坤) (2008) Alien Invasive Species in Agriculture (农业重大外来入侵生物). Science Press, Beijing. (in Chinese)

[本文引用: 1]

Zhang K (张锴), Liang J (梁军), Yan DH (严冬辉), Zhang XY (张星耀) (2010)

Research advances of pine wood nematode disease in China

World Forestry Research (世界林业研究), 23, 59-63. (in Chinese with English abstract)

[本文引用: 1]

Zhang R, Li Y, Liu N, Porter SD (2007)

An overview of the red imported fire ant (Hymenoptera: Formicidae) in mainland China

Florida Entomologist, 90, 723-731.

[本文引用: 1]

Zhang RZ (张润志), Liu N (刘宁), Li YC (李颖超) (2010)

Colorado potato beetle, an disastrous invasive species in China infesting Solanaceae vegetables

Chinese Bulletin of Life Science (生命科学), 22, 1118-1121. (in Chinese with English abstract)

[本文引用: 1]

Zhang RZ (张润志), Wang FX (王福祥), Zhang YL (张雅林), Chen HJ (陈汉杰), Luo JC (罗进仓), Wang QY (王勤英), Liu WX (刘万学), Ainiwaer·M (艾尼瓦尔·木沙), Pu CJ (蒲崇建), Yan YG (严勇敢), Guo JM (郭静敏), Liu XY (刘星月), Chen JG (陈继光), Zhang ZF (张增幅), Yang S (杨森), Xu JJ (许建军), Cui GZ (崔艮中), Xu J (徐婧) (2012)

Progress on monitoring and control of the codling moth, Cydia pomonella (L.)

Chinese Journal of Applied Entomology (应用昆虫学报), 49, 37-42. (in Chinese with English abstract)

[本文引用: 3]

Zhang SS, Jin YL, Tang JJ, Chen X (2009)

The invasive plant Solidago canadensis L, suppresses local soil pathogens through allelopathy

Applied Soil Ecology, 41, 215-222.

[本文引用: 1]

Zhang XD (张骁栋) (2012) The Effects of Smooth Cordgrass and Crab Bioturbation on Plant Interspecific Relationship and Biogeochemistry in a Salt Marsh in the Yangtze Estuary (互花米草与蟹类扰动对崇明东滩植物种间关系及生地化循环的影响). PhD dissertation, Fudan University, Shang- hai. (in Chinese)

[本文引用: 1]

Zhang XW (张学文), Liu WX (刘万学), Wan FH (万方浩), Liu YX (刘亦学), Yang GQ (杨国庆), Zhang W (张惟), Yang XR (杨秀荣) (2007b)

Allelopathy of aqueous leachates of Ageratina adenophora and the indentification of the allelichemicals

Journal of Agricultural University of Hebei (河北农业大学学报), 30, 68-71, 76. (in Chinese with English abstract)

[本文引用: 1]

Zhang XW (张学文), Liu YX (刘亦学), Liu WX (刘万学), Wan FH (万方浩), Zhang W (张惟), Yang XR (杨秀荣) (2007a)

Allelochemicals and its releasing modes

Chinese Agricultural Science Bulletin (中国农学通报), 23, 295-297. (in Chinese with English abstract)

[本文引用: 1]

Zhang YH (张玉虎), Liu MF (刘梅芳), Ling TJ (凌铁军), Wei XY (魏孝义) (2004)

Allelopathic sesquiterpene lactones from Wedelia trilobata

Journal of Tropical and Subtropical Botany (热带亚热带植物学报), 12, 533-537. (in Chinese with English abstract)

[本文引用: 1]

Zhang YJ (张友军), Zhu GR (朱国仁), Chu D (褚栋), Wu QJ (吴青君), Wang SL (王少丽) (2011)

Occurrence, damage and control of important invasive insect pests on the vegetable crops in China

Plant Protection (植物保护), 37, 1-6. (in Chinese with English abstract)

[本文引用: 1]

Zhao BG, Wang HL, Han SF, Han ZM (2003)

Distribution and pathogenicity of bacteria species carried by Bursaphel- enchus xylophilus in China

Nematology, 5, 899-906.

[本文引用: 1]

Zhao GQ (赵广琦), Zhang LQ (张利权), Liang X (梁霞) (2005)

A comparison Spartina alterniflora and an indigen- ous plant Phragmites australis

Acta Ecologica Sinica (生态学报), 25, 1604-1611. (in Chinese with English abstract)

[本文引用: 1]

Zhao L (赵林), Li BP (李保平), Meng L (孟玲) (2008)

Effects of fertilization and grass-cutting on competition between Lolium perenne and the invasive exotic Eupatorium adenophorum at seedling stage

Acta Prataculturae Sinica (草业学报), 17, 151-155. (in Chinese with English abstract)

[本文引用: 1]

Zhao YJ, Qing H, Zhao CJ, Zhou CF, Zhang WG, Xiao Y, An SQ (2010)

Phenotypic plasticity of Spartina alterniflora and Phragmites australis in response to nitrogen addition and intraspecific competition

Hydrobiologia, 637, 143-155.

[本文引用: 3]

Zhao YX, Kang L (2000)

Cold tolerance of the leaf miner Liriomyza sativae (Dipt., Agromyzidae)

Journal of Applied Entomology, 124, 185-189.

[本文引用: 1]

Zheng CH (郑春寒), Liu Q (刘强), Li J (李坚), Dong K (董昆), Feng JN (冯纪年), Zhang YL (张雅林), Wang D (王敦) (2011)

Virulence determination for Cydia pomonella Granulovirus Zhangye Strain (CypoGV-zy)

Journal of Northwest Forestry University (西北林学院学报), 26, 121-123. (in Chinese with English abstract)

[本文引用: 1]

Zhou CF, An SQ, Deng ZF, Yin DQ, Zhi YB, Sun ZY, Zhao H, Zhou LX, Fang C, Qian C (2009)

Sulfur storage changed by exotic Spartina alterniflora in coastal saltmarshes of China

Ecological Engineering, 35, 536-543.

[本文引用: 1]

Zhou ZS, Guo JY, Chen HS, Wan FH (2010a)

Effects of temperature on survival, development, longevity, and fecundity of Ophraella communa (Coleoptera: Chrysom- elidae), a potential biological control agent against Ambrosia artemisiifolia (Asterales: Asteraceae)

Environmental Ento- mology, 39, 1021-1027.

[本文引用: 1]

Zhou ZS, Guo JY, Chen HS, Wan FH (2010b)

Effect of humidity on the development and fecundity of Ophraella communa (Coleoptera: Chrysomelidae)

BioControl, 55, 313-319.

[本文引用: 1]

Zhou ZS, Guo JY, Zheng XW, Luo M, Chen HS, Wan FH (2011)

Reevaluation of biosecurity of Ophraella communa against sunflower (Helianthus annuus)

Biocontrol Science and Technology, 21, 1147-1160.

[本文引用: 1]

Zhou ZS (周忠实), Guo JY (郭建英), Li BP (李保平), Meng L (孟玲), Fu JW (傅建炜), Chen HS (陈红松), Ma MY (马明勇), Shi MZ (史梦竹), Li M (李敏), Guo W (郭薇), Luo M (罗敏), Zheng XW (郑兴汶), Zheng HY (郑海燕), Luo YH (罗源华), Wan FH (万方浩) (2011a)

Distribution of Ambrosia artemisiifolia and Alternanthera philoxcroides and their regional sustainable management strategies

Journal of Biosafety (生物安全学报), 20, 263-266. (in Chinese)

[本文引用: 1]

Zhou ZS (周忠实), Chen HS (陈红松), Zheng XW (郑兴汶), Guo JY (郭建英), Wan FH (万方浩) (2011b)

Combined control of common ragweed, Ambrosia artemisiifolia with Ophraella communa and Epiblema strenuana in Laibin, Guangxi Province, China

Journal of Biosafety (生物安全学报), 20, 267-269. (in Chinese with English abstract)

[本文引用: 1]

Zhou ZS (周忠实), Chen HS (陈红松), Guo JY (郭建英), Guo W (郭薇), Luo YH (罗源华), Zheng XW (郑兴汶), Luo M (罗敏), Zheng HY (郑海燕), Wan FH (万方浩) (2011c)

A field trial of biological control of Ambrosia artemisiifolia with two herbivorous insects in Miluo, Hunan Province, China

Journal of Biosafety (生物安全学报), 20, 186-191. (in Chinese with English abstract)

[本文引用: 1]

Zhou ZS, Guo JY, Guo W, Zheng HY, Wan FH (2012a)

Effects of morphological traits, age and copulation experience on sexual selection in Ophraella communa

Biocontrol Science and Technology, 22, 81-91.

Zhou ZS, Guo JY, Michaud JP, Li M, Wan FH (2012b)

Variation in cold hardiness among geographic populations of the ragweed beetle, Ophraella communa LeSage (Coleoptera: Chrysomelidae), a biological control agent of Ambrosia artemisiifolia L. (Asterales: Asteraceae), in China

Biological Invasions, 13, 659-667.

Zhu DH, Zhu J, Peng ZP, Wan FH (2012)

Effects of photoperiod and temperature on reproductive diapause in Ophraella communa (Coleoptera: Chrysomelidae), a potential biocontrol agent against Ambrosia artemisiifolia

Insect Science, 19, 286-294.

[本文引用: 1]

Zhu HW (朱宏伟), Meng L (孟玲), Li BP (李保平) (2007)

Relative competitive ability of loum perenne and the alien invasive weed, Eupatorium adenophorum (Compositae) at seedling stage

Chinese Journal of Applied and Environ- mental Biology (应用与环境生物学报), 13, 29-32. (in Chinese with English abstract)

[本文引用: 2]

Zhu P, Tang XQ, Xu HG (2005)

Model study on loss assessment of invasion species

World Journal Modelling Simulation, 1, 19-26.

[本文引用: 1]

Zhu HY (朱虹昱), Liu W (刘伟), Cui GZ (崔艮中), Zhang ZF (张增福), Zhang RZ (张润志) (2012a)

Field trials using domestic sex pheromone dispensers to disrupt Cydia pomonella mating behavior

Chinese Journal of Applied Entomology (应用昆虫学报), 49, 121-129. (in Chinese with English abstract)

[本文引用: 1]

Zhu HY (朱虹昱), Du L (杜磊), Xu J (徐婧), Liu W (刘伟), Zhang RZ (张润志) (2012b)

Effective duration of sex pheromone in Delta traps for monitoring the codling moth

Chinese Journal of Applied Entomology (应用昆虫学报), 49, 114-120. (in Chinese with English abstract)

[本文引用: 1]

Zhu L, Sun O, Sang W, Li Z, Ma K (2007)

Predicting the spatial distribution of an invasive plant species ( Eupatorium adenophorum) in China

Landscape Ecology, 22, 1143-1154.

Zhu X, Zhang J, Ma K (2011)

Soil biota reduce allelopathic effects of the invasive Eupatorium adenophorum

PLoS ONE, 6, e25393.

DOI:10.1371/journal.pone.0025393      URL     PMID:21980442      [本文引用: 1]

Allelopathy has been hypothesized to play a role in exotic plant invasions, and study of this process can improve our understanding of how direct and indirect plant interactions influence plant community organization and ecosystem functioning. However, allelopathic effects can be highly conditional. For example allelopathic effects demonstrated in vivo can be difficult to demonstrate in field soils. Here we tested phytotoxicity of Eupatorium adenophorum (croftonweed), one of the most destructive exotic species in China, to a native plant species Brassica rapa both in sand and in native soil. Our results suggested that natural soils from different invaded habitats alleviated or eliminated the efficacy of potential allelochemicals relative to sand cultures. When that soil is sterilized, the allelopathic effects returned; suggesting that soil biota were responsible for the reduced phytotoxicity in natural soils. Neither of the two allelopathic compounds (9-Oxo-10,11-dehydroageraphorone and 9b-Hydroxyageraphorone) of E. adenophorum could be found in natural soils infested by the invader, and when those compounds were added to the soils as leachates, they showed substantial degradation after 24 hours in natural soils but not in sand. Our findings emphasize that soil biota can reduce the allelopathic effects of invaders on other plants, and therefore can reduce community invasibility. These results also suggest that soil biota may have stronger or weaker effects on allelopathic interactions depending on how allelochemicals are delivered.

Zhuang YQ (庄义庆) (2009) Exploratory Development of Pathogenic Fungi-Fusarium Stoveri Against Alligator Weed (Alternanthera philoxeroides) (蕉斑镰刀菌在水花生生物防治上的应用研究). PhD dissertation, Nanjing Agricultural University, Nanjing. (in Chinese)

[本文引用: 1]

Zuo P (左平), Liu CA (刘长安), Zhao SH (赵书河), Wang CH (王春红), Liang YB (梁玉波) (2009)

Distribution of Spartina plantations along the China’s coast

Acta Oceanologica Sinica (海洋学报), 31, 101-111. (in Chinese with English abstract)

URL     [本文引用: 1]

S.anglica,S.alterniflora,S.patens and S.cynosuriodes,have been introduced to China since 1963.S.anglica and S.alterniflora have been widely planted along the China's coast and arousing hot discussions and arguments about their effects and characteristics of invasion since 2000.The survey shows that S.anglica has been greatly degraded and only 16 hm2 are left along the China's coast.However,S.alterniflora has widely spread to Liaoning Province,Hebei Province,Tianjin Municipality,Shandong Province,Jiangsu Province,Shanghai Municipality,Zhejiang Province,Fujian Province,Guangdong Province,the Zhuangzu.Autonomous Region of Guangxi and HongKong with an area of 34 451 hm2 till 2007.The distribution of S.alterniflora can be divided into three parts according to its ecological niche and the characteristics of the China's coast.One is the tidal flats to the south part of Fuding,Fujian Province,which shows potential competition with mangrove ecosystem.The other is estuaries,especially in the Changjiang Estuary,which shows obvious competition with native reeds and Scirpusmari queter.The third is coastal tidal flats in the China's coast,which shows ambiguous effects among its economic,environmental,ecological benefits and invasive threats against local ecosyst em.S.alterniflora has shown different colonization capacity and spreading capacity in different areas in China's coast according to its distribution.Therefore,further studies and assessments on the ecological effects caused by S.alterniflora in the China's coast should be scientifically evaluated.]]>

/