生境异质性及源株密度对互花米草入侵力的影响

外来物种互花米草(Spartina alterniflora)被引种到我国海岸带后,已带来一系列生态危害,严重影响了入侵地生态系统结构和功能,深入了解其入侵机制是实施有效控制的基础和理论依据。长江口的滩涂湿地是我国典型的异质潮滩,拥有多种生境。为了探究异质潮滩环境中互花米草初始源株对其入侵力的影响,研究通过野外移栽实验,模拟不同源株密度,探讨潮滩湿地生境异质性及源株密度对互花米草定居、生长和扩散的影响。结果表明:1)生境异质性及源株密度均对互花米草存活率、株高及种群增长率有极显著影响(P0.01)。2)至生长季末,植被区内互花米草存活率、株高均显著高于前沿光滩(P0.05)。此外,淤泥质潮滩植被区内互花米草株高、种群增长率也显著高于粉砂质潮滩植被区(P0.05)。3)互花米草入侵植被区不受源株密度的限制,但随源株密度增加,互花米草种群增长率显著降低(P0.05),而株高无显著差异(P0.05);在光滩区,只有高源株密度才可入侵成功。4)异质生境的冲淤动态、淹水时间和淹水深度是影响互花米草存活和生长的主要环境因子。丰富了河口湿地互花米草入侵力的机理研究,也为我国开展互花米草防控及海岸带保护、修复和管理提供了科学依据,具有重要的理论和实践意义。     

入侵植物互花米草——生物学、生态学及管理

近20余年来, 互花米草Spartina alterniflora Loisel.在我国沿海及河口滩涂快速扩散, 已成为我国海岸盐沼中最重要的入侵植物。本文介绍了互花米草的形态学、繁殖生物学、生物系统学、分布及全球范围内的入侵途径和历史, 探讨了互花米草与沿海滩涂非生物环境、生物环境之间的相互作用及人类活动在互花米草成功入侵中的作用, 并得出如下结论: 互花米草的成功入侵是在人类活动的影响下, 其入侵力(互花米草自身的生物学特性)与入侵生境的可入侵性(滩涂环境对入侵的抵抗力)相互作用的结果, 其入侵可能对被入侵地的自然环境、生物多样性、生态系统乃至经济生活带来一系列影响。最后, 提出对互花米草的控制只能通过早期预测并保持监控、一旦发现便及时根除的方法来进行。同时,我国应加强立法与相关知识的普及, 最大限度地降低互花米草乃至其他外来生物入侵的可能性。     

中国东部沿海互花米草种群生活史特征的纬度变异与可塑性

互花米草(Spartina alterniflora)于20世纪70年代被引入中国,目前已在东部沿海盐沼湿地中广泛分布,成为海岸带盐沼中危害严重的入侵植物之一.为了研究互花米草在中国入侵区中的适应机制,揭示遗传分化和表型可塑性在该物种成功入侵中的作用,本研究沿纬度梯度在南起广东(22°N)、北至天津(39°N)的沿海样带上采集了10个种群的样本,通过同质园实验比较了不同纬度来源的种群在,生活史和生长特征方面是否存在遗传分化,并平行设置高低两个水位处理以比较互花米草对水位变化(不同高程生境条件)的可塑性反应.结果表明,在所研究的互花米草17个性状中有12个存在显著的种群间差异.其中,平均开花日期和相对生长率(植株高度)表现出显著的纬度梯度变异:随着纬度的升高,开花时间提前,相对生长速率(植株高度)趋于增加.同时17个性状中有9个在不同水位处理之间存在显著差异.这些结果表明,遗传分化可能是互花米草能够快速占据广阔分布区的重要原因之一,而表型可塑性可能对互花米草在小尺度上占据不同高程环境的过程有重要作用.     

上海崇明东滩外来物种互花米草二次入侵过程

选取上海崇明东滩“刈割+水位调节”集成技术治理互花米草示范工程样地,对互花米草二次入侵过程进行了2年的监测.结果表明:大量互花米草实生苗在春季随潮水沿潮沟等通道向恢复自然水文区扩散定居,2年即可形成与周边群落无明显差异的二次入侵群落,而群落边缘通过无性繁殖的扩散距离有限.在保持淹水控制区,互花米草两年累计扩散距离小于1 m.在物理隔离区,2年未监测到互花米草的二次入侵现象.实生苗在春季的快速拓植是互花米草实现种群二次入侵的关键.彻底消除治理区周边互花米草扩散源,设置隔离带以阻止其向治理区的扩散,或采用种植芦苇等土著物种进行生物替代,是防止互花米草二次入侵的主要措施.     

互花米草群落功率最大化倾向

植物新种进入新区域后,通过扰动或搅局,改变生态系统的结构,形成有特色的自组 织生态系统,趋向新条件下的功率最大化.本文应用能值分析的方法,分别对苏北互花米草 生态系统和光滩生态系统进行能值分析和系统评估.结果表明：互花米草生态系统每年能值 产出比光滩生态系统高1.52E+18 sej.能值密度是光滩的4.72倍,基础能值产出率约为光滩生态系统的5倍.互花米草生态系统能更有效地利用能量,增加系统内部的 能值贮存,实现系统内部的功率最大化.互花米草入侵某区域滩涂后,通过自组织促使整个互花米草生态系统趋于功率最大化,充分发挥生态服务功能,但由于其繁殖扩散过快,导致种群爆发,产生了一系列负面效应,如抑制本土物种、侵占航道、危害贝类养殖等.     

基于个体的空间显性模型和遥感技术模拟入侵植物扩张机制

基于个体的空间显性模型和遥感技术,以互花米草为例,模拟了自1997到2010年的种群扩张动态,揭示了土地利用变化与潮间带高程的影响;并通过全局敏感性分析揭示了种子扩散、成体存活率、有性和无性繁殖等种群统计学特征对互花米草种群扩张的相对重要性。研究结果发现:1)有性繁殖与无性繁殖共同决定互花米草种群快速扩张;2)潮间带高程和土地利用变化显著影响模型预测的精度,对互花米草种群扩张有非常重要的影响;3)成体存活率与种子长距离扩散是影响互花米草种群扩张速度最重要的因素;无性繁殖比有性繁殖对种群扩张的影响更大;种子长距离扩散比本地扩散更为重要,同时,小概率的种子长距离扩散事件对种群扩张有非常重要的影响。为了经济有效地控制外来入侵植物的扩张,应该抑制种子的长距离扩散和移除种子长距离扩散形成的位于入侵前沿的小斑块。     

土著昆虫素毒蛾对入侵植物互花米草地理种群的选择性

外来植物入侵后会改变其对入侵地植食性昆虫的防御能力以应对入侵地生物环境的变化,因此,对土著昆虫防御能力变化的研究将有助于解释外来植物成功入侵的机制。互花米草(Spartina alterniflora)是广泛入侵中国东部沿海地区的外来植物,研究其入侵后对本地植食性昆虫的响应,可从一个侧面部分地回答其成功入侵的生态机制。利用Y型嗅觉仪,结合室内取食实验,我们比较了中国的土著昆虫素毒蛾(Laelia coenosa)对互花米草3个原产地种群和5个入侵地种群的选择偏好,这些种群分别来自美国的德克萨斯(Texas Point)、卡纳维拉尔国家海岸(Canaveral National Seashore)、佛罗里达大西洋大学(Florida Atlantic University),以及中国的唐海、天津、盐城、崇明、珠海。结果表明,虽然素毒蛾幼虫对互花米草不同地理种群叶片气味没有显著的选择偏好,但对原产地种群的取食相对选择系数显著高于入侵地种群,说明互花米草入侵地种群对素毒蛾的抵抗能力相对较强。这从某种层面上可以推测互花米草入侵中国东部沿海地区以后,其对植食性昆虫取食的防御能力有所增强,而这种能力将在一定程度上减少素毒蛾等入侵地植食动物对它的攻击。     

互花米草入侵对鸟类的生态影响

生物入侵威胁本地物种生存,破坏生态系统的结构和功能,是导致全球生物多样性丧失的重要原因之一。外来植物是入侵生物中的重要一类,可以显著改变本地植被群落,并影响其他生物类群。鸟类作为生态系统中的较高营养级,对由入侵植物引起的栖息地变化十分敏感。互花米草自引入中国沿海以来,其分布区域不断扩散,多数研究认为互花米草入侵造成本地生物多样性降低和生态系统退化。系统梳理了互花米草入侵对鸟类栖息地、食物资源、繁殖、群落等方面的生态影响。主要负面影响有:(1)植被群落结构不利于鸟类栖息、筑巢、觅食;(2)鸟类食物资源丰度和多样性下降;(3)本地鸟类种群数量和物种多样性显著下降。在我国东部沿海湿地,互花米草入侵已经显著改变了植被与鸟类分布格局。但随着入侵历史的增长,少数小型雀形目鸟类却可以逐渐适应互花米草生境。互花米草入侵为某些非本地鸟类提供了空白生态位,在一定程度上丰富了本地物种多样性,对互花米草的快速清除反而可能不利于已适应并依赖互花米草生境的鸟类。综上,认为互花米草入侵对鸟类群落甚至整个生态系统的影响可能需要更多研究进行综合评价,应开展长期、大尺度、多因子的监测研究和多物种比较研究,建立生态评价模型并制定科学有效的互花米草管理对策。     

互花米草入侵影响下闽江河口湿地土壤有效硅的时空变化特征

2016年1—12月,选择闽江河口鳝鱼滩的短叶茳芏湿地、互花米草湿地以及二者的交错带湿地为研究对象,采用定位研究方法探讨了互花米草入侵影响下湿地土壤有效硅含量的时空变化特征。结果表明:互花米草入侵影响下3块湿地土壤有效硅含量随时间推移整体呈波动上升趋势;互花米草入侵显著提高了鳝鱼滩湿地30—60 cm土层土壤有效硅含量(P0.01),与短叶茳芏湿地相比,交错带湿地和互花米草湿地30—60 cm土层土壤有效硅含量分别增加了8.56%和19.97%,逐步线性回归分析表明土温和电导是影响其变化的重要因素(P0.01)。研究互花米草入侵影响下湿地土壤有效硅含量的变化特征,对于揭示湿地生态系统生源要素硅生物地球化学循环过程以及互花米草入侵及其扩张机制具有重要意义。     

互花米草黄酮含量分析及其生态学意义

互花米草原产于北美洲大西洋沿岸和墨西哥海湾,经引种和其他原因,已扩散到包括中国在内的多个国家和地区,并对被入侵地生态系统的结构和功能均造成了显著影响。有关该物种的入侵机制和控制策略是当前入侵生态学研究领域的热点。互花米草生物质材料中富含黄酮类物质,与该物种向海扩张的能力可能存在一定联系。鉴于黄酮广泛的应用价值,能否借助黄酮成分的开发带动生物质的收割,以达到有效控制该物种的目的,也是当前研究者所关注的问题。以芦丁为标准参照物,NaNO2-Al（ NO3）3法比较温室互花米草不同器官中总黄酮含量的差异,分析盐城滩涂互花米草越冬芽黄酮含量与所在地理位置之间的关系,采用L9（34）正交试验对干储互花米草叶总黄酮的提取工艺进行优化。互花米草叶片中黄酮类物质远高于根、根状茎、茎、叶鞘、颖稃及种子。盐城滩涂上互花米草越冬芽的黄酮含量随着由陆向海方向基本呈升高趋势。互花米草总黄酮最佳提取工艺为料液比1∶30,乙醇浓度70％,水浴温度80℃,水浴时间2h。互花米草植物材料中,尤其在叶部,黄酮含量十分丰富。该黄酮物质与互花米草在海岸带盐沼生态系统中的适应和向海扩张也存在正相关关系,可作为探索互花米草入侵机制的一个方向。利用本研究中所得优化提取工艺可从互花米草干储生物质中获得稳定的黄酮来源。配合黄酮经济成分的开发而推动生物质材料的收割利用,将成为互花米草生态控制的一项潜在策略。     

局域种群的Allee效应和集合种群的同步性

从包含Allee效应的局域种群出发,建立了耦合映像格子模型,即集合种群模型.通过分析和计算机模拟表明:(1)当局域种群受到Allee效应强度较大时,集合种群同步灭绝;(2)而当Allee效应强度相对较弱时,通过稳定局域种群动态(减少混沌)使得集合种群发生同步波动,而这种同步波动能够增加集合种群的灭绝风险;(3)斑块间的连接程度对集合种群同步波动的发生有很大的影响,适当的破碎化有利于集合种群的续存.全局迁移和Allee效应结合起来增加了集合种群同步波动的可能,从而增加集合种群的灭绝风险.这些结果对理解同步性的机理、利用同步机理来制定物种保护策略和害虫防治都有重要的意义.     

Evolution of dispersal traits along an invasion route in the wind‐dispersed Senecio inaequidens (Asteraceae)

In introduced organisms, dispersal propensity is expected to increase during range expansion. This prediction is based on the assumption that phenotypic plasticity is low compared to genetic diversity, and an increase in dispersal can be counteracted by the Allee effect. Empirical evidence in support of these hypotheses is however lacking. The present study tested for evidence of differentiation in dispersal‐related traits and the Allee effect in the wind‐dispersed invasive Senecio inaequidens (Asteraceae). We collected capitula from individuals in ten field populations, along an invasion route including the original introduction site in southern France. In addition, we conducted a common garden experiment from field‐collected seeds and obtained capitula from individuals representing the same ten field populations. We analysed phenotypic variation in dispersal traits between field and common garden environments as a function of the distance between populations and the introduction site. Our results revealed low levels of phenotypic differentiation among populations. However, significant clinal variation in dispersal traits was demonstrated in common garden plants representing the invasion route. In field populations, similar trends in dispersal‐related traits and evidence of an Allee effect were not detected. In part, our results supported expectations of increased dispersal capacity with range expansion, and emphasized the contribution of phenotypic plasticity under natural conditions.     

Invasion speed is affected by geographical variation in the strength of Allee effects

Allee effects can play a critical role in slowing or preventing the establishment of low density founder populations of non-indigenous species. Similarly, the spread of established invaders into new habitats can be influenced by the degree to which small founder populations ahead of the invasion front are suppressed through Allee effects. We develop an approach to use empirical data on the gypsy moth, a non-indigenous invader in North America, to quantify the Allee threshold across geographical regions, and we report that the strength of the Allee effect is subject to spatial and temporal variability. Moreover, we present what is to our knowledge the first empirical evidence that geographical regions with higher Allee thresholds are associated with slower speeds of invasion.     

Clustered or scattered? The impact of habitat quality clustering on establishment and early spread

The match between the environmental conditions of an introduction area and the preferences of an introduced species is the first prerequisite for establishment. Yet, introduction areas are usually landscapes, i.e. heterogeneous sets of habitats that are more or less favourable to the introduced species. Because individuals are able to disperse after their introduction, the quality of the habitat surrounding the introduction site is as critical to the persistence of introduced populations as the quality of the introduction site itself. Moreover, demographic mechanisms such as Allee effects or dispersal mortality can hamper dispersal and affect spread across the landscape, in interaction with the spatial distribution of favourable habitat patches. In this study, we investigate the impact of the spatial distribution of heterogeneous quality habitats on establishment and early spread. First, we simulated introductions in one‐dimensional landscapes for different dispersal rates and either dispersal mortality or Allee effects. The landscapes differed by the distribution of favourable and less favourable habitats, which were either clustered into few large aggregates of the same quality or scattered into multiple smaller ones. Second, we tested the predictions of simulations by performing experimental introductions of hymenopteran parasitoids (Trichogramma chilonis) in ‘clustered’ and ‘scattered’ microcosm landscapes. Results highlighted two impacts of the clustering of favourable habitat: by decreasing the risks of dispersal from the introduction site to unfavourable habitat early during the invasion, it increased establishment success. However, by increasing the distance between favourable habitat patches, it also hindered the subsequent spread of introduced species over larger areas.     

The role of Allee effects in gypsy moth, <Emphasis Type="Italic">Lymantria dispar</Emphasis> (L.), invasions

Allee effects have been applied historically in efforts to understand the low-density population dynamics of rare and endangered species. Many biological invasions likewise experience the phenomenon of decreasing population growth rates at low population densities because most founding populations of introduced nonnative species occur at low densities. In range expansion of established species, the initial colonizers of habitat beyond the organism’s current range are usually at low density, and thus could be subject to Allee dynamics. There has been consistent empirical and theoretical evidence demonstrating, and in some cases quantifying, the role of Allee dynamics in the gypsy moth, Lymantria dispar (L.), invasion of North America. In this review, we examine the potential causes of the Allee effect in the gypsy moth and highlight the importance of mate-finding failure as a primary mechanism behind an Allee effect, while the degree to which generalist predators induce an Allee effect remains unclear. We then explore the role of Allee effects in the establishment and spread dynamics of the gypsy moth system, which conceptually could serve as a model system for understanding how Allee effects manifest themselves in the dynamics of biological invasions.     

The downward spiral: eco‐evolutionary feedback loops lead to the emergence of ‘elastic’ ranges

In times of severe environmental changes and resulting shifts in the geographical distribution of animal and plant species it is crucial to unravel the mechanisms responsible for the dynamics of species’ ranges. Without such a mechanistic understanding, reliable projections of future species distributions are difficult to derive. Species’ ranges may be highly dynamic. One particularly interesting phenomenon is range contraction following a period of expansion, referred to as ‘elastic’ behaviour. It has been proposed that this phenomenon occurs in habitat gradients, which are characterized by a negative cline in selection for dispersal from the range core towards the margin, as one may find, for example, with increasing patch isolation. Using individual‐based simulations and numerical analyses we show that Allee effects are an important determinant of range border elasticity. If only intra‐specific processes are considered, Allee effects are even a necessary condition for ranges to exhibit elastic behavior. The eco‐evolutionary interplay between dispersal evolution, Allee effects and habitat isolation leads to lower colonization probability and higher local extinction risk after range expansions, which result in an increasing amount of marginal sink patches and consequently, range contraction. We also demonstrate that the nature of the gradient is crucial for range elasticity. Gradients which do not select for lower dispersal at the margin than in the core (especially gradients in patch size, demographic stochasticity and extinction rate) do not lead to elastic range behavior. Thus, we predict that range contractions are likely to occur after periods of expansion for species living in gradients of increasing patch isolation, which suffer from Allee effects.     

Population cycles produce periodic range boundary pulses

Classical theories of biological invasions predict constant rates of spread that can be estimated from measurable life history parameters, but such outcomes depend strongly on assumptions that are often unmet in nature. Subsequent advances have demonstrated how relaxing assumptions of these foundational models results in other spread patterns seen in nature, including invasions that accelerate through time, or that alternate among periods of expansion, retraction, and stasis of range boundaries. In this paper, we examine how periodic population fluctuations affect temporal patterns of range expansion by coupling empirical data on the gypsy moth invasion in North America with insights from a model incorporating population cycles, Allee effects, and stratified diffusion. In an analysis of field data, we found that gypsy moth spread exhibits pulses with a period of 6 yr, which field data and model simulations suggest is the result of a 6‐yr population cycle in established populations near the invasion front. Model simulations show that the development of periodic behavior in range expansion depends primarily on the period length of population cycles. The period length of invasion pulses corresponded to the population cycle length, and the regularity of invasion pulses tended to decline with increases in population cycle length. A key insight of this research is that dynamics of established populations, behind the invasion front, can have strong effects on spread. Our findings suggest that coordination between separate management programs targeting low‐density spreading and established outbreaking populations, respectively, could increase the efficacy of efforts to mitigate gypsy moth impacts. Given the variety of species experiencing population fluctuations, Allee effects, and stratified diffusion, insights from this study are potentially important to understanding how the range boundaries of many species change.     

Allee效应对具有生境恢复的集合种群的影响

Allee效应与种群的灭绝密切相关,其研究对生态保护和管理至关重要。Allee效应对物种续存是潜在的干扰因素,濒危物种更容易受其影响,可能会增加生存于生境破碎化斑块的濒危物种的死亡风险,因此研究Allee效应对种群的动态和续存的影响是必要的。从包含由生物有机体对环境的修复产生的Allee效应的集合种群模型出发,引入由其他机制形成的Allee效应,建立了常微分动力系统模型和基于网格模型的元胞自动机模型。通过理论分析和计算机模拟表明:(1)强Allee效应不利于具有生境恢复的集合种群的续存;(2)生境恢复有利于种群续存;(3)局部扩散影响了集合种群的空间结构、动态行为和稳定性,生境斑块之间的局部作用将会减缓或消除集合种群的Allee效应,有利于集合种群的续存。     

氮水平和竞争对互花米草与芦苇叶特征的影响

 互花米草 (Spartina alterniflora) 和芦苇 (Phragmites australis) 是滨海盐沼湿地的多年生草本植物,从世界范围来看,它们二者具有区域性的相互入侵特征,因此研究生境条件对两物种互侵机制的影响是一个十分有意义的生态学命题。该文运用随机区组实验设计方法,模拟海滩环境、构建人工种群、控制可变因子,研究了外来种互花米草与本地种芦苇分别单种和混种时,叶特征对不同氮水平、不同植株密度的响应。结果表明：随着氮水平的升高,互花米草和芦苇的叶面积无论是在单种还是混种情况下都显著增加 (p<0.05),但混种条件下芦苇的叶面积在高氮水平下增幅减少,这与高氮状况下互花米草与芦苇的竞争加剧有关;氮水平对单种中两种植物的叶数影响最显著 (p<0.01),对混种中互花米草的叶数和芦苇的叶宽影响最大 (p<0.05)。植株密度增加导致种内和种间竞争加剧,无论在单种还是混种处理下,都造成两种植物叶面积的显著减少 (p<0.05)。单种处理中, 两物种的叶数受密度的响应最显著 (p<0.05);而混种处理中芦苇对互花米草的竞争显著减小了互花米草的叶宽和叶数(p<0.05),互花米草对芦苇的竞争则显著减小了芦苇 的叶长、叶宽和叶数 (p<0.05)。两种植物的竞争结果受到氮营养的调控,低、高氮水平下互花米草的种间竞争能力大于芦苇,中氮水平下则是芦苇的种间竞争能力大于互花米草。高氮水平下互花米草通过叶面积的快速增加抑制了芦苇的叶生长,使其叶面积减少,从而在竞争中占据优势,这可能是互花米草入侵我国海滩芦苇种群的机制之一。