上海崇明东滩互花米草种子产量及其萌发对温度的响应

外来物种互花米草通过快速的无性和有性繁殖,已在崇明东滩形成了大面积的单优势种群。研究了崇明东滩互花米草入侵区种子产量及温度对其萌发的影响。结果表明,互花米草在潮间带的中潮间带种子产量最高,为44523粒/m²,高潮间带次之,为31150粒/m²,低潮间带最低,为17250粒/m²。各潮间带的种子在10 30℃范围内均能萌发,而且随着温度的升高,萌发率和萌发速度总体呈现先增大后减小的趋势。互花米草种子在25℃恒温与20/30℃变温处理下的萌发率最高和萌发速度最快,而且互花米草种子在变温环境下的萌发率高于恒温处理。研究表明,崇明东滩中潮间带互花米草的种子产量和活性最高,是提供有性扩散种源的主要区域,适合种子萌发温度的4-5月是互花米草进行有性扩散的主要季节。研究结果对了解互花米草有性繁殖扩散的时空格局和入侵生态学过程,以及有效控制和管理入侵物种互花米草提供了科学依据。     

基质及水位对入侵种互花米草生长的影响

为了揭示基质和水位对互花米草种群扩张过程的影响,通过室内模拟试验分析了不同基质和地下水位对互花米草幼苗存活率、高生长和生物量及分配等表型参数的影响。结果表明:基质、水位及其交互作用对互花米草幼苗生长影响显著;基质类型和地下水位的交互作用显著影响幼苗的存活率;同种基质中地下水位的降低显著促进了幼苗根长的增加;由于不同基质持水力和营养的差异,沙中生长的幼苗随地下水位的降低地上部分表现出显著的生长抑制,而沙土和壤土中的幼苗在中等水位生长最好;沙中幼苗根状茎的长度显著低于沙土和壤土,而随水位的降低,沙土和壤土根茎长度显著下降。由此可以推论,互花米草种群扩张强度因生境而异,在沙土和壤土基质中互花米草可能表现出更强的入侵性和扩散能力。     

不同纬度地区互花米草生长性状及适应性研究

互花米草是我国沿海潮间带分布面积最广的入侵植物, 研究不同纬度地区间互花米草的生长特征和适应对策有助于揭示该植物对大范围异质生境的入侵适应机制。对我国沿海10 个省市26 个地区互花米草生长情况的调查发现:1)不同地区间互花米草的株高、节长、基茎、单株地上生物量、种群密度等性状特征均存在显著差异(P<0.01), 其变化规律与纬度密切相关: 随着纬度的升高, 互花米草的株高、节长、基茎、单株地上生物量、种群地上生物量总体呈现先升后降的规律, 而种群密度呈现为先降后升的规律。2)边缘区的互花米草种群密度优势明显: 植物个体矮小细密,种群密度最高达到2668 株·m–2, 是分布区中部的20 多倍, 较高的种群密度大幅度增加了个体或分株数量, 提高了生存概率, 降低了因种群面积过小带来的种群灭绝风险; 分布区中部的互花米草植株个体优势明显: 植株种群密度较低, 但植株个体高大粗壮, 单株地上生物量最高达到48.81 g, 是分布区边缘的30 倍; 表明中心区和边缘区的互花米草种群分别表现出“表现最大化”和“种群维持”的适应对策。3)温度是影响不同地区间互花米草生长差异的主要因素, 降水次之, 而pH 和盐度影响较小。     

氮、硫互作对克隆植物互花米草繁殖和生物量累积与分配的影响

氮、硫均是影响入侵植物互花米草（Spartina alterniflora Loisel.）生长与繁殖的重要营养元素,研究其交互作用可能有助于揭示互花米草成功入侵机制。本文在人工控制条件下,以河沙为栽培基质,研究了氮、硫互作对互花米草繁殖和生物量积累与分配的影响。实验结果表明：中氮和高氮处理下的互花米草分支强度、初级分株数、次级分株数、根状茎数及其总长要显著高于低氮,不同硫水平及氮硫互作对互花米草无性繁殖没有显著影响;互花米草的种子数和种子生物量受硫以及氮硫交互作用的影响显著,在高氮水平下,中硫处理的互花米草种子数和种子生物量要远远高于其他处理。中氮和高氮处理下的互花米草叶、茎、根、根状茎和总生物量显著高于低氮处理;只有在高氮水平下,高硫处理的生物量才显著低于其他处理。在中氮和高氮营养条件下,高硫处理会促使互花米草对地上部分投入更多的生物量。中硫-高氮作用对互花米草有性繁殖的促进,使互花米草能在硫浓度较高的滩地保持高有性繁殖率;虽然过高的硫会抑制互花米草的有性繁殖,但却能促进根状茎数目的增加,从而加速其向外短距离扩张;这些特性均有利于互花米草对滨海盐沼的入侵。     

外来种互花米草入侵模式与爆发机制

互花米草（Spartina alterniflora Loisel）因其促淤造陆和消浪护堤作用显著而被许多国家引种,如今却在侵入地快速蔓延并呈现爆发趋势,对生态系统造成了极大危害,被认为是研究生物入侵生态学和遗传学的模式植物。从种群的入侵力、生态系统可入侵性和入侵通道3个方面探讨互花米草的爆发机制,研究结果表明高遗传分化和基因渗入能力是互花米草爆发的遗传基础,对逆境的高抗性和强竞争力是其快速扩张的保障,而高繁殖系数是互花米草爆发的源泉。我国互花米草种群的早期扩散人为影响超过了自然过程,快速扩张呈现出点源扩散和多点爆发的特点,从而为其种群控制带来困难,同时种子的跳跃式和连续式扩散在互花米草种群维持、更新和爆发中有重要作用,强有力的克隆生长能力也为互花米草种群的连续扩张提供了保障。现阶段要完全控制和根除互花米草是不实际的,但在及时预测预警的基础上,应用成本一效益分析方法,采取有序控制和综合开发利用的策略,仍可望妥善解决互花米草入侵所带来的负面效应。     

除草剂对黄河三角洲入侵植物互花米草的影响

互花米草(Spartina alterniflora)作为我国危害最严重的外来入侵植物之一,严重威胁滨海湿地生态系统安全。筛选可高效灭除互花米草的除草剂,为互花米草防治提供技术支持。2017年7月在黄河三角洲潮间带对互花米草茎叶喷施不同除草剂,在施药当年和次年对互花米草生长状况和大型底栖动物密度进行跟踪调查。研究结果表明:1)高效氟吡甲禾灵对互花米草的灭除效果最好,既能杀死互花米草地上部分从而完全抑制有性繁殖,又可完全抑制次年的无性繁殖;2)氰氟草酯对互花米草的灭除效果也比较好,可以完全抑制互花米草的生长和结穗,但抑制根状茎无性繁殖能力的效果稍差,草甘膦可以完全抑制互花米草的有性繁殖,但无法抑制次年互花米草的无性繁殖;3)施用除草剂在短期内会毒害某些底栖动物,但在1年后底栖动物种群数量与对照处理无显著差异。除草剂对环境的影响程度与除草剂用量和施用时间息息相关,未来研究中,应在保证灭草效果的前提下,探索最佳用药时间和最低用量以最大限度地降低环境影响。     

基于CA模型的上海九段沙互花米草和芦苇种群扩散动态

基于1997年以来上海九段沙栽种芦苇和互花米草种群扩散格局的多年现场调查与遥感解译分析,并结合3S技术,构建了适合滩涂盐沼植物种群动态的元胞自动机（CA）模型．结果表明：该模型能较好地模拟九段沙上芦苇和互花米草种群扩散的格局和趋势,并验证了互花米草和芦苇的空间抢先占有模型以及锋面状连续扩散格局;土著植物芦苇与外来植物互花米草占据相同的生态位,而互花米草的种群扩散速度是芦苇的3—5倍,随着九段沙的不断淤涨,互花米草种群的快速扩展还将持续．构建的CA模型有助于深入研究外来物种扩散格局与其生态学过程之间的相互关系,对湿地生物多样性保护和资源管理具有重要意义．     

崇明东滩湿地芦苇和互花米草种群的分布格局及其与生境的相关性

应用GIS技术与实地调查相结合的方法,对上海崇明东滩湿地芦苇〔Phragmites australis(Cav.)Trin.exSteud.〕和互花米草(Spartina alterniflora Loisel.)种群斑块的分布格局和沿潮位梯度的分布特征及其与部分环境参数的相关性进行了研究。结果显示:在1 500 m×100 m尺度内共有42个斑块;其中,互花米草种群斑块数量较少(13个)但面积和周长均较大,呈集中化分布特征;芦苇种群斑块数量最多(25个)但面积和周长均较小,呈现破碎化特征;芦苇种群的斑块密度和边缘密度均大于互花米草种群,但二者的聚集度指数及连通度指数均较高。沿潮位降低,芦苇种群数量减少,互花米草种群数量增多;中、低潮位以互花米草种群为主,中、高潮位以芦苇种群为主。不同潮位二者活体与立枯体株数有明显差异且具有不同程度的相关性,其中,互花米草总株数与芦苇活体株数、互花米草立枯体株数与芦苇立枯体株数均呈显著负相关(P<0.05)。该区域内各景观和斑块类型格局指数数值均较高,Shannon多样性和均匀性指数均较大。不同潮位的环境参数差异明显且与距堤坝的距离有不同程度的相关性;中潮位相对高程最高(9.2 m),低潮位土壤盐度最高(37.17 ng.L-1);随潮位降低土壤总氮和总磷含量总体上下降,但中高潮位土壤总磷含量最高、高潮位土壤总氮含量最高。芦苇活体株数与距堤坝的距离及土壤盐度、互花米草活体株数与相对高程及土壤总磷含量、互花米草总株数与相对高程均呈显著负相关(P<0.05)。研究结果揭示:该区域景观已出现破碎化现象;互花米草种群规模逐年增大,与芦苇种群有明显的竞争,二者呈明显的镶嵌分布格局。高潮位的相对高程低而土壤盐度高,这种微生境有利于互花米草生长却能抑制芦苇生长,对二者的扩散和分布格局产生复杂的局部性影响。     

Geographical variation in germination traits of the salt-marsh cordgrass Spartina alterniflora in its invasive and native ranges

入侵地和原产地盐沼植物互花米草种子萌发性状的地理变异 种子萌发是植物早期生活史中最重要的阶段,决定了植物的生态位和地理分布范围,对外来植物的入侵潜力有重要影响。盐沼植物互花米草(Spartina alterniflora)在中国沿海滩涂的入侵范围最大,并已入侵到比原产地更低的纬度范围,这为我们研究互花米草在不同地理区域之间以及沿纬度梯度的萌发性状差异和适应提供了契机。在控温培养箱中淡水培养条件下,我们比较研究了来自入侵地(19°–40° N)10 个地点和原产地(27°–43° N)16个地点不同纬度互花米草种群的种子萌发性状差异,以及这种差异与各种 群来源地潮差和气候因素的相关性。原产地互花米草种群种子的萌发率和萌发指数比入侵地种群分别高10%和20%,但入侵地互花米草种群的萌发速度比原产地快3 d。入侵地互花米草种群的萌发率和萌发 指数随着纬度升高呈现线性递增的变化趋势,而原产地呈现线性递减的变化趋势。入侵地和原产地互花米草种群的平均萌发时间都与纬度呈现线性负相关。入侵地互花米草种群的萌发率和萌发指数与年日均温、年日最低均温、和年日最高均温呈现负相关,而在原产地呈现相反的相关关系。入侵地和原产地互花米草种群的平均萌发时间分别与年日均温、年日最低均温和年日最高均温呈现正相关关系。我们的研究结果表明,入侵地和原产地互花米草种群的萌发率和萌发指数已沿纬度进化出不同的渐变群格局,但平均萌发时间进化出与原产地一致的纬度渐变群格局,即在生物入侵过程中沿纬度梯度种子萌发策略会随着入侵时间和过程而发生变化。     

文冠果种群繁殖方式及其在种群更新中的作用

以黄土高原丘陵区文冠果天然种群为研究对象,通过样地调查,分析了不同生境文冠果种群的有性和无性繁殖方式及其在种群更新中的地位.结果表明：不同坡向文冠果种群均能进行有性繁殖和无性繁殖,在阳坡、半阴坡生境,文冠果以有性繁殖为主,实生苗数量和占据空间的能力均大于萌生苗,有性繁殖在这2个生境中对种群的更新贡献较大;在半阳坡生境,文冠果以无性繁殖为主,萌生苗数量和占据空间的能力均大于实生苗,无性繁殖对种群更新的贡献较大.在相同生境条件下,文冠果幼苗生长状况均表现为萌生苗＞实生苗.同一起源的文冠果幼苗生长状况均表现为阳坡＞半阳坡＞半阴坡.2种繁殖的瓶颈期在实生苗的种子-幼苗阶段和萌生苗的幼树-成株阶段.黄土丘陵区文冠果种群在新生境定居下来的过程中,首先以实生苗入侵,当水分、光照、养分条件较好时,文冠果又以无性繁殖方式迅速占领空间,繁衍后代.     

The relative importance of sexual and asexual reproduction in the spread of Spartina alterniflora using a spatially explicit individual-based model

This study investigates a spatially explicit, individual-based model for simulating the spread of invasive smooth cordgrass (Spartina alterniflora) in Yancheng coastal wetlands from 1995 to 2010. The model, which considers the landscape heterogeneity and changes detected by remote sensing, also reveals the relative importance of sexual and asexual reproduction in the spread by global sensitivity analysis. The model was verified as suitable for simulating the range expansion of S. alterniflora. The results show that: (1) although seedling recruitment is low, it significantly contributes to the range expansion of S. alterniflora. Removing sexual propagation greatly reduces the expansion rate. Rapid expansion requires both sexual and asexual reproduction; (2) in the global sensitivity analysis, the most significant affecters of S. alterniflora invasion were seed dispersal distance, adult survival rate and asexual recruitment survival rate. Sexual propagation contributes much more significantly to quick range expansion than asexual reproduction, but asexual reproduction is the main source of recruitment. Invasion control strategies should target a single reproduction mode. Here, limiting the germination and dispersal of seeds is suggested as a realistic strategy for controlling and managing invasion by this species.     

Optimum reproduction and dispersal strategies of a clonal plant in a metapopulation: a simulation study with Hieracium pilosella

Clonal spread is favoured in many plants at the expense of seed production in order to expand rapidly into open habitats or to occupy space by forming dense patches. However, for the dynamics of a population in a patchy landscape seed dispersal remains important even for clonal plants. We used a spatially explicit individual-based metapopulation model to examine the consequences of two trade-offs in Hieracium pilosella L: first, between vegetative and sexual reproduction, and second, between short and far-distance dispersal of seeds. Our main question was, what are the environmental conditions that cause a mixed strategy of vegetative and sexual reproduction to be optimal. The model was parameterised with field data on local population dynamics of H. pilosella. Patch dynamics were given firstly by disturbance events that opened patches in a matrix of a clonal grass that were colonisable for H. pilosella, and secondly by the gradual disappearance of H. pilosella patches due to the expanding grass. Simulations revealed opposing selection pressures on traits determined by the two trade-offs. Vegetative reproduction is favoured by local dynamics, i.e. the need for maintenance and expansion of established populations, whereas seed production is favoured by the necessity to colonise empty habitats. Similar pressures act on the proportion of seeds dispersed over short and far distances. Optimum reproductive and dispersal strategies depended on habitat quality (determined by seedling establishment probability), the fraction of dispersed seeds, and the fraction of seeds lost on unsuitable ground. Under habitat conditions supporting moderate to low seedling establishment, between 20% and 40% of reproductive effort in H. pilosella should be devoted to sexual reproduction with at least 10% of the seeds dispersed over distances suitable to attain empty patches. We conclude that in a spatially heterogeneous landscape sexual seed production in a clonal plant is advantageous even at the expense of local vegetative growth.     

Evolutionary increase in sexual and clonal reproductive capacity during biological invasion in an aquatic plant Butomus umbellatus (Butomaceae)

To test the hypothesis that increased allocation to reproduction is selected during biological invasion, we compared germination, survival, growth, and reproduction of native vs. introduced populations of the invasive aquatic plant Butomus umbellatus in a common greenhouse environment. Although seedling emergence and establishment did not differ consistently, survival thereafter was twice as high for eight introduced North American than eight native European populations. As predicted, introduced plants were more likely to produce sexual inflorescences and clonal asexual vegetative bulbils, and they invested much more biomass in both reproductive modes. Higher reproductive investment was due to higher proportional allocation of biomass rather than larger plant size. These results are consistent with selection for increased reproduction during range expansion. However, population genetic surveys indicate that recruitment from seed rarely occurs in introduced populations. Hence increased sexual allocation is not an adaptive response to invasion. Although increased clonal reproduction may be advantageous in expanding populations, genetic evidence from introduced populations of B. umbellatus suggests that increased clonal allocation may have arisen via stochastic processes during long-distance transport or a selective filter right at introduction, rather than incremental natural selection during range expansion.     

Mutation accumulation in space and the maintenance of sexual reproduction

The maintenance of sexual reproduction remains one of the major puzzles of evolutionary biology, since, all else being equal, an asexual mutant should have a twofold fitness advantage over the sexual wildtype. Most theories suggest that sex helps either to purge deleterious mutations, or to adapt to changing environments. Both mechanisms have their limitations if they act in isolation because they require either high genomic mutation rates or very virulent pathogens, and it is therefore often thought that they must act together to maintain sex. Typically, however, these theories have in common that they are not based on spatial processes. Here, we show that local dispersal and local competition can explain the maintenance of sexual reproduction as a means of purging deleterious mutations. Using a spatially explicit individual-based model, we find that even with reasonably low genomic mutation rates and large total population sizes, asexual clones cannot invade a sexual population. Our results demonstrate how spatial processes affect mutation accumulation such that it can fully erode the twofold benefit of asexuality faster than an asexual clone can take over a sexual population. Thus, the cost of sex is generally overestimated in models that ignore the effects of space on mutation accumulation.     

Rapid evolution of dispersal‐related traits during range expansion of an invasive vine Mikania micrantha

Rapid range expansion of invasive plants provides a unique opportunity to explore evolutionary changes of dispersal‐related traits during the invasion process. Increasing evidence now suggests that a higher dispersal rate is favored at the invasion front. However, little is known about the role of genetic differentiation and phenotypic plasticity on patterns of dispersal ability during the invasion process. In this study, we combined a field survey and a common garden transplant experiment to test for evidence of genetically based dispersal ability in Mikania micrantha, a highly invasive vine, across its invaded range in southern China. Three dispersal‐related traits, plume loading, seed mass and pappus radius, were measured in both natural and common garden populations. We found that in natural conditions, plume loading and seed mass significantly decreased with expanding distance from the source population, but in controlled conditions, these two traits exhibited a significant humped trend against percent field cover, indicating that dispersal ability of M. micrantha was selected for during range expansion and that the related traits were likely to be under genetic control. Furthermore, rebounding dispersal ability was detected in highly competitive sites in the range core, which suggested that this evolutionary process was likely partially driven by intraspecific competition. Because more and more plant species are under spatial nonequilibirum due to climate change, this study can serve to provide hints at the fate of spatially fluctuant populations.     

Reid's paradox revisited: the evolution of dispersal kernels during range expansion

Current approaches to modeling range advance assume that the distribution describing dispersal distances in the population (the "dispersal kernel") is a static entity. We argue here that dispersal kernels are in fact highly dynamic during periods of range advance because density effects and spatial assortment by dispersal ability ("spatial selection") drive the evolution of increased dispersal on the expanding front. Using a spatially explicit individual-based model, we demonstrate this effect under a wide variety of population growth rates and dispersal costs. We then test the possibility of an evolved shift in dispersal kernels by measuring dispersal rates in individual cane toads (Bufo marinus) from invasive populations in Australia (historically, toads advanced their range at 10 km/year, but now they achieve >55 km/year in the northern part of their range). Under a common-garden design, we found a steady increase in dispersal tendency with distance from the invasion origin. Dispersal kernels on the invading front were less kurtotic and less skewed than those from origin populations. Thus, toads have increased their rate of range expansion partly through increased dispersal on the expanding front. For accurate long-range forecasts of range advance, we need to take into account the potential for dispersal kernels to be evolutionarily dynamic.     

Multi-scale methods predict invasion speeds in variable landscapes

Spread rates of invasive plant species depend heavily on variable seed/seedling survivorships over various habitat types as well as on variability in seed dispersal induced by rapid transport of propagules in open areas and slow transport in vegetated areas. The ability to capture spatial variability in seed survivorship and dispersal is crucial to accurately predict the rate of spread of plants in real world landscapes. However, current analytic methods for predicting spread rates are not suited for arbitrary, spatially heterogeneous systems. Here, we analyze invasion rates of the invasive plant Phragmites australis (common reed) over variable wetland landscapes. Phragmites is one of the most pervasive perennial grasses, outcompeting native vegetation, providing poor wildlife habitat, and proving difficult to eradicate across its invasive range in North America. Phragmites spreads sexually via seeds and asexually via underground (rhizomes) and aboveground (stolons) stems. We construct a structured integrodifference equation model of the Phragmites life cycle capturing variable seed survivorship in a seed bank, sexual and asexual recruitment into a juvenile age class, and differential competition among all classes with adults. The demographic model is coupled with a homogenized ecological diffusion/settling seed dispersal model that allows for seed deposition that varies with habitat type. The dispersal kernel we develop does not require local normalization and can be implemented efficiently using standard computational techniques. The model generates a traveling wave of isolated patches, establishing only in suitable habitats. We use the method of multiple scales to predict invasion speed as a solvability condition at large scales and test the predictions numerically. Accurate predictions are generated for a wide range of landscape parameters, indicating that invasion speeds can be understood in landscapes of arbitrary structure using this approach.     

The Meuse river as a corridor for range expansion of the exotic plant species <Emphasis Type="Italic">Sisymbrium austriacum</Emphasis>: evidence for long-distance seed dispersal

Riparian habitats are particularly prone to invasion of non-indigenous plant species and several species have been shown to rapidly expand their range along river networks, possibly mediated by the occurrence of frequent long-distance seed dispersal events. However, there is still relatively little empirical evidence for long-distance seed dispersal along river networks and most studies to date are inconclusive with regards to the direction (upstream vs. downstream) of seed movement. Using assignment analyses based on dominant AFLP markers, we provide empirical evidence that downstream long-distance seed dispersal has facilitated range expansion of the exotic plant Sisymbrium austriacum along the Meuse River. Of 242 sampled individuals, 13 (5.4%) were allocated to a population other than the one from which it was sampled. Of these, nine (3.7%) individuals were assigned to a known population within the area, the furthest being more than 20 km away from the population from which it was sampled. All putative source populations were located upstream, thus providing strong evidence for downstream migration of propagules. These results support the general view that river systems may serve as efficient transport vectors of plant species and thus may play an important role in increasing the spatial spread and range expansion of exotic plant species.     

The evolution of conditional dispersal and reproductive isolation along environmental gradients

Dispersal modulates gene flow throughout a population's spatial range. Gene flow affects adaptation at local spatial scales, and consequently impacts the evolution of reproductive isolation. A recent theoretical investigation has demonstrated that local adaptation along an environmental gradient, facilitated by the evolution of limited dispersal, can lead to parapatric speciation even in the absence of assortative mating. This and other studies assumed unconditional dispersal, so individuals start dispersing without regard to local environmental conditions. However, many species disperse conditionally; their propensity to disperse is contingent upon environmental cues, such as the degree of local crowding or the availability of suitable mates. Here, we use an individual-based model in continuous space to investigate by numerical simulation the relationship between the evolution of threshold-based conditional dispersal and parapatric speciation driven by frequency-dependent competition along environmental gradients. We find that, as with unconditional dispersal, parapatric speciation occurs under a broad range of conditions when reproduction is asexual, and under a more restricted range of conditions when reproduction is sexual. In both the asexual and sexual cases, the evolution of conditional dispersal is strongly influenced by the slope of the environmental gradient: shallow environmental gradients result in low dispersal thresholds and high dispersal distances, while steep environmental gradients result in high dispersal thresholds and low dispersal distances. The latter, however, remain higher than under unconditional dispersal, thus undermining isolation by distance, and hindering speciation in sexual populations. Consequently, the speciation of sexual populations under conditional dispersal is triggered by a steeper gradient than under unconditional dispersal. Enhancing the disruptiveness of frequency-dependent selection, more box-shaped competition kernels dramatically lower the speciation-enabling slope of the environmental gradient.