生物多样性, 2019, 27(3): 306-313 doi: 10.17520/biods.2018269

研究报告

利用昆虫携带的花粉初探西藏入侵植物印加孔雀草在当地传粉网络中的地位

土艳丽1, 王力平2,3, 王喜龙1, 王林林,2,3, 段元文2

1 西藏自治区高原生物研究所, 拉萨 850001

2 中国科学院昆明植物研究所, 昆明 650201

3 中国科学院大学, 北京 100049

Status of invasive plants on local pollination networks: A case study of Tagetes minuta in Tibet based on pollen grains from pollinators

Yanli Tu1, Liping Wang2,3, Xilong Wang1, Linlin Wang,2,3, Yuanwen Duan2

1 Tibet Plateau Institute of Biology, Lhasa 850001

2 Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201

3 University of Chinese Academy of Sciences, Beijing 100049

通讯作者: * E-mail: wanglinlin0328@mail.kib.ac.cn

编委: 黄双全

责任编辑: 时意专

收稿日期: 2018-10-11   接受日期: 2019-03-2   网络出版日期: 2019-03-20

基金资助: 西藏自治区科技计划项目(ZD20170021)

Received: 2018-10-11   Accepted: 2019-03-2   Online: 2019-03-20

摘要

入侵植物在新生境中成功定殖后, 通过利用当地传粉昆虫促进繁殖可以更好地保证种群的扩张, 但是入侵植物在当地传粉网络中的角色和地位仍不是很清楚。本文利用西藏近年发现的入侵植物印加孔雀草(Tagetes minuta), 分析其访花昆虫所携带的植物花粉种类, 构建了植物花粉-传粉者网络, 探讨印加孔雀草快速入侵和扩张的可能机制。结果表明印加孔雀草为泛化传粉系统, 共有13种昆虫访花, 其中12种携带有印加孔雀草花粉, 所有花粉中印加孔雀草花粉数量占比为89.89%。12种印加孔雀草传粉昆虫中, 4种泛化传粉昆虫(1种蜂、2种食蚜蝇和1种蝇)是其主要传粉昆虫。本研究揭示印加孔雀草在较短时间内已经成功利用多种当地泛化传粉昆虫为其授粉, 已顺利融入当地的传粉网络, 今后需要更加重视对印加孔雀草的防控。

关键词: 青藏高原 ; Tagetes minuta ; 入侵植物 ; 传粉网络 ; 花粉

Abstract

Invasive plants that use local pollinators would better ensure their expansion in the new environment, but the role and status of the invasive plants on local pollination network is still unclear. In this paper, we analyzed the pollen grains of Tagetes minuta carried by pollinators and constructed a plant-pollinator network, to explore the roles and impacts of T. minuta in local pollination network. The results showed that pollination system of T. minuta was generalized, and there were 13 insect species visiting its flowers, 12 of which carried pollen grains of T. minuta, and the pollen grains of T. minuta accounted for 89.89%. Among the 12 pollinator species of T. minuta, one species of bees, two species of scorpion flies and one species of flies are the main pollinators. This study reveals that T. minuta has successfully used a variety of local pollinators for pollination in a relatively short period of time, indicating that T. minuta has strong adaptability and diffusion ability, and it is necessary to pay more attentions to the prevention and control of T. minuta in the future.

Keywords: Qinghai-Tibet Plateau ; Tagetes minuta ; invasive plants ; pollination network ; pollen

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引用本文

土艳丽, 王力平, 王喜龙, 王林林, 段元文. 利用昆虫携带的花粉初探西藏入侵植物印加孔雀草在当地传粉网络中的地位. 生物多样性, 2019, 27(3): 306-313 doi:10.17520/biods.2018269

Yanli Tu, Liping Wang, Xilong Wang, Linlin Wang, Yuanwen Duan. Status of invasive plants on local pollination networks: A case study of Tagetes minuta in Tibet based on pollen grains from pollinators. Biodiversity Science, 2019, 27(3): 306-313 doi:10.17520/biods.2018269

生物入侵是指外来生物在扩散到新的栖息地后种群发生暴发性扩张, 最终引起严重的经济损失并可能危害生态系统。作为21世纪五大全球性问题之一(Millennium Ecosystem Assessment, 2005), 生物入侵严重威胁着全球生态系统的功能和服务(Weber & Li, 2008; Jeschke et al, 2015; Kaiser-Bunbury et al, 2017; 孙士国等, 2018)。入侵植物大量侵占当地生境, 能迅速改变农业生态系统并带来严重的经济损失(Kearns et al, 1998)。此外, 入侵植物能打破入侵地生态系统平衡, 改变动植物多样性及其相互关系(Lopezaraiza-Mikel et al, 2007)。当入侵植物能够在新的栖息地建立种群并利用当地的传粉昆虫开始繁殖后, 当地植物与传粉昆虫的传粉网络被打破, 进而影响当地的生态功能(Eviner et al, 2012; Potts et al, 2016)。例如, Kaiser-Bunbury等(2017)通过对塞舌尔马赫岛上有无入侵植物的植物群落的植物-传粉者网络比较发现, 存在入侵物种的群落具有更低的传粉者物种数量、访问次数和互作多样性, 植物与传粉者互作关系也更为特化, 并且入侵植物的存在降低了当地部分野生植物的繁殖成功。

我国是生物多样性大国, 存在复杂而多样的地理环境、气候和生态系统类型, 使外来物种极易定殖并扩散, 从而成为入侵物种(Xie et al, 2001)。据统计, 我国当前入侵生物数量已达529种(鞠瑞亭等, 2012), 其中入侵植物种类占所有入侵生物比例高达51.04% (Weber et al, 2008)。由于多数菊科植物具有生长发育快、生态幅广、瘦果产量高且易于传播, 并能产生化感物质抑制周边植物生长等特点, 成为我国主要的入侵植物类群(朱世新等, 2005)。已有研究表明, 对依赖传粉者才能繁殖成功的入侵植物来说, 成功利用当地传粉者为其传粉是决定入侵植物能否入侵和扩散的关键(McKinney & Goodell, 2011; Powell et al, 2011)。而菊科植物特有的头状花序能吸引多达25种昆虫为其传粉(Schemske, 1983), 如此泛化的传粉系统为其在入侵地繁殖成功及快速扩张提供了条件, 并可能对群落的稳定性和功能产生重大影响(Bascompte et al, 2006)。但是不同的传粉者(比如膜翅目、双翅目和鳞翅目昆虫)在群落中所处的地位不同, 不同传粉功能群对泛化的入侵植物的访花频率和传粉效率可能存在差异, 从而在传粉网络上表现出不同的连接强度(Olesen, 2007; Wang et al, 2017)。但是到目前为止, 利用花粉网络来分析入侵植物与本土传粉者关系的研究相对较少。

本研究以西藏新近发现的入侵植物印加孔雀草(Tagetes minuta)为对象, 通过分析其访花昆虫所携带的花粉网络特征, 试图解答以下科学问题: (1)印加孔雀草的主要传粉者是什么?与本土哪些传粉者构建了连接?(2)印加孔雀草在本土花粉网络上的角色如何, 是否有可能成为群落的核心物种?本研究在群落水平上通过量化入侵植物印加孔雀草的访花昆虫所携带的花粉, 为探究入侵植物如何快速融入当地传粉网络提供证据。

1 材料与方法

1.1 研究材料

印加孔雀草为菊科万寿菊属(Tagetes)一年生草本植物, 株高0.1-2.5 m, 具有万寿菊属特有的芳香气味。印加孔雀草原产于南美洲南部温带草原和山区, 近年来已在中国台湾、江苏、山东、北京、西藏等地成功定殖(董振国等, 2013; 张劲林等, 2014; 许敏和扎西次仁, 2015)。近年来在西藏拉萨市周边以及山南、林芝等地区发现大量印加孔雀草种群。印加孔雀草在西藏的开花时间为9月初至11月初, 花期长达2个月。盛花期为9月中旬至10月中旬, 这期间当地开花植物主要有龙胆科、唇形科、石竹科和马先蒿属(Pedicularis)植物。本实验选择在林芝市朗县附近的一个印加孔雀草群落(92°31° E, 29°8° N, 海拔3,294 m)进行。该群落位于当地栽种苹果树的农田中, 印加孔雀草个体数量较多, 其中正在开花的植株大于1,000株, 大部分植株高0.5-1.5 m, 部分植株高达2.5 m。在印加孔雀草样方内, 本土植物几乎均被排挤, 而在样方附近主要有龙胆科、唇形科以及马先蒿属植物。

1.2 印加孔雀草访花昆虫收集

为了确定印加孔雀草的主要访花昆虫, 我们在群落中选取一个50 m× 60 m的样地, 样地内正在开花的植物主要为印加孔雀草。采用截面式观察方法, 沿着样方四周缓慢步行, 观察并捕捉沿途2 m范围内印加孔雀草的访花昆虫。在印加孔雀草的盛花期进行观察, 具体时间为2017年9月24-25日, 观察时间为9:00-19:00, 每次步行时间为60 min, 共计观察10 h。早于9:00和晚于19:00, 由于高原温度较低限制昆虫活动, 少有访花昆虫。当昆虫接触花药或者柱头时才被认定为传粉昆虫并进行捕捉。双翅目和膜翅目昆虫分别放入装有95%酒精的离心管中, 如果采集的传粉昆虫具有携粉足, 则把携粉足取下后保存。鳞翅目昆虫放入装有硅胶的三角包中带回实验室放入冰箱低温保存。观察期间捕捉到13种传粉昆虫计55只, 同时采集正在散粉的花并将花药固定在95%酒精中作为对照。

1.3 花粉分析

除鳞翅目昆虫直接取下头部喷金进行花粉分析外, 其他保存在95%酒精内的昆虫用超声波振荡器把虫体上的花粉进行洗脱, 然后取出保存在装有95%酒精的离心管中。花粉悬浊液在5,000转速的离心机上离心10 min, 移除上清液后把花粉样品转移到电镜样品台上, 喷金后在扫描电镜下观察并拍照。由于蜂类的携粉足上的花粉通常不具传粉作用, 我们只统计了除携粉足外其他部位的花粉数量和形态。为了分析印加孔雀草的传粉者所携带花粉的植物种类, 我们首先通过扫描印加孔雀草的花粉确定其形态特征, 然后再统计每一种昆虫虫体上的花粉类别。主要通过花粉的萌发孔数量、花粉大小和表面纹饰来区别植物种类。由于收集的蜂类样品较多且携带花粉数量较大, 只选择其中的20只进行电镜扫描分析, 最终用于分析的昆虫个体共43只。

1.4 统计分析

首先利用花粉类型和昆虫种类构建数据矩阵, 其中植物花粉类型作为行数据, 传粉昆虫类型作为列数据, 第i行与第j列对应的单元格则为第j种传粉昆虫所携带的第i种植物的平均花粉数量; 如果所有昆虫均不携带这种花粉, 则记为0。利用R软件中的bipartite包来计算花粉网络在网络和物种水平上的相关参数。

在网络水平上主要关注植物和传粉昆虫种类以及两者之间的连接数量和连接度(connectance), 以及网络的嵌套结构(nestedness)和特化水平(H°2)。网络的连接度和嵌套结构可以指示网络的稳定性, 一般来说网络的连接度和网络的嵌套结构越高, 网络的稳定性就越强(Bascompte et al, 2003)。嵌套结构用网络的嵌套温度(nestedness temperature)表示, 嵌套温度取值范围为0-100, 0表示网络具有完全嵌套的结构, 100表示完全不存在嵌套现象。另外我们还计算了网络的加权嵌套度(weighted nestedness), 取值范围为0-1, 0表示网络不存在嵌套结构, 1表示网络具有完美的嵌套结构。网络水平特化指数(H°2)描述的是网络物种间生态位互补程度(niche complementarity), 取值范围为0-1, 0表示最泛化, 1表示最特化。

在物种水平上我们主要关注传粉昆虫以及植物的连接数量和比例, 以及物种强度(species strength)和特化水平()。物种强度是单一物种所有连接的强度之和, 又可以分为动物对植物的依赖性和植物对动物的依赖性两种(Bascompte et al, 2006)。物种特化水平指数()则描述网络内的某个物种与其他物种生态位的互补程度(Bluthgen et al, 2006)。的取值范围为0-1, 0表示最泛化, 1表示最特化。以上所有参数利用R (3.4.1)中的bipartite包(2.08)进行计算。

2 结果

2.1 物种分析

捕获的印加孔雀草访花昆虫共有13种, 其中膜翅目1种(32只), 鳞翅目2种(3只), 双翅目9种(19只), 鞘翅目1种(1只) (图1, 表1)。除东方菜粉蝶(Pieris canidia)外, 其他12种昆虫均携带有印加孔雀草的花粉(表2)。13种访花昆虫所携带的花粉共存在12种形态(植物物种) (图2)。所有昆虫上共发现2,614枚花粉, 其中印加孔雀草花粉占89.89%, 其他11种花粉占总花粉数的10.11%, 主要属于龙胆科、唇形科和石竹科等植物类群(图2)。

图1

图1   西藏印加孔雀草种群与传粉昆虫。(A)印加孔雀草种群; (B)膜翅目分舌蜂科一种; (C)双翅目蝇类花蝇科一种; (D)双翅目蝇类丽蝇科一种; (E)鳞翅目红灰蝶; (F)鞘翅目拟步甲科一种。

Fig. 1   Population of Tagetes minuta in Tibet and its pollinators. (A) T. minuta population; (B) Colletidae sp.; (C) Anthomyiidae sp.; (D) Calliphoridae sp.; (E) Lycaena phlaeas; (F) Tenebrionidae sp.


表1   印加孔雀草种群的传粉网络的群落水平参数

Table 1  Community level parameters of pollination network of Tagetes minuta population

特征 Traits网络参数 Parameters
植物种类数 No. of plant species12
访花昆虫种类数 No. of visiting species13
植物与昆虫的连接数量 No. of interactions63
连接度 Connectance0.404
嵌套度 Nestedness temperature14.57
加权嵌套度 Weighted nestedness0.683
特化水平 Specialization level (H°2)0.147

新窗口打开| 下载CSV


表2   印加孔雀草种群的传粉网络中植物物种水平的几个参数

Table 2  Several parameters of plants at the species levels in the pollination network of Tagetes minuta population

代码
Codes
植物
Plant species
传粉者种类
Degree
传粉昆虫比例
Normalised degree
物种强度
Species strength
特化水平
Specialization level ()
A印加孔雀草 Tagetes minuta120.92310.3120.034
B狭叶荆芥 Nepeta souliei80.6150.850.095
C无心菜 Arenaria serphyllifolia50.3850.0860.11
D马先蒿属一种 Pedicularis sp.60.4620.0540.085
E龙胆属一种 Gentiana sp.40.3080.7790.467
F紫草科一种 Boraginaceae sp.60.4620.1070.046
G蓝钟花属一种 Cyananthus sp.50.3850.3280.289
H唇形科一种 Lamiaceae sp.40.3080.1970.099
I唇形科一种 Lamiaceae sp.10.0770.0020.045
J唇形科一种 Lamiaceae sp.30.2310.0370.332
K百合科一种 Liliaceae sp.30.2310.3520.563
L甘青老鹳草 Geranium pylzowianum60.4620.0630.117

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图2

图2   印加孔雀草的传粉昆虫所携带的花粉扫描图。(A)印加孔雀草; (B)狭叶荆芥; (C)无心菜; (D)马先蒿属一种; (E)龙胆属一种; (F)紫草科一种; (G)蓝钟花属一种; (H-J)唇形科; (K)百合科一种; (L)甘青老鹳草。

Fig. 2   A scan of pollens carried by pollinators of Tagetes minuta. (A) Tagetes minuta; (B) Nepeta souliei; (C) Arenaria serphyllifolia; (D) Pedicularis sp.; (E) Gentiana sp.; (F) Boraginaceae sp.; (G) Cyananthus sp.; (H-J) Lamiaceae spp.; (K) Liliaceae sp.; (L) Geranium pylzowianum.


2.2 花粉网络分析

13种昆虫与12种植物共存在63个连接, 占可能的总连接数的40.4% (表1, 图3)。传粉网络的嵌套度为14.57, 网络的特化水平为0.147。印加孔雀草的物种强度高达10.312, 而其他植物的物种强度都小于1; 且印加孔雀草的特化水平在所有物种中最小, 仅为0.034 (表2)。印加孔雀草的传粉者多达12种, 占观察到的访花昆虫的92.3%。12种传粉昆虫中有8种携带超过3种植物的花粉, 表明印加孔雀草的传粉昆虫主要为泛化传粉者。印加孔雀草具有多种有效传粉者, 其中1种膜翅目昆虫(分舌蜂科)和3种双翅目昆虫(食蚜蝇科2种、丽蝇科1种)的物种强度大于1 (表3, 图3)。以上结果表明印加孔雀草的主要传粉者为泛化传粉昆虫中的3种双翅目昆虫和1种膜翅目昆虫。

图3

图3   印加孔雀草的所有访花昆虫携带的花粉网络。上部每个矩形框表示一种传粉者, 下部每个矩形框表示一种植物, 框的宽度与花粉数量成正比。灰线表示植物与传粉者之间的连接。植物及传粉者各编号代表的物种见表2和表3。

Fig. 3   Pollen network carried by pollinators of Tagetes minuta. Upper bars represent insect species and lower bars represent plant species. Bar width is proportional to the frequency of interactions. Lines are relative to the interaction between plants and insects. The codes of species are shown in Table 2 and Table 3.


表3   印加孔雀草种群的传粉网络中传粉昆虫物种水平的几个参数

Table 3  Several parameters of pollinators at the species level in the pollination network of the Tagetes minuta population

代码
Codes
传粉者
Pollinators
昆虫数量
Samples
植物物种
Degree
植物物种比例
Normalised degree
物种强度
Species strength
特化水平
Bee sp.分舌花蜂科Colletidae32110.9171.6110.006
Syr sp.1长尾管蚜蝇Eristalis tenax5100.8332.1940.102
Syr sp.2黑带蚜蝇属Episyrpus370.5830.640.024
Syr sp.3黑带蚜蝇属Episyrpus370.5831.3190.012
Syr sp.4食蚜蝇科Syrphidae120.1670.0130
Fly sp.1丽蝇科Calliphoridae280.6674.0720.016
Fly sp.2丽蝇科Calliphoridae120.1670.2990.139
Fly sp.3蝇科Muscidae110.0830.0140.017
Fly sp.4花蝇科Anthomyiidae140.3330.3030.034
Sar sp.麻蝇科Sarcophadidae240.3330.8230.056
Pier sp.1东方菜粉蝶Pieris canidia120.1670.5010.803
Pier sp.2红灰蝶Lycaena phlaeas210.0830.0030.005
Col sp.拟步甲科Tenebrionidae140.3330.2290.119

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3 讨论

我们的结果表明印加孔雀草的物种强度比群落中其他所有植物的物种强度之和还高, 而特化水平却极低, 仅为0.034 (表2)。同时印加孔雀草利用当地12种昆虫为其传粉, 携带印加孔雀草花粉最多的几种昆虫都为泛化传粉者, 且其中4种传粉昆虫的物种强度大于1, 表明印加孔雀草属于泛化传粉系统并已顺利利用当地传粉昆虫为其传粉。

印加孔雀草能在较短时间内快速占据新生境并融入当地传粉网络, 可能原因有以下几点: (1)生境破碎化和农业开垦等人为干扰使原生植被遭到破坏, 为外来生物入侵创造了有利条件(万方浩等, 2002; 鞠瑞亭等, 2012)。我们对西藏入侵植物的调查发现印加孔雀草主要占据人为干扰比较严重的生境, 比如公路两侧和农业生态系统, 这是因为此类生境可以为先锋入侵物种的奠基提供空余生态位(Weber & Li, 2008), 从而使入侵种能更好地利用资源并建立种群(鞠瑞亭等, 2012)。(2)印加孔雀草具有快速在入侵地建立种群的能力。尽管部分菊科植物的交配系统是自交不亲和的(Brennan et al, 2005; Campbell & Husband, 2007; Hao et al, 2011), 但是有研究表明我国入侵菊科植物中许多种类具有自交亲和或者无融合生殖特性(Hao et al, 2011), 可以保证它们入侵初期在缺少传粉者的情况下快速建立种群。虽然没有对印加孔雀草的交配系统开展详细研究, 但是对印加孔雀草进行套网处理, 结果发现它在套网情况下仍然可以顺利结实, 表明印加孔雀草可能具有自动自交或无融合生殖的能力。(3)印加孔雀草具有万寿菊属植物特有的挥发性物质, 具有浓烈的气味, 不仅对其他植物的生长发育有抑制作用, 还对病虫害和食草动物等有很强的抵制能力, 而且其头状花序多、结果量大、瘦果小且具冠毛, 极易繁殖和传播(张劲林等, 2014)。由于以上因素, 当印加孔雀草进入新生境后, 既能快速建立种群, 还能通过化感作用排斥其他植物, 从而在入侵地形成单一种群(图1A)。

入侵植物在新生境成功定殖后, 通过与当地植物竞争有限的传粉者, 会降低当地植物的传粉频率和效率, 进而影响当地植物的繁殖成功(Lopezaraiza-Mikel et al, 2007; Kaiser-Bunbury et al, 2017)。例如入侵植物千屈菜(Lythrum salicaria)通过大量开花来吸引当地传粉昆虫为其传粉(Goodell & Parker, 2017), 同时导致同域分布的多种本土植物的传粉者访问频率降低, 进而使部分本土植物的结实率显著降低(Flanagan et al, 2010; Goodell & Parker, 2017)。另外一个著名的例子是Kaiser-Bunbury等(2017)通过对塞舌尔马赫岛上4个移除入侵植物和4个没有移除入侵植物的植物-传粉者网络比较发现, 入侵植物不仅显著降低群落内传粉者的物种数量、访问次数和互作多样性, 而且也降低了当地部分野生植物的繁殖成功。这是因为入侵植物可以和当地的泛化传粉者构建稳定连接, 从而融入当地的传粉网络(Padrón et al, 2009), 甚至成为当地群落的核心物种(Lopezaraiza-Mikel et al, 2007), 从而改变当地植物-传粉者网络的结构和功能。在我们的研究样地内, 印加孔雀草花丰富度占所有开花植物的98%以上, 当地昆虫主要访问印加孔雀草, 导致本地开花植物的访花昆虫种类、数量及昆虫的访花频率都可能降低, 从而影响当地传粉网络结构和功能。另外, 印加孔雀草种群花期较晚, 主要在每年的9月初至11月初, 并且是大量植株同时开花, 单株开花数量最多可达上千朵(图1)。我们的结果表明印加孔雀草在西藏主要利用当地的泛化传粉者例如蜂、蝇、食蚜蝇等昆虫为其传粉(图1, 表3)。泛化传粉者会在多种植物之间混访, 由于入侵植物引起的异种花粉干扰结实的不利影响比本土植物更严重(Arceo-Gómez & Ashman, 2016), 因此印加孔雀草花粉对当地植物花粉转移网络和雌性和雄性适合度方面都可能产生负面影响。

不过本研究仍有一些需要注意的地方: (1)我们是通过形态学和孢粉学手段进行的物种鉴定, 不能排除高估种内花粉形态差异大的物种或低估动植物近缘种的物种多度的现象(郎丹丹等, 2018)。今后可结合DNA条形码与高通量测序技术(超级条形码)对昆虫以及花粉混合样品进行定性和定量分析来克服物种的鉴定误差(Richardson et al, 2015)。(2)本研究是通过观察印加孔雀草的传粉者和分析其所携带的花粉种类和数量来构建的传粉者-花粉网络, 很可能高估印加孔雀草在群落中的作用, 同时低估群落中其他植物的作用。不过我们野外调查发现在印加孔雀草群落中极少见到其他植物生存, 可能的原因是印加孔雀草产生的化感物质对其他植物的生存具有很强的抑制作用(张劲林等, 2014)。(3)本研究只探讨了入侵植物印加孔雀草种群的访花昆虫所携带的花粉-植物网络的结构, 而没有选取未受印加孔雀草入侵的群落进行花粉网络对比分析。在今后的研究中需要通过对比分析西藏有无印加孔雀草群落的传粉网络结构变化, 进一步阐明印加孔雀草对当地定性和定量传粉网络(比如对传粉昆虫丰度和多样性、访花频率以及当地植物繁殖成功等)的影响。目前, 我国西藏已发现较多的印加孔雀草群落, 且表现出局部生态危害(许敏和扎西次仁, 2015)。因此, 在群落水平上量化分析入侵植物在当地传粉网络中的地位和功能和对当地传粉网络的影响(方强和黄双全, 2012), 有助于了解其入侵机制并为当地植物群落恢复和生物多样性保护提供理论依据。

附录 Supplementary Material

附录1 印加孔雀草花粉数据矩阵

Appendix 1The matrix of the pollen grains of Tagetes minuta and other plant species carried by pollinators

http://www.biodiversity-science.net/fileup/PDF/2018269-1.pdf

致谢:

感谢王浩在花粉扫描中提供的帮助, 感谢两位匿名审稿人和编辑对文稿提出的宝贵意见和建议。

The authors have declared that no competing interests exist.
作者已声明无竞争性利益关系。

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报道了中国大陆归化植物新记录刺毛峨参(Anthriscus caucalis M.Bieb.)和印加孔雀草(Tagetes minuta L.),并进行了形态描述,提供了凭证标本和活植物照片。

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报道了中国大陆归化植物新记录刺毛峨参(Anthriscus caucalis M.Bieb.)和印加孔雀草(Tagetes minuta L.),并进行了形态描述,提供了凭证标本和活植物照片。

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植物与传粉者之间相互作用, 构成了复杂的传粉网络。近年来, 社会网络分析技术的发展使得复杂生态网络的研究成为可能。从群落水平上研究植物与传粉者之间的互惠关系, 为理解群落的结构和动态以及花部特征的演化提供了全新的视角。传粉网络的嵌套结构说明自然界的传粉服务存在冗余, 而且是相对泛化的物种主导了传粉。在多年或者多季度的传粉网络中, 虽然有很高的物种替换率, 但是其网络结构仍然保持相对稳定, 说明传粉网络对干扰有很强的抗性。尽管有关网络结构和动态的研究逐渐增多, 但传粉网络维持的机制仍不清楚。网络结构可以部分由花部特征与传粉者的匹配来解释, 也受到系统发生的制约, 影响因素还包括群落构建的时间和物种多样性, 以及物种在群落中的位置。开展大尺度群落动态的研究, 为探索不同时间尺度、不同物种多样性水平上的传粉网络的生态学意义提供了条件。但已有的研究仍存在不足, 比如基于访问观察的网络无法准确衡量传粉者的 访问效率和植物间的花粉流动, 以及结果受到调查精度区域研究不平衡的制约等。目前的研究只深入到传粉者携带花粉构成成分的水平, 传粉者访问植物的网络不能代表植物的整个传粉过程。因此, 研究应当更多地深入到物种之间关系对有性生殖的切实影响上。

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传粉网络的研究进展: 网络的结构和动态

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植物与传粉者之间相互作用, 构成了复杂的传粉网络。近年来, 社会网络分析技术的发展使得复杂生态网络的研究成为可能。从群落水平上研究植物与传粉者之间的互惠关系, 为理解群落的结构和动态以及花部特征的演化提供了全新的视角。传粉网络的嵌套结构说明自然界的传粉服务存在冗余, 而且是相对泛化的物种主导了传粉。在多年或者多季度的传粉网络中, 虽然有很高的物种替换率, 但是其网络结构仍然保持相对稳定, 说明传粉网络对干扰有很强的抗性。尽管有关网络结构和动态的研究逐渐增多, 但传粉网络维持的机制仍不清楚。网络结构可以部分由花部特征与传粉者的匹配来解释, 也受到系统发生的制约, 影响因素还包括群落构建的时间和物种多样性, 以及物种在群落中的位置。开展大尺度群落动态的研究, 为探索不同时间尺度、不同物种多样性水平上的传粉网络的生态学意义提供了条件。但已有的研究仍存在不足, 比如基于访问观察的网络无法准确衡量传粉者的 访问效率和植物间的花粉流动, 以及结果受到调查精度区域研究不平衡的制约等。目前的研究只深入到传粉者携带花粉构成成分的水平, 传粉者访问植物的网络不能代表植物的整个传粉过程。因此, 研究应当更多地深入到物种之间关系对有性生殖的切实影响上。

Flanagan RJ, Mitchell RJ, Karron JD ( 2010)

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Oecologia, 164, 445-454.

DOI:10.1007/s00442-010-1693-2      URL     PMID:20585807      [本文引用: 1]

When exotic plant species share pollinators with native species, competition for pollination may lower the reproductive success of natives by reducing the frequency and/or quality of visits they receive. Exotic species often become numerically dominant in plant communities, and the relative abundance of these potential competitors for pollination may be an important determinant of their effects on the pollination and reproductive success of co-occurring native species. Our study experimentally tests whether the presence and abundance of an invasive exotic, Lythrum salicaria L. (Lythraceae), influences reproductive success of a co-flowering native species, Mimulus ringens L. (Phrymaceae). We also examine the mechanisms of competition for pollination and how they may be altered by changes in competitor abundance. We found that the presence of Lythrum salicaria lowered mean seed number in Mimulus ringens fruits. This effect was most pronounced when the invasive competitor was highly abundant, decreasing the number of seeds per fruit by 40% in 2006 and 33% in 2007. Reductions in the number of seeds per fruit were likely due to reduced visit quality resulting from Mimulus pollen loss when bees foraged on neighboring Lythrum plants. This study suggests that visit quality to natives may be influenced by the presence and abundance of invasive flowering plants.

Goodell K, Parker IM ( 2017)

Invasion of a dominant floral resource: Effects on the floral community and pollination of native plants

Ecology, 98, 57-69.

DOI:10.1002/ecy.1639      URL     PMID:28052387      [本文引用: 2]

Abstract Through competition for pollinators, invasive plants may suppress native flora. Community-level studies provide an integrative assessment of invasion impacts and insights into factors that influence the vulnerability of different native species. We investigated effects of the non-native herb Lythrum salicaria on pollination of native species in 14 fens of the eastern United States. We compared visitors per flower for 122 native plant species in invaded and uninvaded fens and incorporated a landscape-scale experiment, removing L. salicaria flowers from three of the invaded fens. Total flower densities were more than three times higher in invaded than uninvaded or removal sites when L. salicaria was blooming. Despite an increase in number of visitors with number of flowers per area, visitors per native flower declined with increasing numbers of flowers. Therefore, L. salicaria invasion depressed visitation to native flowers. In removal sites, visitation to native flowers was similar to uninvaded sites, confirming the observational results and also suggesting that invasion had not generated a persistent buildup of visitor populations. To study species-level impacts, we examined effects of invasion on visitors per flower for the 36 plant species flowering in both invaded and uninvaded fens. On average, the effect of invasion represented about a 20% reduction in visits per flower. We measured the influence of plant traits on vulnerability to L. salicaria invasion using meta-analysis. Bilaterally symmetrical flowers experienced stronger impacts on visitation, and similarity in flower color to L. salicaria weakly intensified competition with the invader for visitors. Finally, we assessed the reproductive consequences of competition with the invader in a dominant flowering shrub, Dasiphora fruticosa. Despite the negative effect of invasion on pollinator visitation in this species, pollen limitation of seed production was not stronger in invaded than uninvaded sites, suggesting little impact of competition for pollinators on its population demography. Negative effects on pollination of native plants by this copiously-flowering invader appeared to be mediated by increases in total flower density that were not matched by increases in pollinator density. The strength of impact was modulated across native species by their floral traits and reproductive ecology. This article is protected by copyright. All rights reserved.

Hao J, Sheng Q, Thomas C, Mark VK, Liu Q ( 2011)

A test of baker’s law: Breeding systems of invasive species of Asteraceae in China

Biological Invasions, 13, 571-580.

DOI:10.1007/s10530-010-9850-4      URL     [本文引用: 2]

Invasive alien plant species are frequently characterized by a high fecundity. However, because suitable pollinators and/or mates may be absent in the new range, it is not clear how they achieve high seed production. According to Baker Law, species capable of uniparental reproduction are more likely to establish after long-distance dispersal than species that rely on suitable mates and pollinators. To test whether self-compatible species, and particularly species capable of autonomous seed set, are more likely to establish and spread, we experimentally assessed the breeding systems of 12 species of Asteraceae that are invasive in China. Among these 12 species of Asteraceae, the percentages of self-compatible species (66.7%) and species capable of autonomous seed set (83.3%), which included self-fertilizing and apomictic species, were significantly larger than expected from the percentages of such species in global data sets of Asteraceae (36.8% and 46.0%, respectively). Furthermore, the number of Chinese provinces in which the invasive alien species occur was significantly positively correlated with the proportion seed set on bagged capitula (i.e. with the degree of autonomous seed set). Among 36 species of Asteraceae that are invasive in China and for which we found breeding-system data in the literature, we also found a higher than expected percentage of self-compatible species (65.7%), and that these self-compatible species are more widespread in China than self-incompatible species. These results support the predictions of Baker Law that self-compatible species, and particularly those capable of autonomous seed production, are more likely to establish and spread in a new range. Therefore, breeding systems of plants should be included as one of the key elements in risk assessment protocols for plant invasiveness.

Jeschke JM, Bacher S, Blackburn TM, Dick JTA, Essl F, Evans T, Gaertner M, Hulme PE, Kühn I, Mrugała A ( 2015)

Defining the impact of non-native species

Conservation Biology, 28, 1188-1194.

DOI:10.1111/cobi.12299      PMID:4282110      [本文引用: 1]

Abstract Non-native species cause changes in the ecosystems to which they are introduced. These changes, or some of them, are usually termed impacts; they can be manifold and potentially damaging to ecosystems and biodiversity. However, the impacts of most non-native species are poorly understood, and a synthesis of available information is being hindered because authors often do not clearly define impact. We argue that explicitly defining the impact of non-native species will promote progress toward a better understanding of the implications of changes to biodiversity and ecosystems caused by non-native species; help disentangle which aspects of scientific debates about non-native species are due to disparate definitions and which represent true scientific discord; and improve communication between scientists from different research disciplines and between scientists, managers, and policy makers. For these reasons and based on examples from the literature, we devised seven key questions that fall into 4 categories: directionality, classification and measurement, ecological or socio-economic changes, and scale. These questions should help in formulating clear and practical definitions of impact to suit specific scientific, stakeholder, or legislative contexts. Definiendo el Impacto de las Especies No-Nativas Resumen Las especies no-nativas pueden causar cambios en los ecosistemas donde son introducidas. Estos cambios, o algunos de ellos, usualmente se denominan como impactos ; estos pueden ser variados y potencialmente da09inos para los ecosistemas y la biodiversidad. Sin embargo, los impactos de la mayoría de las especies no-nativas están pobremente entendidos y una síntesis de información disponible se ve obstaculizada porque los autores continuamente no definen claramente impacto . Discutimos que definir explícitamente el impacto de las especies no-nativas promoverá el progreso hacia un mejor entendimiento de las implicaciones de los cambios a la biodiversidad y los ecosistemas causados por especies no-nativas; ayudar a entender cuáles aspectos de los debates científicos sobre especies no-nativas son debidos a definiciones diversas y cuáles representan un verdadero desacuerdo científico; y mejorar la comunicación entre científicos de diferentes disciplinas y entre científicos, administradores y quienes hacen las políticas. Por estas razones y basándonos en ejemplos tomados de la literatura, concebimos siete preguntas clave que caen en cuatro categorías: direccionalidad, clasificación y medida, cambios ecológicos o socio-económicos, y escala. Estas preguntas deberían ayudar en la formulación de definiciones claras y prácticas del impacto para encajar mejor con contextos científicos, de las partes interesadas o legislativos específicos.

Ju RT, Li H, Shi CJ, Li B ( 2012)

Progress of biological invasions research in China over the last decade

Biodiversity Science, 20, 581-611. (in Chinese with English abstract)

DOI:10.3724/SP.J.1003.2012.31148      URL     [本文引用: 3]

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。

[ 鞠瑞亭, 李慧, 石正人, 李博 ( 2012)

近十年中国生物入侵研究进展

生物多样性, 20, 581-611.]

DOI:10.3724/SP.J.1003.2012.31148      URL     [本文引用: 3]

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。

Kaiser-Bunbury CN, Mougal J, Whittington AE, Valentin T, Gabriel R, Olesen JM, Bluthgen N ( 2017)

Ecosystem restoration strengthens pollination network resilience and function

Nature, 542, 223-227.

DOI:10.1038/nature21071      URL     PMID:28135718      [本文引用: 3]

Land degradation results in declining biodiversity and the disruption of ecosystem functioning worldwide, particularly in the tropics. Vegetation restoration is a common tool used to mitigate these impacts and increasingly aims to restore ecosystem functions rather than species diversity. However, evidence from community experiments on the effect of restoration practices on ecosystem functions is scarce. Pollination is an important ecosystem function and the global decline in pollinators attenuates the resistance of natural areas and agro-environments to disturbances. Thus, the ability of pollination functions to resist or recover from disturbance (that is, the functional resilience) may be critical for ensuring a successful restoration process. Here we report the use of a community field experiment to investigate the effects of vegetation restoration, specifically the removal of exotic shrubs, on pollination. We analyse 64 plant-pollinator networks and the reproductive performance of the ten most abundant plant species across four restored and four unrestored, disturbed mountaintop communities. Ecosystem restoration resulted in a marked increase in pollinator species, visits to flowers and interaction diversity. Interactions in restored networks were more generalized than in unrestored networks, indicating a higher functional redundancy in restored communities. Shifts in interaction patterns had direct and positive effects on pollination, especially on the relative and total fruit production of native plants. Pollinator limitation was prevalent at unrestored sites only, where the proportion of flowers producing fruit increased with pollinator visitation, approaching the higher levels seen in restored plant communities. Our results show that vegetation restoration can improve pollination, suggesting that the degradation of ecosystem functions is at least partially reversible. The degree of recovery may depend on the state of degradation before restoration intervention and the proximity to pollinator source populations in the surrounding landscape. We demonstrate that network structure is a suitable indicator for pollination quality, highlighting the usefulness of interaction networks in environmental management.

Kearns CA, Inouye DW, Waser NM ( 1998)

Endangered mutualisms: The conservation of plant-pollinator interactions

Annual Review of Ecology & Systematics, 29, 83-112.

DOI:10.1146/annurev.ecolsys.29.1.83      URL     [本文引用: 1]

The pollination of flowering plants by animals represents a critical ecosystem service of great value to humanity, both monetary and otherwise. However, the need for active conservation of pollination interactions is only now being appreciated. Pollination systems are under increasing threat from anthropogenic sources, including fragmentation of habitat, changes in land use, modern agricultural practices, use of chemicals such as pesticides and herbicides, and invasions of non-native plants and animals. Honeybees, which themselves are non-native pollinators on most continents, and which may harm native bees and other pollinators, are nonetheless critically important for crop pollination. Recent declines in honeybee numbers in the United States and Europe bring home the importance of healthy pollination systems, and the need to further develop native bees and other animals as crop pollinators. The "pollination crisis" that is evident in declines of honeybees and native bees, and in damage to webs of plant-pollinator interaction, may be ameliorated not only by cultivation of a diversity of crop pollinators, but also by changes in habitat use and agricultural practices, species reintroductions and removals, and other means. In addition, ecologists must redouble efforts to study basic aspects of plant-pollinator interactions if optimal management decisions are to be made for conservation of these interactions in natural and agricultural ecosystems.

Lang DD, Tang M, Zhou X ( 2018)

Qualitative and quantitative molecular construction of plant-pollinator network: Application and prospective

Biodiversity Science, 26, 445-456. (in Chinese with English abstract)

DOI:10.17520/biods.2018058      URL     [本文引用: 1]

传粉者是重要的生态功能提供者,在维持稳定的生态系统和高效的农业生产力中发挥着重要作用。因此,传粉网络的构建和监测工作对评价生态系统平衡和调控农业生产至关重要。该工作的基础就是通过对传粉者及植物的物种鉴定构建其相关性。传统的形态学物种鉴定对分类学专家的专业知识、时间和经验都提出较高的要求。DNA条形码和高通量测序技术(high-throughput sequencing,HTS)的发展及其在传粉网络研究中的应用,提供了高效、准确鉴定传粉者与植物的方法,大大提高了传粉网络构建的效率。本文阐述了传粉网络研究相关的研究方法和技术进展,并提出利用高通量测序技术结合无PCR扩增(PCR-free)的"超级条形码"技术,有望实现以更高的灵敏度和分辨率对混合物种样品进行定性及相对定量的监测。该方法的有效性已在其他生物多样性研究中得以验证,在传粉网络研究中虽处于初始阶段,但应用前景广阔。

[ 郎丹丹, 唐敏, 周欣 ( 2018)

传粉网络构建的定性定量分子研究: 应用与展望

生物多样性, 26, 445-456.]

DOI:10.17520/biods.2018058      URL     [本文引用: 1]

传粉者是重要的生态功能提供者,在维持稳定的生态系统和高效的农业生产力中发挥着重要作用。因此,传粉网络的构建和监测工作对评价生态系统平衡和调控农业生产至关重要。该工作的基础就是通过对传粉者及植物的物种鉴定构建其相关性。传统的形态学物种鉴定对分类学专家的专业知识、时间和经验都提出较高的要求。DNA条形码和高通量测序技术(high-throughput sequencing,HTS)的发展及其在传粉网络研究中的应用,提供了高效、准确鉴定传粉者与植物的方法,大大提高了传粉网络构建的效率。本文阐述了传粉网络研究相关的研究方法和技术进展,并提出利用高通量测序技术结合无PCR扩增(PCR-free)的"超级条形码"技术,有望实现以更高的灵敏度和分辨率对混合物种样品进行定性及相对定量的监测。该方法的有效性已在其他生物多样性研究中得以验证,在传粉网络研究中虽处于初始阶段,但应用前景广阔。

Lopezaraiza-Mikel ME, Hayes RB, Whalley MR, Memmott J ( 2007)

The impact of an alien plant on a native plant-pollinator network: An experimental approach

Ecology Letters, 10, 539-550.

DOI:10.1111/j.1461-0248.2007.01055.x      URL     PMID:17542933      [本文引用: 3]

Studies of pairwise interactions have shown that an alien plant can affect the pollination of a native plant, this effect being mediated by shared pollinators. Here we use a manipulative field experiment, to investigate the impact of the alien plant Impatiens glandulifera on an entire community of coflowering native plants. Visitation and pollen transport networks were constructed to compare replicated I. glandulifera invaded and I. glandulifera removal plots. Invaded plots had significantly higher visitor species richness, visitor abundance and flower visitation. However, the pollen transport networks were dominated by alien pollen grains in the invaded plots and consequently higher visitation may not translate in facilitation for pollination. The more generalized insects were more likely to visit the alien plant, and Hymenoptera and Hemiptera were more likely to visit the alien than Coleoptera. Our data indicate that generalized native pollinators can provide a pathway of integration for alien plants into native visitation systems.

McKinney AM, Goodell K ( 2011)

Plant-pollinator interactions between an invasive and native plant vary between sites with different flowering phenology

Plant Ecology, 212, 1025-1035.

DOI:10.1007/s11258-010-9882-y      URL     [本文引用: 1]

Floral displays of invasive plants have positive and negative impacts on native plant pollination. Invasive plants may also decrease irradiance, which can lead to reduced pollination of native plants. The effects of shade and flowers of invasive plant species on native plant pollination will depend on overlap in flowering phenologies. We examined the effect of the invasive shrub Lonicera maackii on female reproductive success of the native herb Hydrophyllum macrophyllum at two sites: one with asynchronous flowering phenologies (slight overlap) and one with synchronous (complete overlap). At each site, we measured light availability, pollinator visitation, pollen deposition, and seed set of potted H. macrophyllum in the presence and absence of L. maackii. At both sites, understory light levels were lower in plots containing L. maackii. At the asynchronous site, H. macrophyllum received fewer pollinator visits in the presence of L. maackii, suggesting shade from L. maackii reduced visitation to H. macrophyllum. Despite reduced visitation, H. macrophyllum seed set did not differ between treatments. At the synchronous site, H. macrophyllum received more pollinator visits and produced more seeds per flower in the presence of co-flowering L. maackii compared to plots in which L. maackii was absent, and conspecific pollen deposition was positively associated with seed set. Our results support the hypothesis that co-flowering L. maackii shrubs facilitated pollination of H. macrophyllum, thereby mitigating the negative impacts of shade, leading to increased seed production. Phenological overlap appears to influence pollinator-mediated interactions between invasive and native plants and may alter the direction of impact of L. maackii on native plant pollination.

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

[本文引用: 1]

Olesen JM, Bascompte J, Dupont YL, Jordano P ( 2007)

The modularity of pollination networks. Proceedings of the National Academy of Sciences,

USA, 104, 19891-19896.

DOI:10.1073/pnas.0706375104      URL     PMID:18056808      [本文引用: 1]

In natural communities, species and their interactions are often organized as nonrandom networks, showing distinct and repeated complex patterns. A prevalent, but poorly explored pattern is ecological modularity, with weakly interlinked subsets of species (modules), which, however, internally consist of strongly connected species. The importance of modularity has been discussed for a long time, but no consensus on its prevalence in ecological networks has yet been reached. Progress is hampered by inadequate methods and a lack of large datasets. We analyzed 51 pollination networks including almost 10,000 species and 20,000 links and tested for modularity by using a recently developed simulated annealing algorithm. All networks with >150 plant and pollinator species were modular, whereas networks with <50 species were never modular. Both module number and size increased with species number. Each module includes one or a few species groups with convergent trait sets that may be considered as coevolutionary units. Species played different roles with respect to modularity. However, only 15% of all species were structurally important to their network. They were either hubs (i.e., highly linked species within their own module), connectors linking different modules, or both. If these key species go extinct, modules and networks may break apart and initiate cascades of extinction. Thus, species serving as hubs and connectors should receive high conservation priorities.

Padrón B, Traveset A, Biedenweg T, Díaz D, Nogales M, Olesen JM ( 2009)

Impact of alien plant invaders on pollination networks in two archipelagos

PLoS ONE, 4, e6275.

DOI:10.1371/journal.pone.0006275      URL     PMID:2707600      [本文引用: 1]

Mutualistic interactions between plants and animals promote integration of invasive species into native communities. In turn, the integrated invaders may alter existing patterns of mutualistic interactions. Here we simultaneously map in detail effects of invaders on parameters describing the topology of both plant-pollinator (bi-modal) and plant-plant (uni-modal) networks. We focus on the invader Opuntia spp., a cosmopolitan alien cactus. We compare two island systems: Tenerife (Canary Islands) and Menorca (Balearic Islands). Opuntia was found to modify the number of links between plants and pollinators, and was integrated into the new communities via the most generalist pollinators, but did not affect the general network pattern. The plant uni-modal networks showed disassortative linkage, i.e. species with many links tended to connect to species with few links. Thus, by linking to generalist natives, Opuntia remained peripheral to network topology, and this is probably why native network properties were not affected at least in one of the islands. We conclude that the network analytical approach is indeed a valuable tool to evaluate the effect of invaders on native communities.

Potts SG, Imperatrizfonseca V, Ngo HT, Aizen MA, Biesmeijer JC, Breeze TD, Dicks LV, Garibaldi LA, Hill R, Settele J ( 2016)

Safeguarding pollinators and their values to human well-being

Nature, 540, 220-229.

DOI:10.1038/nature20588      URL     PMID:27894123      [本文引用: 1]

Wild and managed pollinators provide a wide range of benefits to society in terms of contributions to food security, farmer and beekeeper livelihoods, social and cultural values, as well as the maintenance of wider biodiversity and ecosystem stability. Pollinators face numerous threats, including changes in land-use and management intensity, climate change, pesticides and genetically modified crops, pollinator management and pathogens, and invasive alien species. There are well-documented declines in some wild and managed pollinators in several regions of the world. However, many effective policy and management responses can be implemented to safeguard pollinators and sustain pollination services.

Powell KI, Krakos KN, Knight TM ( 2011)

Comparing the reproductive success and pollination biology of an invasive plant to its rare and common native congeners: A case study in the genus Cirsium (Asteraceae)

Biological Invasions, 13, 905-917.

DOI:10.1007/s10530-010-9878-5      URL     [本文引用: 1]

Previous studies have examined an association between reproductive success and pollination biology of rare versus widespread species through pair-wise comparisons of native and invasive congeners or rare and common congeners. To determine the importance of reproductive success and pollination biology for an invasive thistle, Cirsium vulgare , we compared it in its invaded range to five, co-occurring native Cirsium species that range from rare to common. Native study species include C. fontinale var. fontinale , C. andrewsii , C. brevistylum , C. occidentale , and C. quercetorum . We compared all species鈥 reproductive success, insect visitation rate and composition, autonomous self-pollination, and level of pollen limitation in multiple populations. Species differed in their reproductive success; the invasive C. vulgare produced more flower heads per plant than most native species. C. vulgare attracted more visitors than its congeners. In addition, reproductive success and insect visitation significantly varied between populations within species, mainly due to aphid infestation in one population of C. occidentale . Unlike the rare species ( C. fontinale and andrewsii ), C. vulgare did not require a pollinator for high-levels of seed production. The remaining native species set fewer seeds than C. vulgare without a pollinator. However, differences in insect visitation and autonomous self-pollination did not lead to differences in pollen limitation across species or between populations. This result suggests that factors other than pollination biology determine the difference in reproductive success of these species. However, high levels of autonomous self-pollination and generalist insect visitation may allow the invasive C. vulgare to easily establish new populations from low numbers of propagules. Our study provides one contrast that should build towards a larger comparative analysis to examine general patterns in the relationship between reproductive success, pollination biology, rare and invasive species, and our ability to predict biological invasions in introduced species.

Richardson RT, Lin CH, Sponsler DB, Quijia JO, Goodell K, Johnson RM ( 2015)

Application of ITS2 metabarcoding to determine the provenance of pollen collected by honey bees in an agroecosystem

Applications in Plant Sciences, 3, 235-250.

DOI:10.3732/apps.1400066      URL     PMID:25606352      [本文引用: 1]

Abstract PREMISE OF THE STUDY: Melissopalynology, the identification of bee-collected pollen, provides insight into the flowers exploited by foraging bees. Information provided by melissopalynology could guide floral enrichment efforts aimed at supporting pollinators, but it has rarely been used because traditional methods of pollen identification are laborious and require expert knowledge. We approach melissopalynology in a novel way, employing a molecular method to study the pollen foraging of honey bees (Apis mellifera) in a landscape dominated by field crops, and compare these results to those obtained by microscopic melissopalynology. 61 METHODS: Pollen was collected from honey bee colonies in Madison County, Ohio, USA, during a two-week period in midspring and identified using microscopic methods and ITS2 metabarcoding. 61 RESULTS: Metabarcoding identified 19 plant families and exhibited sensitivity for identifying the taxa present in large and diverse pollen samples relative to microscopy, which identified eight families. The bulk of pollen collected by honey bees was from trees (Sapindaceae, Oleaceae, and Rosaceae), although dandelion (Taraxacum officinale) and mustard (Brassicaceae) pollen were also abundant. 61 DISCUSSION: For quantitative analysis of pollen, using both metabarcoding and microscopic identification is superior to either individual method. For qualitative analysis, ITS2 metabarcoding is superior, providing heightened sensitivity and genus-level resolution.

Schemske DW ( 1983) Limits to Specialization and Coevolution in Plant-Animal Mutualisms. Chicago University Press, Chicago.

[本文引用: 1]

Sun SG, Lu B, Lu XM, Huang SQ ( 2018)

On reproductive strategies of invasive plants and their impacts on native plants

Biodiversity Science, 26, 457-467. (in Chinese with English abstract)

DOI:10.17520/biods.2017294      URL     [本文引用: 1]

理解入侵生物的繁殖策略是阐明生物入侵机制的一个重要方面。入侵植物常表现出一些共同的繁殖特征,如以两性花为主的性系统、自动自交为主的繁育系统或不依赖传粉媒介的无融合生殖和无性繁殖以及高生殖投资的资源配置策略等。成功入侵的外来植物通过影响本土的传粉者,在种群和群落水平上影响本土植物的有性繁殖,甚至促使某些本土植物在繁殖对策和表型性状上发生快速转变。目前,入侵植物繁殖策略及其生态效应的研究多侧重于入侵种的快速演化,而有关外来植物与本土植物间的相互影响及其可能存在的协同适应研究还较为缺乏。探讨本土植物在外来种入侵压力下的繁殖对策和响应机制,将丰富人们对物种间竞争、共存及群落构建等机制的深入了解。从繁殖和适应的角度探求入侵植物与本土植物之间的复杂关系,将有助于解析生物入侵的机制及人类干扰下的物种演化规律,也为预测和防控入侵植物提供科学依据。

[ 孙士国, 卢斌, 卢新民, 黄双全 ( 2018)

入侵植物的繁殖策略以及对本土植物繁殖的影响

生物多样性, 26, 457-467.]

DOI:10.17520/biods.2017294      URL     [本文引用: 1]

理解入侵生物的繁殖策略是阐明生物入侵机制的一个重要方面。入侵植物常表现出一些共同的繁殖特征,如以两性花为主的性系统、自动自交为主的繁育系统或不依赖传粉媒介的无融合生殖和无性繁殖以及高生殖投资的资源配置策略等。成功入侵的外来植物通过影响本土的传粉者,在种群和群落水平上影响本土植物的有性繁殖,甚至促使某些本土植物在繁殖对策和表型性状上发生快速转变。目前,入侵植物繁殖策略及其生态效应的研究多侧重于入侵种的快速演化,而有关外来植物与本土植物间的相互影响及其可能存在的协同适应研究还较为缺乏。探讨本土植物在外来种入侵压力下的繁殖对策和响应机制,将丰富人们对物种间竞争、共存及群落构建等机制的深入了解。从繁殖和适应的角度探求入侵植物与本土植物之间的复杂关系,将有助于解析生物入侵的机制及人类干扰下的物种演化规律,也为预测和防控入侵植物提供科学依据。

Wan FH, Guo JY, Wang DH ( 2002)

Alien invasive species in China: Their damages and management strategies

Biodiversity Science, 10, 119-125. (in Chinese with English abstract)

DOI:10.3321/j.issn:1005-0094.2002.01.015      URL     [本文引用: 1]

本文探讨了外来入侵生物的概念及其在我国的危害状况、入侵原因 ,提出了外来入侵生物的预防及管理对策。随着国际贸易往来和旅游业的发展 ,生物入侵在我国不断加剧 ,正在成为威胁我国生物多样性与生态环境的重要因素之一。外来入侵种的生态代价是造成本地物种多样性不可弥补的消失以及物种的灭绝 ,其经济代价是农林牧渔业产量与质量的惨重损失与高额的防治费用。生物入侵在我国大部分是由于人为因素引起的。这些因素包括 :缺乏对引进种的利益与风险进行评估、淡薄的生态意识与不顾生态后果的经济利益驱使下的盲目引进、缺乏严格的科学监管体系或监管不力、缺乏全面检疫的体系与机制。外来入侵生物的综合性与系统性研究已成为当今我国生态环境保护、农业生产和经济可持续发展的重大研究领域。我国对外来入侵生物的预防与管理应着重于国家能力、研究能力、监测与管理能力三大体系的建设上。根据我国国情和目前的紧急现状应制定出优先行动计划。对特定外来种的入侵生物学基础研究、特定生态系统或地理区域入侵种现状及影响的关键评估研究、特定外来入侵生物对生态环境影响的风险评估体系及经济损失的模式研究、发展控制外来有害生物的环保型技术与方法研究、外来生物受控制后生态系统的恢复与栖息生境的复

[ 万方浩, 郭建英, 王德辉 ( 2002)

中国外来入侵生物的危害与管理对策

生物多样性, 10, 119-125.]

DOI:10.3321/j.issn:1005-0094.2002.01.015      URL     [本文引用: 1]

本文探讨了外来入侵生物的概念及其在我国的危害状况、入侵原因 ,提出了外来入侵生物的预防及管理对策。随着国际贸易往来和旅游业的发展 ,生物入侵在我国不断加剧 ,正在成为威胁我国生物多样性与生态环境的重要因素之一。外来入侵种的生态代价是造成本地物种多样性不可弥补的消失以及物种的灭绝 ,其经济代价是农林牧渔业产量与质量的惨重损失与高额的防治费用。生物入侵在我国大部分是由于人为因素引起的。这些因素包括 :缺乏对引进种的利益与风险进行评估、淡薄的生态意识与不顾生态后果的经济利益驱使下的盲目引进、缺乏严格的科学监管体系或监管不力、缺乏全面检疫的体系与机制。外来入侵生物的综合性与系统性研究已成为当今我国生态环境保护、农业生产和经济可持续发展的重大研究领域。我国对外来入侵生物的预防与管理应着重于国家能力、研究能力、监测与管理能力三大体系的建设上。根据我国国情和目前的紧急现状应制定出优先行动计划。对特定外来种的入侵生物学基础研究、特定生态系统或地理区域入侵种现状及影响的关键评估研究、特定外来入侵生物对生态环境影响的风险评估体系及经济损失的模式研究、发展控制外来有害生物的环保型技术与方法研究、外来生物受控制后生态系统的恢复与栖息生境的复

Wang H, Cao GX, Wang LL, Yang YP, Zhang ZQ, Duan YW ( 2017)

Evaluation of pollinator effectiveness based on pollen deposition and seed production in a gynodieocious alpine plant, Cyananthus delavayi

Ecology and Evolution, 7, 8156-8160.

DOI:10.1002/ece3.3391      URL     [本文引用: 1]

Examining variations in pollinator effectiveness can enhance our understanding of how pollinators and plants interact. Pollen deposition and seed production after a single visit by a pollinator are often used to estimate pollinator effectiveness. However, seed production is not always directly related to pollen deposition because not all pollen grains that are deposited on a stigma are compatible or conspecific. In the field, we tested pollinator effectiveness based on pollen deposition and the resulting seed production after single visits by different pollinator groups in a gynodieocious alpine plant Cyananthus delavayi (Campanulaceae). Our results showed that mean pollen deposition was generally inconsistent with mean seed production when comparisons were performed among different pollinator groups and sexes. In general, the correlations were not significant between pollen deposition and seed production in both perfect and female flowers after single visits by halictid bees, bumble bees, and hoverflies. We suggest seed set of virgin flowers after single visits is a more reliable indicator of pollinator effectiveness than pollen deposition and would be a better indicator of pollinator effectiveness for future studies.

Weber E, Li B ( 2008)

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

BioScience, 58, 437-444.

DOI:10.1016/j.str.2009.09.008      URL     [本文引用: 3]

Developing and transitional countries undergoing rapid economic development will face growing problems with biological invasions because international commerce will bring new invaders. We assessed the potential for plant invasions in China by comparing the country's current invasive flora with that of the United States, a nation of similar size and latitudinal span but with a different history of plant introductions. Invasive plant species richness in the United States is about twice as high as it is in China. The remarkably lower fraction of invasive woody plants in China suggests that more alien trees and shrubs could invade Chinese habitats. Road density correlated with invasive plant species density among geographical units, with numbers for China lower than those for the United States. The data suggest that China has been invaded less than the United States has, and that the potential for new plant invasions in China is high. Measures toward preventing biological invasions are needed and timely teps taken now can prevent adverse impacts from future invasions.

Weber E, Sun SG, Li B ( 2008)

Invasive alien plants in China: Diversity and ecological insights

Biological Invasions, 10, 1411-1429.

DOI:10.1007/s10530-008-9216-3      URL    

China current invasive alien plant species were analyzed with regard to their floristic status, biological attributes and invasion status elsewhere. Most of the 270 species identified were annuals, followed by perennial herbs. Woody perennials made only about 10% of the species. The invasives were comprised of 59 families, the largest being Asteraceae, Poaceae, and Brassicaceae. The genera with most invasive species were Amaranthus , Ipomoea , and Solanum . Most of the species originated from the New World, notably from South America. About one-third of the species were serious invaders of natural habitats in countries other than China. The proportion of invasive alien plants in province floras ranged from 0.5 to 3.8%, absolute numbers from nine to 117 species per province. Density of invasive species was correlated positively with native species density at provincial scale. The results demonstrate that in China invasive plants are present throughout the country, with a particularly high species richness in the Southeast. The ecological diversity of invasive plants suggests wide ranging impacts which need to be assessed.

Xie Y, Li ZY, Gregg WP, Dianmo L ( 2001)

Invasive species in China—An overview

Biodiversity and Conservation, 10, 1317-1341.

DOI:10.1023/A:1016695609745      URL     [本文引用: 1]

China is a vast country with rich biodiversity, which makes China especially vulnerable to invasive species. It has a long history of introduction of non-native species, especially those with perceived beneficial impacts. Its rapid economic development, including an explosive growth in international trade and transportation, has increased the potential for new introductions. Currently, alien species are widespread in the country, occur in many ecosystems, represent most major taxonomic groups, and are introduced unintentionally as well as intentionally for cultivation. The paper lists various cases of invasive species which have caused significant threats or damages to local natural or artificial ecosystems, and indicates that two example industries (fresh water fisheries and lawn grasses) have brought or tend to bring in many invasive species and hence have caused or will cause changes and loss of biodiversity in local ecosystems. Based on these studies, it is suggested that China combat the problem through enhancing awareness, development of a database on invasive species, strengthening international co-operation, preparing case studies and introducing the necessary legislation, regulations and monitoring.

Xu M, Tashi T ( 2015)

A newly naturalized plant in Qinghai-Tibet Plateau

Guihaia, 35, 554-555. (in Chinese with English abstract)

DOI:10.11931/guihaia.gxzw201310020      URL     [本文引用: 2]

该文报道了青藏高原一归化植物新纪录印加孔雀草(Tagetes minuta L.).

[ 许敏, 扎西次仁 ( 2015)

青藏高原一新归化种

广西植物, 35, 554-555.]

DOI:10.11931/guihaia.gxzw201310020      URL     [本文引用: 2]

该文报道了青藏高原一归化植物新纪录印加孔雀草(Tagetes minuta L.).

Zhang JL, YF, Bian Y, Liu RS, Jiang L ( 2014)

A new kind of invasive plant from mainland China—Tagetes minuta L

Plant Quarantine, 28(2), 65-67. (in Chinese with English abstract)

URL     [本文引用: 3]

菊科万寿菊属的印加孔雀草国内报道极少,只FloraofChina中记载,2006年台湾发现了归化的印加孔雀草。2011年,在北京郊区发现印加孔雀草野生群落,局部呈现生态危害。本文介绍了万寿菊属的属下分类和印加孔雀草的形态特征和生物学特性、地理分布、入侵性分析、危害、传入途径分析和控制等。

[ 张劲林, 吕玉峰, 边勇, 刘若思, 江璐 ( 2014)

中国境内(内地)一种新的入侵植物——印加孔雀草

植物检疫, 28(2), 65-67.]

URL     [本文引用: 3]

菊科万寿菊属的印加孔雀草国内报道极少,只FloraofChina中记载,2006年台湾发现了归化的印加孔雀草。2011年,在北京郊区发现印加孔雀草野生群落,局部呈现生态危害。本文介绍了万寿菊属的属下分类和印加孔雀草的形态特征和生物学特性、地理分布、入侵性分析、危害、传入途径分析和控制等。

Zhu SX, Qin HN, Chen YL ( 2005)

Alien species of Compositae in China

Guihaia, 25, 69-76. (in Chinese with English abstract)

DOI:10.3969/j.issn.1000-3142.2005.01.014      URL     [本文引用: 1]

全面整理了中国菊科植物外来种,确认迄今为止共计有74属148种(含亚种),并根据外来种的类型和用途进行了归类,分析了菊科外来入侵种的生物学特性,中国菊科外来种引种原因和途径及对我国生态系统的影响,为全面研究我国外来种、消除外来入侵种提供参考.

[ 朱世新, 覃海宁, 陈艺林 ( 2005)

中国菊科植物外来种概述

广西植物, 25, 69-76.]

DOI:10.3969/j.issn.1000-3142.2005.01.014      URL     [本文引用: 1]

全面整理了中国菊科植物外来种,确认迄今为止共计有74属148种(含亚种),并根据外来种的类型和用途进行了归类,分析了菊科外来入侵种的生物学特性,中国菊科外来种引种原因和途径及对我国生态系统的影响,为全面研究我国外来种、消除外来入侵种提供参考.

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