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

AimsClonal 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.     

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。     

克隆整合和石油污染对芦苇湿地土壤理化性质的影响

克隆整合是克隆植物重要的性状之一。它不仅能够提高分株对环境胁迫的耐受能力,而且可能影响分株周围的土壤属性。为检验克隆整合对土壤属性的影响,在黄河三角洲芦苇湿地开展克隆整合和石油污染的两因子的野外实验。每年将0、5或10 mm厚的原油添加到直径为60 cm的圆形芦苇群落样方内来模拟无污染、轻度或重度石油污染,并通过保留或切断样方内外芦苇根状茎的连接来实现克隆整合的有或无。实验开始于2014年,并于2016年10月采集样方内土壤样品,测定土壤团聚体组成、pH值、电导率、总碳、总氮、总磷和有机碳含量。石油污染显著增加了土壤粗大团聚体(粒径:2 mm)、pH值、总氮和有机碳含量,降低了土壤微团聚体(粒径:0.053—0.25 mm)占比以及电导率。克隆整合显著降低了土壤pH值,提高了土壤电导率和氮磷比。克隆整合和石油污染的交互作用仅对电导率有显著效应。因此,石油污染和克隆整合都可以影响湿地土壤的理化性质,而克隆整合对分株周围土壤理化性质的影响可能进一步影响克隆植物的优势度。     

Effects of salinity and clonal integration on the amphibiousplant Paspalum paspaloides: growth,photosynthesis and tissue ion regulation

Aims Clonal integration can increase performance of clonal plants suffering from environmental stress, and clonal plants in many wetlands commonly face stress of flooding accompanied by salinity. However, few studies have tested roles of clonal integration in amphibious plants expanding from terrestrial to aquatic saline habitats.     

克隆植物对种间竞争的适应策略

克隆植物种群因其寿命的持久性、空间上的可移动性和繁殖方式的多样化等特征与非克隆植物有很大区别, 在自然生态系统中占有重要地位, 甚至成为优势种或者建群种。该文通过归纳有关克隆植物的种间竞争适应策略研究案例, 阐述了克隆植物的竞争能力差异和影响竞争力的因素; 论述了克隆植物在构件形态、克隆构型、繁殖对策等方面对种间竞争的响应, 以及生理整合作用与种间竞争的关系; 分析了导致某些同类研究的结论不一致的原因, 认为实验对象差异、实验设计、生境条件与克隆植物形态及生理上的时空动态变化等都可能影响实验结果; 提出了全球变化背景下的克隆植物种间竞争及其分子生态学机制等可能是今后需要重点关注的问题。     

Short‐term fitness benefits of physiological integration in the clonal herb Hydrocotyle peduncularis

Abstract We test whether physiological integration enhances the short‐term fitness of the clonal herb Hydrocotyle peduncularis (Apiaceae, R. Brown ex A. Richards) subjected to spatial variation in water availability. Our measures of fitness and costs and benefits are based on the relative growth rate of fragmented genets. Physiological integration over a gradient in soil moisture resulted in a highly significant net benefit to genet growth of 0.015 g g^?1 day^?1. This net benefit represents a significant enhancement of the average fitness of fragmented genets spanning the moisture gradient relative to the average of those growing in homogeneous moist or dry conditions. Sections of genet fragments growing in dry conditions in spatially heterogeneous treatments had significantly higher growth than the sections they were connected to that were growing in moist conditions. Within fragments, older (parent) sections growing in moist conditions experienced significant costs from connection to younger (offspring) sections growing in dry conditions. In contrast, offspring sections with ample water did not experience any costs when connected to parent sections growing in dry conditions. However, the net benefit of physiological integration was similar for parent and offspring sections, suggesting that parent and offspring sections contributed equally to the net benefit of physiological integration to genet growth and short‐term fitness.     

Two types of division of labour in clonal plants: benefits, costs and constraints

Clonal plants spread vegetatively within their habitats by forming rooted ramets on stolons or rhizomes. Each of these ramets is capable of an independent existence after establishment. Nevertheless, ramets remain physically connected by stolon or rhizome internodes for variable periods of time, thereby allowing for resource movement and signal transduction within clones.Interconnected ramets of clonal plants, though potentially independent and totipotent, can specialize functionally in the performance of limited numbers of tasks such as the uptake of resources from above- vs below-ground sources, carbohydrate storage, vegetative spread and sexual reproduction. Such specialization and cooperation is comparable to a division of labour in economic systems or in colonies of social animals. The ecological significance of division of labour in clonal plants may be found in the increased efficiency of entire clones in exploiting their environments.Two different types of division of labour in clonal plants will be discussed in this review. The first type is an environmentally-induced specialization of ramets in the uptake of locally abundant resources (plastic division of labour), which can be found in several stoloniferous species. Evidence exists that this response increases resource uptake in spatially heterogeneous environments. The second type of division of labour, which occurs mainly in rhizomatous species, relates to a developmentally-programmed specialization and cooperation between interconnected ramets. This response pattern is thought to enhance plant performance by restricting the number of tasks for individual ramets and thereby significantly increasing the efficiency of task performance. In some plants, such an inherent division of labour is likely to contribute to nutrient extraction from poor and unpredictably variable sources.In this article not only benefits but also potential costs and constraints on division of labour in clonal plants are shown. The aim is to provide a review of existing knowledge and to develop concepts and hypotheses for future research.     

芦苇克隆整合对石油污染湿地土壤微生物群落结构和生物量的影响

克隆植物形体相连的无性个体(分株)之间可以进行水分、养分和光合产物的传递和共享,并且这种克隆整合可以显著提高分株对环境胁迫的耐受能力,从而可能进一步影响分株周围的土壤微生物群落结构和生物量。尽管国内外已经开展了大量有关克隆整合对分株耐受胁迫能力影响的研究,但克隆整合对土壤微生物群落结构和生物量影响的研究却十分缺乏。以黄河三角洲芦苇(Phragmites australis)湿地生态系统为研究对象,将直径60 cm的圆形样方进行三个水平的石油添加处理(不添加石油或每年添加5 mm或10 mm厚的石油),同时通过切断或不切断样方内外芦苇根状茎的连接来控制克隆整合的有无。实验连续开展了两年(2014—2015年),每年8月份在样方内进行土壤样品取样,在实验室内采用磷酸脂肪酸(PLFA)法测定土壤微生物总量及主要微生物类群含量,并测定土壤微生物生物量碳和氮含量。取样时间显著影响土壤微生物PLFA总量、微生物碳和氮含量,这3个变量在2015年均显著高于2014年。石油添加在2015年显著增加了土壤微生物PLFA总量,但在2014年却无显著效应;同时,石油添加在2014年降低了土壤微生物碳和氮含量,而在2015年却增加了其含量。然而,无论在2014年还是2015年,芦苇的克隆整合对土壤微生物PLFA含量、微生物碳和氮含量均没有显著影响。土壤微生物PLFA总量与土壤微生物碳和氮含量呈正相关关系。这些结果表明,石油污染可以显著影响湿地土壤微生物动态,但克隆整合却无显著效应。     

Effects of clonal integration on allelopathy of invasive plant Wedelia trilobata under heterogeneous light conditions

异质光照条件下克隆整合对入侵植物南美蟛蜞菊化感作用的影响 植物入侵已成为全球生态系统最严重的威胁之一。当具有克隆生长能力的入侵植物入侵或定殖到新的生境时,它们相互连接的分株可能受到异质光照的影响。在异质光照条件下,克隆整合对入侵植物化感作用的影响尚不清楚。为研究异质光照条件下克隆整合对入侵植物南美蟛蜞菊(Wedelia trilobata) 化感作用的影响,采用两个连续分株的克隆片段进行了盆栽试验。较老的分株暴露在全光下,而年轻的分株则受到20%的全光照。同时,每个克隆片段的年轻分株与目标植株(一个番茄苗)在盆栽中相邻生长。南美蟛蜞菊的两个连续分株之间的匍匐茎设置切断和不切断两种处理。另外,两株番茄幼苗 (一株作为目标植株)在盆栽中相邻生长作为对照。研究结果表明,当与目标植物相邻生长的南美蟛蜞菊分株之间的匍匐茎保持完整时,目标植株的生物量积累、叶片叶绿素和氮含量、叶绿素荧光参数和净光合速率及其根长和活性相较于匍匐茎切断处理显著降低。异质光照条件下连续两个分株之间碳水化合物的运输或共享可以增强20%全光处理下年轻分株的化感作用。克隆整合在异质光照条件下对具有克隆生长能力的入侵植物的入侵或定殖具有重要意义。     

Effects of soil heterogeneity and clonal integration on Alternanthera philoxeroides populations with a radial ramet aggregation

Some clonal plants can spread their ramet populations radially, and soil heterogeneity and clonal integration may greatly affect the establishment of these types of populations. We constructed Alternanthera philoxeroides populations with a radial ramet aggregation, allowing old ramets of clonal fragments to concentrate in central pots and younger ramets to root in peripheral pots. The peripheral pots were supplemented either with three different levels (high, medium and low) of soil nutrients to simulate a heterogeneous soil environment, or only one medium level of soil nutrients to simulate a homogeneous environment. Stolon connections between the central older ramets and the peripheral younger ramets were left intact or severed to test the effect of clonal integration. The maintenance of stolon connection could induce the division of labor between different‐aged ramets, by increasing the root investment in central ramets and the above‐ground growth in peripheral ramets. The maintenance of stolon connection could improve the growth of the central and peripheral ramets, clonal fragments and even the whole population. However, the positive consequence in peripheral ramets and whole fragments was only detected in the high‐nutrient patch of heterogeneous treatment. In sum, in the population with the radial ramet aggregation, clonal integration can play a key role in the rapid recruitment of young ramets of A. philoxeroides fragments, as well as the expansion of the whole population. The magnitude of clonal integration also became more obvious in the peripheral young ramets and whole fragments that experienced high‐nutrient patches.     

Shifting effects of physiological integration on performance of a clonal plant during submergence and de-submergence

Background and Aims

Submergence and de-submergence are common phenomena encountered by riparian plants due to water level fluctuations, but little is known about the role of physiological integration in clonal plants (resource sharing between interconnected ramets) in their adaptation to such events. Using Alternanthera philoxeroides (alligator weed) as an example, this study tested the hypotheses that physiological integration will improve growth and photosynthetic capacity of submerged ramets during submergence and will promote their recovery following de-submergence.

Methods

Connected clones of A. philoxeroides, each consisting of two ramet systems and a stolon internode connecting them, were grown under control (both ramet systems untreated), half-submerged (one ramet system submerged and the other not submerged), fully submerged (both ramet systems submerged), half-shaded (one ramet system shaded and the other not shaded) and full-shaded (both ramet systems shaded) conditions for 30 d and then de-submerged/de-shaded for 20 d. The submerged plants were also shaded to very low light intensities, mimicking typical conditions in turbid floodwater.

Key Results

After 30 d of submergence, connections between submerged and non-submerged ramets significantly increased growth and carbohydrate accumulation of the submerged ramets, but decreased the growth of the non-submerged ramets. After 20 d of de-submergence, connections did not significantly affect the growth of either de-submerged or non-submerged ramets, but de-submerged ramets had high soluble sugar concentrations, suggesting high metabolic activities. The shift from significant effects of integration on both submerged and non-submerged ramets during the submergence period to little effect during the de-submergence period was due to the quick recovery of growth and photosynthesis. The effects of physiological integration were not found to be any stronger under submergence/de-submergence than under shading/de-shading.

Conclusions

The results indicate that it is not just the beneficial effects of physiological integration that are crucial to the survival of riparian clonal plants during periods of submergence, but also the ability to recover growth and photosynthesis rapidly after de-submergence, which thus allows them to spread.     

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。     

Resource transport between ramets alters soil resource pattern: a simulation study on clonal growth

Clonal plants spread horizontally, and can transport nutrients between ramets. Decaying biomass feeds back nutrients into the soil, but importantly, the place of deposition may differ from the place of uptake. To our knowledge, the present model is the first attempt to couple population dynamics with resource dynamics with the consideration of lateral transport. The simulations start from various initial resource patterns. Six types of clonal plants are compared, which differ in the birth and survival rates of ramets. Size of the ramet population and the amount of translocated resource are recorded over time. In addition, we consider the pattern of gaps in the canopy of the clonal plant from the aspect of two colonizer species: a strong and a weak competitor. The results suggest that the most important factor determining the impact of a clonal plant on its environment is ramet survival; the rate of ramet production is only secondary. Phenotypic plasticity in the production of ramets does not appear to be important: it has only minor effect on resource translocation and on the availability of colonizable gaps.     

两种践踏胁迫下克隆整合对入侵植物空心莲子草生长的影响

研究了两种践踏胁迫下克隆整合对入侵植物空心莲子草生长的影响.结果表明:(1)切断分株间的匍匐茎连接,会降低先端分株的生物量、分株数、总匍匐茎长度和总叶片数,但会显著增强基端分株的生物量.(2)对先端分株的践踏胁迫会显著降低先端分株叶片的叶绿素相对含量,对基端分株的践踏胁迫会显著降低基端分株的生物量和总匍匐茎长度.(3)对于基端分株的分株数、总匍匐茎长度和总叶片数来说,当进行基端分株践踏胁迫时,匍匐茎连接对其影响不大,而当进行先端分株践踏胁迫时,则明显对其不利.(4)对于整个克隆片段,践踏胁迫的差异和匍匐茎是否切断对其生长没有显著影响.     

分株比例对异质光环境下美丽箬竹克隆系统光合生理的影响

分株数量或生物量比例差异会明显影响克隆系统对资源异质性环境的生态适应性, 地下茎木质化、连接稳固的竹类植物在生长过程中相连克隆分株通常会生活在异质光环境中, 但其叶片光合生理特性对异质光环境的响应及其分株比例效应则未见报道。该研究以地下茎相连的美丽箬竹(Indocalamus decorus)克隆系统为实验材料, 设置2个遮光率(分别为50% ± 5%和75% ± 5%)和3个分株比例(遮光与未遮光分株比例分别为1:3、2:2、3:1)处理, 分株数量为4株。分别测定了遮光处理后30、90、150天遮光和未遮光分株叶片光响应特征、气体交换参数、光合色素含量, 分析了异质光环境下美丽箬竹光合生理的变化规律。结果显示: 分株比例对美丽箬竹光合生理有显著影响, 且其与遮光、处理时间交互作用显著。美丽箬竹克隆系统遮光分株比例越大, 即遮光相对分株数量越多, 其表观量子效率(AQE)、光饱和点、最大净光合速率(P_{n max})、净光合速率(P_n)、气孔导度(G_s)、蒸腾速率(T_r)、水分利用效率越大, 光补偿点、暗呼吸速率越小, 其光合效率越高, 光能利用能力越强, 而与之相连的未遮光分株则相反; 随遮光分株比例的增大, 遮光分株叶片叶绿素a、叶绿素b含量呈先升高而后下降的变化趋势, 类胡萝卜素(Car)含量则持续下降, 而与之相连的未遮光分株叶片光合色素含量则呈下降趋势; 遮光率提高, 相同分株比例美丽箬竹克隆系统遮光分株叶片AQE、P_{n max}、P_n、G_s、胞间CO₂浓度(C_i)和光合色素含量总体升高, 而与之相连的未遮光分株叶片P_{n max}、C_i以及Car含量则总体下降。研究结果表明异质光环境下, 遮光分株光合效率和弱光利用能力明显增强, 而未遮光分株则相反, 克隆系统内分株间发生了明显的克隆分工, 且2:2、3:1分株比例克隆系统较1:3分株比例对异质光环境具有更好的适应能力。美丽箬竹克隆系统可通过差异性调节分株光合生理特性和光合色素含量, 提高遮光分株光能利用和光合效率来适应异质光环境, 以提高克隆系统的适合度。     

Clonal integration facilitates spread of Paspalum paspaloides from terrestrial to cadmium‐contaminated aquatic habitats

• Cadmium (Cd) is a hazardous environmental pollutant with high toxicity to plants, which has been detected in many wetlands. Clonal integration (resource translocation) between connected ramets of clonal plants can increase their tolerance to stress. We hypothesised that clonal integration facilitates spread of amphibious clonal plants from terrestrial to Cd‐contaminated aquatic habitats.
• The spread of an amphibious grass Paspalum paspaloides was simulated by growing basal older ramets in uncontaminated soil connected (allowing integration) or not connected (preventing integration) to apical younger ramets of the same fragments in Cd‐contaminated water.
• Cd contamination of apical ramets of P. paspaloides markedly decreased growth and photosynthetic capacity of the apical ramets without connection to the basal ramets, but did not decrease these properties with connection. Cd contamination did not affect growth of the basal ramets without connection to the apical ramets, but Cd contamination of 4 and 12 mg·l^?1 significantly increased growth with connection. Consequently, clonal integration increased growth of the apical ramets, basal ramets and whole clones when the apical ramets were grown in Cd‐contaminated water of 4 and 12 mg·l^?1. Cd was detected in the basal ramets with connection to the apical ramets, suggesting Cd could be translocated due to clonal integration. Clonal integration, most likely through translocation of photosynthates, can support P. paspaloides to spread from terrestrial to Cd‐contaminated aquatic habitats.
• Amphibious clonal plants with a high ability for clonal integration are particularly useful for re‐vegetation of degraded aquatic habitats caused by Cd contamination.

     

Fitness and evolution in clonal plants: the impact of clonal growth

Seeds have often been emphasized in estimates of plant fitness because they are the units that carry genes to the next generation, disperse, and found new populations. We contend that clonal growth also needs to be considered when estimating fitness in clonal plants, regardless of whether fitness is measured from a genet or ramet perspective. Clonal growth affects genet fitness through both genet persistence and seed production. It affects ramet fitness through new ramet production, because both seeds and clonal propagants are considered offspring. The differential production of clonal propagants will contribute to fitness differences among individuals which may result in population-level changes in allele frequencies (i.e. microevolution). We describe a form of selection unique to clonal organisms, genotypic selection, that can result in evolution. Genotypic selection occurs when genotypically based traits are associated with differences in the rate of ramet production. It can lead to evolutionary change in quantitative trait means both directly and indirectly. It leads directly to change in the ramet population by increasing the proportion of ramets with more advantageous trait values. From the genet perspective, it leads indirectly to evolution within and among populations whenever significant portions of the genetic effect on a trait are inherited through seed. We argue that under most conditions, clonal growth will play a major role in the microevolution of clonal plants.     

Nutrient addition does not increase the benefits of clonal integration in an invasive plant spreading from open patches into plant communities

• One benefit of clonal integration is that resource translocation between connected ramets enhances the growth of the ramets grown under stressful conditions, but whether such resource translocation reduces the performance of the ramets grown under favourable conditions has not produced consistent results. In this study, we tested the hypothesis that resource translocation to recipient ramets may reduce the performance of donor ramets when resources are limiting but not when resources are abundant.
• We grew Mikania micrantha stolon fragments (each consisting of two ramets, either connected or not connected) under spatially heterogeneous competition conditions such that the developmentally younger, distal ramets were grown in competition with a plant community and the developmentally older, proximal ramets were grown without competition. For half of the stolon fragments, slow‐release fertiliser pellets were applied to both the distal and proximal ramets.
• Under both the low and increased soil nutrient conditions, the biomass, leaf number and stolon length of the distal ramets were higher, and those of the proximal ramets were lower when the stolon internode was intact than when it was severed. For the whole clone, the biomass, leaf number and stolon length did not differ between the two connection treatments. Connection did not change the biomass of the plant communities competing with distal ramets of M. micrantha.
• Although clonal integration may promote the invasion of M. micrantha into plant communities, resource translocation to recipient ramets of M. micrantha will induce a cost to the donor ramets, even when resources are relatively abundant.

     

Effects of physiological integration on defense strategies against herbivory by the clonal plant Alternanthera philoxeroides

Aims The plant–herbivore interaction is one of the most fundamental interactions in nature. Plants are sessile organisms, and consequently rely on particular strategies to avoid or reduce the negative impact of herbivory. Here, we aimed to determine the defense strategies against insect herbivores in the creeping invasive plantAlternanthera philoxeroides.     

Effect of physiological integration in self/non-self genotype recognition on the clonal invader Carpobrotus edulis

Aims Biological invasions represent one of the most important threats to the conservation of biodiversity; however, the mechanisms underlying successful invaders remain unsolved. Many of the most aggressive invaders show clonal growth, and capacity for clonal integration has been pointed out recently as an important trait explaining the success of invasive plants. We aim to determine the role of physiological integration in the capacity for self/non-self genotype recognition in the clonal invader Carpobrotus edulis and the implications of this capacity for the expansion of this aggressive invader.Methods We used connected and severed ramets of identical or different genotype and we determined the capacity for self/non-self recognition by comparing changes in biomass partitioning to avoid competition for resources between pairs of ramets.Important findings Physiological integration allowed self/non-self genotype recognition in the invader C. edulis. Results showed a significant effect of physiological integration on the biomass allocated to roots by genetically identical ramets: older ramets specialize in acquisition of soil-based resources and younger ramets specialize in lateral expansion. This specialization could be considered a form of division of labour, which reduce intra-genotype competition. This is the first evidence that division of labour could be interpreted as a form of self/non-self recognition between genetically identical ramets. Capacity for self/non-self discrimination could contribute to increase the colonization capacity of the aggressive invader C. edulis. This is the first study showing an association between self/non-self recognition and invasiveness in a clonal plant.