Reduced tolerance to simulated herbivory on clonal organs in alien genotypes: a multi-species experiment with native and introduced origins

Clonal growth may increase the likelihood for alien plants becoming invasive, as it is an efficient foraging and spatial exploration strategy. Here, we investigated the effect of artificial herbivory on organs of clonal growth and its potential to drive post-introduction evolutionary change. Based on the assumption that tolerance traits are costly and that clonal alien species may benefit from investing freed resources into growth, fecundity or nutrient acquisition, we tested the hypothesis of lower tolerance to herbivory on organs of clonal growth in alien plants. In a common-garden experiment we studied divergence in plants from native German and alien New Zealand populations of six species with different clonal growth forms. A nutrient treatment testing the plant’s acquisition abilities, was combined with artificial herbivory on clonal organs. We investigated origin-dependent differences in sexual reproduction, plant growth and the production of clonal organs. For aboveground and clonal organ biomass, alien plants showed lower tolerance to artificial herbivory on clonal organs than native plants. In the combined herbivory and nutrient treatment, alien plants of four species grew fewer clonal organs when compared to the nutrient treatment alone. Alien plants of the other two species produced more clonal organs, regardless of treatment. All species revealed significant differences in flower production between origins, with five of them producing more flowers on alien than on native plants. The results support the hypothesis that a release of herbivory on clonal organs has lead to subtle evolutionary changes in tolerance of alien plants and to a species-dependent increase in plant vigour, clonal growth and/or sexual reproduction that may enhance their invasive success.     

莲草直胸跳甲取食对空心莲子草和莲子草克隆整合的影响

很多外来入侵植物都具有克隆生长习性,探究克隆整合特性与外来克隆植物入侵性间的关系对阐明其生态适应性及入侵机制具有重要的意义。本研究以入侵植物空心莲子草及其本地同属种莲子草为研究对象,比较在生防昆虫莲草直胸跳甲的取食下,克隆整合对两种植物先端分株、基端分株及整个克隆片段生长和生物量分配的影响。结果表明: 在莲草直胸跳甲取食下,有克隆整合的空心莲子草先端分株的叶片数、茎长、分株数及整个克隆片段的地径均显著高于无克隆整合植株,其基端分株及整个克隆片段的地下生物量和总生物量相较于无克隆整合植株分别降低了78.2%、60.9%和48.7%、37.2%;有克隆整合的莲子草先端分株的地径及整个克隆片段的叶片数与无克隆整合植株相比显著增加,其基端分株数显著降低了21.7%,而其先端分株、基端分株及整个克隆片段的生物量均无显著差异。耗益分析表明,在莲草直胸跳甲取食下,空心莲子草先端分株的分株数与生物量及莲子草先端分株的分株数均能通过克隆整合显著受益,而两种植物基端的分株数、生物量的耗益则不受克隆整合处理的影响。这些结果表明,克隆整合虽能在一定程度缓解莲草直胸跳甲对于两种植物先端分株的取食压力,且空心莲子草的克隆整合作用要强于莲子草,但在整个克隆片段水平上,两种植物并不能通过克隆整合显著获益。     

Invasive alien plants in China: role of clonality and geographical origin

Biological invasions have become a significant threat to the global environment. Unfortunately, to date there is no consensus on invasion mechanisms and predictive models. Controversies range from whether we can reliably predict which species may become invasive to which species characteristics (e.g., life history, taxonomic groups, or geographic origin) contribute to the invasion processes. We examined 126 invasive alien plant species in China to understand the role of clonality and geographical origin in their invasion success. These species were categorized into three groups (I, II, III) based on their invasiveness in terms of current spatial occupation and the degree of damage to invaded habitats. Clonal plants consisted of almost half (44%) of the 126 invasive species studied, and consisted of 66% of 32 the most invasive alien plant species (Group I). There was a significant positive relationship between clonality and species invasiveness. A 68% of the 126 species studied originated in the continent of America (North and/or South America). These preliminary findings support that America is the primary geographical origin of invasive alien plant species in China and that clonality of the invasive plant species contributed significantly to the their invasiveness. The results suggest an urgent need at the global scale to investigate the mechanisms whereby plant clonal growth influences plant invasions, and the need for a focus at regional scale to examine factors affecting the exchange of invasive plant species between America and China.     

植物克隆整合生态效应及潜在应用

克隆植物因特殊的克隆整合和空间拓展特性,在异质生境下展示出较高的生态适应性及适合度,这是其广泛存在于各类生态系统的一个重要原因。目前对克隆整合在个体或种群水平的生态学效应已有较深认识,而对群落及生态系统的影响及作用机制则明显关注不足。前期研究表明,克隆整合对土壤理化性质、根际微生物及个体竞争力均有显著影响,从而有利于克隆植物的成功入侵、生境修复及植被重建等。对群落及生态系统水平的克隆整合生态学效应的研究进行归纳和总结,分析了克隆整合对植物群落结构和生产力、根际微生物和土壤动物、生态系统碳固持、养分循环等的影响;阐释了克隆整合及空间拓展特性对退化生态系统的修复及作用机制,并指出今后克隆整合的研究应同时考虑微观(根际过程)和宏观(群落及生态系统)层次的效应以及短期与长期的效应。可将克隆整合与植物-土壤反馈等其他生态过程相联系,综合探究克隆整合的生态学意义。     

外来入侵植物和本地植物核DNA C-值的比较及其与入侵性的关系

用流式细胞仪测定了8科10属13种外来入侵植物、6种本地植物和1种外来非入侵植物的核DNAC-值。结果表明:作为整体,外来入侵植物的平均核DNAC-值显著低于本地种和外来非入侵种,但对同属不同类型植物进行比较,未发现一致的规律;在4个既包含外来入侵种又包含本地种的属中,泽兰属(Eupatorium)和鬼针草属(Bidens)外来入侵种的核DNAC-值显著低于同属本地种,莲子草属(Alternanthera)的2种外来入侵植物中仅有1个种的核DNAC-值显著低于同属本地种,而草胡椒属(Peperomia)外来入侵种的核DNAC-值显著高于同属本地种;表明核DNAC-值与外来植物入侵性无必然联系。     

The cobblers stick to their lasts: pollinators prefer native over alien plant species in a multi-species experiment

The majority of plant species rely, at least partly, on animals for pollination. Our knowledge on whether pollinator visitation differs between native and alien plant species, and between invasive and non-invasive alien species is still limited. Additionally, because numerous invasive plant species are escapees from horticulture, the transition from human-assisted occurrence in urbanized habitats to unassisted persistence and spread in (semi-)natural habitats requires study. To address whether pollinator visitation differs between native, invasive alien and non-invasive alien species, we did pollinator observations for a total of 17 plant species representing five plant families. To test whether pollinator visitation to the three groups of species during the initial stage of invasion depends on habitat type, we did the study in three urbanized habitats and three semi-natural grasslands, using single potted plants. Native plants had more but smaller flower units than alien plants, and invasive alien plants had more but smaller flowers than non-invasive alien plants. After accounting for these differences in floral display, pollinator visitation was higher for native than for alien plant species, but did not differ between invasive and non-invasive alien plant species. Pollinator visitation was on average higher in semi-natural than in urbanized habitats, irrespective of origin or status of the plant species. This might suggest that once an alien species has managed to escape from urbanized into more natural habitats, pollinator limitation will not be a major barrier to establishment and invasion.     

The biogeography of invasive alien plants in California: an application of GIS and spatial regression analysis

The spatial distribution of invasive alien plants has been poorly documented in California. However, with the increased availability of GIS software and spatially explicit data, the distribution of invasive alien plants can be explored. Using bioregions as defined in Hickman (1993 ), I compared the distribution of invasive alien plants (n = 78) and noninvasive alien plants (n = 1097). The distribution of both categories of alien plants was similar with the exception of a higher concentration of invasive alien plants in the North Coast bioregion. Spatial autocorrelation analysis using Moran's I indicated significant spatial dependence for both invasive and noninvasive alien plant species. I used both ordinary least squares (OLS) and spatial autoregressive (SAR) models to assess the relationship between alien plant species distribution and native plant species richness, road density, population density, elevation, area of sample unit, and precipitation. The OLS model for invasive alien plants included two significant effects; native plant species richness and elevation. The SAR model for invasive alien plants included three significant effects; elevation, road density, and native plant species richness. The SAR model for noninvasive alien plants resulted in the same significant effects as invasive alien plants. Both invasive and noninvasive alien plants are found in regions with low elevation, high road density, and high native‐plant species richness. This is in congruity with previous spatial pattern studies of alien plant species. However, the similarity in effects for both categories of alien plants alludes to the importance of autecological attributes, such as pollination system, dispersal system and differing responses to disturbance in the distribution of invasive plant species. In addition, this study emphasizes the critical importance of testing for spatial autocorrelation in spatial pattern studies and using SAR models when appropriate.     

The invasive plant Alternanthera philoxeroides benefits from clonal integration in response to defoliation

Many notorious alien invasive plants have the capacity for vigorous clonal growth, and clonal integration may contribute to their invasiveness in response to various disturbances. Here, it is hypothesized that clonal integration affects the growth, biomass allocation, physiology, and compensatory response of the alien invasive clonal plant Alternanthera philoxeroides when faced with defoliation. To test these hypotheses, a growth experiment was conducted to investigate the effect of clonal integration on the responses of A. philoxeroides to different levels of defoliation. Daughter ramets that had been grown with stolon connections that were either severed from or connected to the mother plant were subjected to four defoliation levels: 0 (control), 30% (mild), 60% (moderate) and 90% (heavy) removal of leaf tissue. Defoliation greatly decreased growth (total biomass, number of ramets and total stolon length) but increased the maximum quantum yield of photosystem II (F_v/F_m) of daughter ramets. Clonal integration significantly increased growth, F_v/F_m and contents of non-structural carbohydrates (soluble sugars and total non-structural carbohydrates) of A. philoxeroides, and these effects were larger under heavier defoliation. Moreover, clonal integration markedly reduced the shoot/root ratio of A. philoxeroides, and these effects tended to increase with increasing levels of defoliation. These results support our hypothesis that A. philoxeroides benefits from clonal integration in response to defoliation, suggesting that clonal integration may be closely related to the invasiveness of A. philoxeroides in natural habitats with frequent disturbances.     

异质光照环境下克隆整合对南美蟛蜞菊和蟛蜞菊种间关系的影响

很多入侵植物具有克隆性,克隆整合对入侵克隆植物生长和繁殖具有重要的贡献。自然界中,植物生长和繁殖所需的各种资源如光照、水分和矿质养分等在空间上分布通常是异质的,但关于异质环境下克隆整合对入侵植物和本土同属植物种间关系影响的研究相对缺乏。通过温室控制实验,将入侵植物南美蟛蜞菊(Wedelia trilobata)和同属本土植物蟛蜞菊(W.chinensis)的分株对单独种植或者混合种植在异质性光照条件下,同时通过保持或者切断分株之间的连接来控制克隆整合效应的有无,研究异质光照环境下克隆整合对南美蟛蜞菊和蟛蜞菊种间关系的影响。克隆整合对南美蟛蜞菊和蟛蜞菊的生长和繁殖都有促进作用,且南美蟛蜞菊比蟛蜞菊从克隆整合中获益更多。与单独种植相比,两者混种对南美蟛蜞菊的叶生物量有显著影响,而对本地种蟛蜞菊的根生物量有显著影响。克隆整合和种间关系对南美蟛蜞菊的总生物量和叶生物量产生了显著的交互作用,而对蟛蜞菊各指标无显著影响。克隆整合状态显著影响了南美蟛蜞菊和蟛蜞菊的种间关系。这些结果表明,异质环境下克隆整合可以改变入侵植物南美蟛蜞菊和本土植物蟛蜞菊的生长性状及种间关系。     

三峡库区消落带外来植物入侵与景观基质组成结构的关联性

外来物种入侵过程不仅受入侵地生境的影响,也受到周边景观基质的影响.厘清景观基质对外来入侵物种的影响,对于外来物种的入侵管控具有重要意义.本研究针对长江三峡库区消落带植物群落开展调查,评估三峡库区消落带外来植物入侵现状;在岸线以上(含消落带)2000 m范围内划分出10个景观格局尺度,采用14个指标分析景观基质组成结构;...     

Little evidence for release from herbivores as a driver of plant invasiveness from a multi‐species herbivore‐removal experiment

Enemy release is frequently posed as a main driver of invasiveness of alien species. However, an experimental multi‐species test examining performance and herbivory of invasive alien, non‐invasive alien and native plant species in the presence and absence of natural enemies is lacking. In a common garden experiment in Switzerland, we manipulated exposure of seven alien invasive, eight alien non‐invasive and fourteen native species from six taxonomic groups to natural enemies (invertebrate herbivores), by applying a pesticide treatment under two different nutrient levels. We assessed biomass production, herbivore damage and the major herbivore taxa on plants. Across all species, plants gained significantly greater biomass under pesticide treatment. However, invasive, non‐invasive and native species did not differ in their biomass response to pesticide treatment at either nutrient level. The proportion of leaves damaged on invasive species was significantly lower compared to native species, but not when compared to non‐invasive species. However, the difference was lost when plant size was accounted for. There were no differences between invasive, non‐invasive and native species in herbivore abundance. Our study offers little support for invertebrate herbivore release as a driver of plant invasiveness, but suggests that future enemy release studies should account for differences in plant size among species.     

Differences in physiological integration between invasive and noninvasive introduced clonal species of Carpobrotus

Aims Clonal growth is associated with invasiveness in introduced plant species, but few studies have compared invasive and noninvasive introduced clonal species to investigate which clonal traits may underlie invasiveness. To test the hypothesis that greater capacity to increase clonal growthviaphysiological integration of connected ramets increases invasiveness in clonal plants, we compared the effects of severing connections on accumulation of mass in the two species of the creeping, succulent, perennial, herbaceous genusCarpobrotusthat have been introduced on sand dunes along the Pacific Coast of northern California, the highly invasive speciesCarpobrotus edulisand the co-occurring, noninvasive speciesCarpobrotus chilensis.     

Effects of clonal integration in the expansion of two alien <Emphasis Type="Italic">Carpobrotus</Emphasis> species into a coastal dune system – a field experiment

What makes a plant a successful invader is one of the most interesting questions in modern ecology. Comparative studies including congeners differing in invasiveness are a straightforward approach to detect potential traits explaining invasions. In this experiment we studied the importance of clonal integration and the capacity to buffer fragmentation in the expansion of two stoloniferous invaders, Carpobrotus edulis, considered more invasive, and Carpobrotus acinaciformis, considered less invasive. In particular we aim to determine whether differences in these clonal traits may explain differences in invasiveness between both species. We report evidence that clonal integration favour the expansion of the two exotic clonal species into a sand dune system. Benefit derived from clonal integration by itself does not explain differences in invasiveness between theses two exotic species. However, our result indicates that the greater invasiveness of C. edulis could be explained by a higher capacity to buffer the negative effect of fragmentation in comparison with C. acinaciformis. To elucidate the real contribution of clonal traits in plant invasions, new comparative studies should be conducted including more clonal species.     

中国外来入侵生物的空间分布格局及其影响因素

不同地理区域影响生物分布格局的因子不同,对外来入侵物种也是如此。在区域尺度上分析外来入侵生物的空间分布格局及其影响因子对预测生物入侵的影响及入侵种的控制管理具有重要意义。本研究应用中国外来入侵物种数据库、自然环境数据库和社会人文环境数据库,分析了我国外来入侵动植物的空间格局;并运用主成分分析（PCA）和典范对应分析（CCA）探讨了自然环境和人类活动等因素对外来入侵动植物分布格局的影响;同时研究了外来入侵物种多样性与本地物种多样性之间的关系。结果表明,我国现有外来入侵动物138种、入侵植物384种,其数量和密度都呈现出由东南沿海向西北内陆减少的趋势,且入侵动物和入侵植物空间格局基本一致;降水（MAP）是决定我国外来入侵动植物分布格局的主要自然环境因子,国民生产总值（GDP）是主要社会经济影响因子。在全国尺度上,外来入侵物种多样性与本地物种多样性之间呈显著的正相关关系,但地域间存在较大差异。不同区域外来入侵物种与本土物种多样性的相关关系表现出不同,与研究尺度有密切关系。     

南四湖湿地外来入侵植物

南四湖是我国重要的淡水湖泊湿地之一。为阐明南四湖湿地外来植物的现状及入侵规律, 为湿地生态系统的重建、生物多样性的保护和湿地资源开发等提供指导, 通过野外调查和参考有关文献资料, 对南四湖湿地外来植物进行了较为深入的研究, 并与青岛、昆嵛山、滨州沿海三地作了比较。结果表明, 南四湖湿地外来入侵植物有41种, 隶属19科35属; 以菊科和豆科为优势科; 外来入侵物种中热带来源与温带来源比例相当, 人为引入导致的外来入侵植物高达39%。南四湖与沿海三地相比, 沿海三地外来入侵植物的种类均比南四湖湿地多, 其中以经济最发达的青岛种类最多; 青岛外来入侵植物中有35种在南四湖湿地未发现; 南四湖湿地的特有外来入侵植物有5种, 体现出湿地外来入侵植物水生特性。群落相似性分析表明, Jaccard指数和Sorenson指数呈现的规律一致, 南四湖与滨州的相似性最高, 其次为昆嵛山, 与青岛的差异最大。通过分析南四湖湿地外来入侵植物种类和分布规律, 提出了相应的防治对策。     

From the backyard to the backcountry: how ecological and biological traits explain the escape of garden plants into Mediterranean old fields

To explain current ornamental plant invasions, or predict future ones, it is necessary to determine which factors increase the probability of an alien species becoming invasive. Here, we focused on the early phases of ornamental plant invasion in order to identify which plant features and cultivation practices may favor the escape of ornamental plants from domestic gardens to abandoned agricultural land sites in the Mediterranean Region. We used an original approach which consisted in visiting 120 private gardens in an urbanizing rural area of the French Mediterranean backcountry, and then visited surrounding old fields to determine which planted species had escaped out of the gardens. We built a database of 407 perennial ornamental alien species (most of which were animal-dispersed), and determined nineteen features that depicted the strength of species’ propagule pressure within gardens, the match between species requirements and local physical environment, and each species’ reproductive characteristics. Using standard and phylogenetic logistic regression, we found that ornamental alien plants were more likely to have escaped if they were planted in gardens’ margins, if they had a preference for dry soil, were tolerant to high-pH or pH-indifferent, and if they showed a capacity for clonal growth. Focusing only on animal-dispersed plants, we found that alien plants were more likely to have escaped if they were abundant in gardens and showed preference for dry soil. This suggests that gardening practices have a primary impact on the probability of a species to escape from cultivation, along with species pre-adaptation to local soil conditions, and capacity of asexual reproduction. Our results may have important implications for the implementation of management practices and awareness campaigns in order to limit ornamental plants to becoming invasive species in Mediterranean landscapes.     

叶物候、构建消耗和偿还时间对入侵植物碳积累的影响

随着全球经济一体化进程的深入,生物入侵已成为严重的全球性问题之一.外来种入侵打破了物种生存的自然平衡,导致生态系统趋于均质化,并给社会经济发展和人类健康造成了无法估量的损失.然而,目前人们对外来种的入侵机理仍不十分清楚.叶物候如叶片发生时间、凋落时间、及由二者决定的叶寿命和叶片发育进程是植物在季节和非季节性变化的环境中为了获得碳收益所采取的适应策略.叶构建消耗是植物碳收获过程中必要的成本投入,反映了植物在叶片水平上的能量分配策略.偿还时间能在一定程度上反映叶片碳的积累情况.从叶物候、构建消耗和偿还时间入手,分析了入侵植物的资源捕获能力和成本对其碳积累的影响,并提出了今后的研究方向.     

北京市建成区绿地植物物种来源分析

为探讨北京市建成区绿地植物的物种构成、外来植物组成及其在不同功能区的分布,采用机械布点和随机抽样法对北京市建成区绿地植物资源进行系统调查。结果表明：（1）北京市建成区绿地植物外来引进种173种,外来归化入侵种20余种：（2）外来引进种以国内引进为主,境外起源物种中美洲和欧亚区系植物占绝对优势：（3）外来种丰度按大小排列为：大型公园〉高等院校〉居住区绿地〉街头绿地／街心花园〉附属绿地〉街道绿地〉片林绿地,而各绿地类型的入侵种数量没有显著差异。     

中国外来入侵植物的分布格局及其与环境因子和人类活动的关系

认识区域尺度上外来入侵植物的分布格局及其成因对预测入侵的影响和入侵种的管理具有重要意义。该文采用聚类分析和排序的方法分析了我国外来入侵植物的空间格局,并利用多元线性逐步回归和典范对应分析探讨了自然环境因子和人类活动强度对中国32个省级空间单位(省市自治区,不包括香港和澳门)中外来入侵植物分布的影响。结果表明,中国各省外来入侵植物物种数从南到北逐渐减少,导致这一格局的主要因子为无霜期;各省外来入侵植物物种密度由东南海岸向内陆递减,造成这一趋势的主要影响因素为交通密度;纬度是解释中国各省外来入侵植物物种组成变异的主要因子,因此中国32个省区可归为低、中、高纬度区3大类型。在此基础上预测中国东南部地区有遭受更多外来植物入侵的可能;此外,交通发达的区域也将成为外来植物入侵的热点区,应该引起有关部门的重视。     

安徽省外来入侵植物的分布格局及其等级划分

区域性外来入侵植物的研究及其入侵等级划分在地方入侵植物管理与防治对策的制定上具有重要的指导意义。本文基于文献报道、标本记录和必要的分类学考证,特别是通过2013-2015年对安徽省外来入侵植物的全面调查,统计分析了安徽省外来入侵植物的物种组成、生活型、原产地、区系组成和水平分布格局,并对其入侵危害进行等级划分。结果显示,安徽省有外来入侵植物37科86属132种,总体上呈现种类丰富、原产于美洲的种类多、泛热带起源的种类多、危害严重的种类多、区域分化明显、入侵途径集中等特征。提出在外来入侵植物的防控中应重点警惕起源于美洲的、泛热带分布属的植物和关注人类活动频繁的热点区域。还对中国外来入侵植物研究中存在的问题进行了探讨,澄清了几个长期以来被忽视的关于外来入侵物种认知的问题。