Cultivated alien plants with high invasion potential are more likely to be traded online in China.

Biological invasions have become a worldwide problem, and measures to efficiently prevent and control invasions are still in development. Like many other parts of the world, China is undergoing a dramatic increase in plant invasions. Most of the currently 933 established (i.e., naturalized) plant species, of which 214 are categorized as invasive, have been introduced into China for cultivation. It is likely that many of those species are still being traded, particularly online, by plant nurseries. However, studies assessing whether naturalized and invasive species are currently being traded more or less than nonnaturalized aliens are rare. We extracted online-trade information for 13,718 cultivated alien plant taxa on 1688.com, the largest website for domestic B2B in China. We analyzed how the presence in online-nursery catalogs, the number of online nurseries that offerred the species for sale, and the product type (i.e., seeds, live plants and vegetative organs) differed among nonnaturalized, naturalized noninvasive, and invasive species. Compared to nonnaturalized taxa, naturalized noninvasive and invasive taxa were 3.7-5.2 times more likely to be available for purchase. Naturalized noninvasive and invasive taxa were more frequently offered as seeds by online nurseries, whereas nonnaturalized taxa were more frequently offered as live plants. Based on these findings, we propose that, to reduce the further spread of invasive and potentially invasive plants, implementation of plant-trade regulations and a monitoring system of the online horticultural supply chain will be essential.

Functional trait trade-offs define plant population stability across different biomes.

Ecological theory posits that temporal stability patterns in plant populations are associated with differences in species' ecological strategies. However, empirical evidence is lacking about which traits, or trade-offs, underlie species stability, especially across different biomes. We compiled a worldwide collection of long-term permanent vegetation records (greater than 7000 plots from 78 datasets) from a large range of habitats which we combined with existing trait databases. We tested whether the observed inter-annual variability in species abundance (coefficient of variation) was related to multiple individual traits. We found that populations with greater leaf dry matter content and seed mass were more stable over time. Despite the variability explained by these traits being low, their effect was consistent across different datasets. Other traits played a significant, albeit weaker, role in species stability, and the inclusion of multi-variate axes or phylogeny did not substantially modify nor improve predictions. These results provide empirical evidence and highlight the relevance of specific ecological trade-offs, i.e. in different resource-use and dispersal strategies, for plant populations stability across multiple biomes. Further research is, however, necessary to integrate and evaluate the role of other specific traits, often not available in databases, and intraspecific trait variability in modulating species stability.

The priming effects of plant leachates on dissolved organic matter degradation in water depend on leachate type and water stability.

The modification of dissolved organic matter (DOM) degradation by plant carbon inputs represents a critical biogeochemical process that controls carbon dynamics. However, the priming effects (PEs) different plant tissues induce on the degradation of DOM pools with different stabilities remain unknown. In this study, PEs, induced by different tissue leachates of Phragmites australis, were evaluated via changes in DOM components and properties of both fresh and tidal water (with different stabilities). The results showed that DOM derived from different plant tissue leachates differed in composition and bioavailability. Inputs of tissue leachates induced PEs with different intensities and directions (negative or positive) on DOM degradation of fresh and tidal water. In fresh water, the PEs of leaf and root leachates were significantly higher than those of stem and rhizome leachates. The PE direction changed for DOM degradation between fresh and tidal water. The addition of leaf and root leachates tended to induce positive PEs on DOM degradation of fresh water, while resulting in negative PEs on DOM degradation of tidal water. Negative PEs for tidal water DOM may be due to preferential utilization of microbes, high salinity, and/or the promotion of exogenous DOM production from plant tissues. The results indicate that intensity and direction of PEs induced by plant leachates depend on both leachate type and water stability. The findings highlight the necessity to examine the nature of exogenous and native DOM when interpreting the interactive processes that regulate DOM degradation.

Synchrony matters more than species richness in plant community stability at a global scale.

The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved. Our analysis of time series from 79 datasets across the world showed that stability was associated more strongly with the degree of synchrony among dominant species than with species richness. The relatively weak influence of species richness is consistent with theory predicting that the effect of richness on stability weakens when synchrony is higher than expected under random fluctuations, which was the case in most communities. Land management, nutrient addition, and climate change treatments had relatively weak and varying effects on stability, modifying how species richness, synchrony, and stability interact. Our results demonstrate the prevalence of biotic drivers on ecosystem stability, with the potential for environmental drivers to alter the intricate relationship among richness, synchrony, and stability.

Effects of resource availability on the growth, Cd accumulation, and photosynthetic efficiency of three hyperaccumulator plant species.

Plants that hyperaccumulate heavy metals such as cadmium (Cd) are important agents of phytoremediation. Availability of resources such as light, nutrients, and water can affect heavy metal accumulation by plants, but the responses of hyperaccumulators to different levels of resource availability remain little studied. To test such responses, three Cd hyperaccumulators, Solanum nigrum, Bidens pilosa, and Taraxacum mongolicum, were grown in Cd-contaminated soil; subjected to three levels of light, nutrient, or water availability; and measured for growth, Cd accumulation, and photosynthetic efficiency. All three species accumulated more total biomass and grew taller if given high than low water or light (each P < 0.001). Species accumulated four to eight times more Cd (190-309 versus 24-68 µg Cd g-1 mass) under high than low light. High water availability increased Cd accumulation by 89% in B. pilosa but decreased it by 31% and 40% in S. nigrum and B. pilosa, respectively. Effects of nutrients on both growth and accumulation varied between species; Cd accumulation by S. nigrum and T. mongolicum was respectively 14% and 54% lower at high than low nutrients, while it was 130% higher in B. pilosa. Light but not water or nutrient availability affected effective and maximum quantum yields and electron transport rate. Findings from this study suggest that low levels of light may constrain phytoremediation of Cd in soil and that testing species of Cd hyperaccumulators individually for responses to levels of water and nutrients will inform selection of species for phytoremediation.

Clonal functional traits favor the invasive success of alien plants into native communities.

Functional traits are frequently proposed to determine the invasiveness of alien species. However, few empirical studies have directly manipulated functional traits and tested their importance in the invasion success of alien species into native plant communities, particularly under global change. We manipulated clonal integration (a key clonal functional trait) of four alien clonal plants by severing inter-ramet connections or keeping them intact and simulated their invasion into native plant communities with two levels of species diversity, population density and nutrient availability. High community diversity and density impeded the invasion success of the alien clonal plants. Clonal integration of the alien plants promoted their invasion success, particularly in the low-density communities associated with low species diversity or nutrient addition, which resulted in a negative correlation between the performance of alien plants and native communities, as expected under global change. Thus, clonal integration can favor the invasion success of alien clonal plants into degraded resident communities with a high degree of disturbance and eutrophication. Our findings confirm the role of clonal functional traits in facilitating alien plant invasions into native plant communities and suggest that clonal functional traits should be considered to efficiently restore degraded communities heavily invaded by alien clonal plants.

Soil Microbe-Mediated N:P Stoichiometric Effects on Solidago canadensis Performance Depend on Nutrient Levels.

Both soil microbes and soil N:P ratios can affect plant growth, but it is unclear whether they can interact to alter plant growth and whether such an interactive effect depends on nutrient levels. Here, we tested the hypothesis that soil microbes can ameliorate the negative effects of nutrient imbalance caused by low or high N:P ratios on plant growth and that such an ameliorative effect of soil microbes depends on nutrient supply levels. We grew individuals of six populations of the clonal plant Solidago canadensis at three N:P ratios (low (1.7), intermediate (15), and high (135)), under two nutrient levels (low versus high) and in the presence versus absence of soil microbes. The presence of soil microbes significantly increased biomass of S. canadensis at all three N:P ratios and under both nutrient levels. Under the low-nutrient level, biomass, height, and leaf number of S. canadensis did not differ significantly among the three N:P ratio treatments in the absence of soil microbes, but they were higher at the high than at the low and the intermediate N:P ratio in the presence of soil microbes. Under the high-nutrient level, by contrast, biomass, height, and leaf number of S. canadensis were significantly higher at the low than at the high and the intermediate N:P ratio in the absence of soil microbes, but increased with increasing the N:P ratio in the presence of soil microbes. In the presence of soil microbes, number of ramets (asexual individuals) and the accumulation of N and P in plants were significantly higher at the high than at the low and the intermediate N:P ratio under both nutrient levels, whereas in the absence of soil microbes, they did not differ significantly among the three N:P ratio regardless of the nutrient levels. Our results provide empirical evidence that soil microbes can alter effects of N:P ratios on plant performance and that such an effect depends on nutrient availability. Soil microbes may, therefore, play a role in modulating ecosystem functions such as productivity and carbon and nutrient cycling via modulating nutrient imbalance caused by low and high N:P ratios.

Litter nitrogen concentration changes mediate effects of drought and plant species richness on litter decomposition.

Biodiversity loss, exotic plant invasion and climatic change are three important global changes that can affect litter decomposition. These effects may be interactive and these global changes thus need to be considered simultaneously. Here, we assembled herbaceous plant communities with five species richness levels (1, 2, 4, 8 or 16) and subjected them to a drought treatment (no, moderate or intensive drought) that was factorially combined with an invasion treatment (presence or absence of the non-native Symphyotrichum subulatum). We collected litter of these plant communities and let it decompose for 9 months in the plant communities from which it originated. Drought decreased litter decomposition, while invasion by S. subulatum had little impact. Increasing species richness decreased litter decomposition except under intensive drought. A structural equation model showed that drought and species richness affected litter decomposition indirectly through changes in litter nitrogen concentration rather than by altering quantity and diversity of soil meso-fauna or soil physico-chemical properties. The slowed litter decomposition under high species diversity originated from a sampling effect, specifically from low litter nitrogen concentrations in the two dominant species. We conclude that effects on litter decomposition rates that are mediated by changing concentrations of the limiting nutrient in litter need to be considered when predicting effects of global changes such as plant diversity loss.

Plant Diversity Enhances Soil Fungal Diversity and Microbial Resistance to Plant Invasion.

Interactions and feedbacks between aboveground and belowground biomes are fundamental in controlling ecosystem functions and stability. However, the relationship between plant diversity and soil microbial diversity is elusive. Moreover, it remains unknown whether plant diversity loss will cause the stability of soil microbial communities to deteriorate. To shed light on these questions, we conducted a pot-based experiment to manipulate the plant richness gradient (1, 2, 4, or 8 species) and plant [Symphyotrichum subulatum (Michx.) G. L. Nesom] invasion status. We found that, in the noninvasion treatment, soil fungal diversity significantly and positively correlated with plant diversity, while the relationship between bacterial and plant diversity was not significant. Under plant invasion conditions, the coupling of plant-fungal alpha diversity relationship was enhanced, but the plant-fungal beta diversity relationship was decoupled. We also found significant positive relationships between plant diversity and soil microbial resistance. The observed positive relationships were determined by turnover (species substitution) and nestedness (species loss) processes for bacterial and fungal communities, respectively. Our study demonstrated that plant diversity enhanced soil fungal diversity and microbial resistance in response to plant invasion. This study expands our knowledge about the aboveground-belowground diversity relationship and the diversity-stability relationship.IMPORTANCE Our study newly showed that plant invasion significantly altered relationships between aboveground and belowground diversity. Specifically, plant richness indirectly promoted soil fungal richness through the increase of soil total carbon (TC) without plant invasion, while plant richness had a direct positive effect on soil fungal richness under plant invasion conditions. Our study highlights the effect of plant diversity on soil fungal diversity, especially under plant invasion conditions, and the plant diversity effect on microbial resistance in response to plant invasion. These novel findings add important knowledge about the aboveground-belowground diversity relationship and the diversity-stability relationship.

Parasitism induces negative effects of physiological integration in a clonal plant.

Clonal integration often increases fitness of clonal plants, but it may decrease it when some but not all connected plants (ramets) within a clone are parasitized. This hypothesis was synthesized in a conceptual model and tested by growing pairs of connected ramets of two congeneric clonal plants, Sphagneticola trilobata and Sphagneticola calendulacea, with and without parasitizing one ramet with Cuscuta australis and with and without severing the connection (allowing or preventing integration). Consistent with the model, integration in S. calendulacea did not affect biomass of the parasitized ramet, decreased biomass of its connected, unparasitized ramet by 60% and of the clone by 40%, and increased biomass of the parasite by 50%. By contrast, integration in S. trilobata did not affect biomass of the clone or the parasite. The parasite increased export of nitrogen-15 from the connected, unparasitized ramet seven-fold in S. calendulacea but did not affect export in S. trilobata. Parasitism can cause clonal integration to negatively affect fitness in clonal plants because parasites can import resources from connected, unparasitized ramets, possibly partly through signaling. This is the first experimental demonstration that clonal integration can decrease fitness in plants induced by parasitism and may help explain community-level effects of parasites.

Effects of fragmentation of clones compound over vegetative generations in the floating plant Pistia stratiotes.

BACKGROUND AND AIMS: Clonal plants dominate many plant communities, especially in aquatic systems, and clonality appears to promote invasiveness and to affect how diversity changes in response to disturbance and resource availability. Understanding how the special physiological and morphological properties of clonal growth lead to these ecological effects depends upon studying the long-term consequences of clonal growth properties across vegetative generations, but this has rarely been done. This study aimed to show how a key clonal property, physiological integration between connected ramets within clones, affects the response of clones to disturbance and resources in an aquatic, invasive, dominant species across multiple generations. METHODS: Single, parental ramets of the floating stoloniferous plant Pistia stratiotes were grown for 3 weeks, during which they produced two or three generations of offspring; connections between new ramets were cut or left intact. Individual offspring were then used as parents in a second 3-week iteration that crossed fragmentation with previous fragmentation in the first iteration. A third iteration yielded eight treatment combinations, zero to three rounds of fragmentation at different times in the past. The experiment was run once at a high and once at a low level of nutrients. RESULTS: In each iteration, fragmentation increased biomass of the parental ramet, decreased biomass of the offspring and increased number of offspring. These effects persisted and compounded from one iteration to another, though more recent fragmentation had stronger effects, and were stronger at the low than at the high nutrient level. Fragmentation did not affect net accumulation of mass by groups after one iteration but increased it after two iterations at low nutrients, and after three iterations at both nutrient levels. CONCLUSIONS: Both the positive and negative effects of fragmentation on clonal performance can compound and persist over time and can be stronger when resource levels are lower. Even when fragmentation has no short-term net effect on clonal performance, it can have a longer-term effect. In some cases, fragmentation may increase total accumulation of mass by a clone. The results provide the first demonstration of how physiological integration in clonal plants can affect fitness across generations and suggest that increased disturbance may promote invasion of introduced clonal species via effects on integration, perhaps especially at lower nutrient levels.

Physiological integration can increase competitive ability in clonal plants if competition is patchy.

Physiological integration of connected plants of the same clone, or ramets, often increases clonal fitness when ramets differ in resource supply. However, review of the literature found that no study has directly tested the hypothesis that integration can increase the ability of clones to compete against other species. To test this, we grew two-ramet clonal fragments of the stoloniferous, perennial herb Fragaria chiloensis in which none, one, or both of the ramets had neighbors of a naturally co-occurring, dominant grass, Bromus carinatus, and connections between ramets were either severed to prevent integration or left intact. We also grew four-ramet fragments in which all ramets had neighbors and connections were severed or intact. Severance decreased the final leaf mass and area of two-ramet fragments by 25% and their final total mass by 15% when just one ramet was grown with B. carinatus. Severance had no significant effect on the total mass of fragments when none or all of the ramets were grown with the grass. This provides the first direct evidence that physiological integration can increase the competitive ability of clonal plant species, though only when competition is spatially heterogeneous. Integration may thus enable plant clones to grow into plant communities and to compete within communities with fine-scale disturbance. However, integration may not increase the competitive ability of clonal plants within uniformly dense communities of taller species.

When facilitation meets clonal integration in forest canopies.

Few studies have explored how - within the same system - clonality and positive plant-plant interactions might interact to regulate plant community composition. Canopy-dwelling epiphytes in species-rich forests provide an ideal system for studying this because many epiphytic vascular plants undertake clonal growth and because vascular epiphytes colonize canopy habitats after the formation of nonvascular epiphyte (i.e. bryophyte and lichen) mats. We investigated how clonal integration of seven dominant vascular epiphytes influenced inter-specific interactions between vascular epiphytes and nonvascular epiphytes in a subtropical montane moist forest in southwest China. Both clonal integration and environmental buffering from nonvascular epiphytes increased survival and growth of vascular epiphytes. The benefits of clonal integration for vascular epiphytes were higher when nonvascular epiphytes were removed. Similarly, facilitation from nonvascular epiphytes played a more important role when clonal integration of vascular epiphytes was eliminated. Overall, clonal integration had greater benefits than inter-specific facilitation. This study provides novel evidence for interactive effects of clonality and facilitation between vascular and nonvascular species, and has implications for our understanding of a wide range of ecosystems where both high levels of clonality and facilitation are expected to occur.

Correlations between genetic, epigenetic and phenotypic variation of an introduced clonal herb.

Heritable epigenetic modifications may occur in response to environmental variation, further altering phenotypes through gene regulation, without genome sequence changes. However, epigenetic variation in wild plant populations and their correlations with genetic and phenotypic variation remain largely unknown, especially for clonal plants. We investigated genetic, epigenetic and phenotypic variation of ten populations of an introduced clonal herb Hydrocotyle vulgaris in China. Populations of H. vulgaris exhibited extremely low genetic diversity with one genotype exclusively dominant, but significantly higher epigenetic diversity. Both intra- and inter-population epigenetic variation were related to genetic variation. But there was no correlation between intra-/inter-population genetic variation and phenotypic variation. When genetic variation was controlled, intra-population epigenetic diversity was related to petiole length, specific leaf area, and leaf area variation, while inter-population epigenetic distance was correlated with leaf area differentiation. Our study provides empirical evidence that even though epigenetic variation is partly under genetic control, it could also independently play a role in shaping plant phenotypes, possibly serving as a pathway to accelerate evolution of clonal plant populations.

Synergistic Effects of Soil Microbes on Solidago canadensis Depend on Water and Nutrient Availability.

Soil microbes may greatly affect plant growth. While plants are commonly associated with diverse communities of soil microbes, complementary roles of different microbial communities that may stimulate synergistic effects on plant growth are not adequately tested. Also, such synergistic effects may vary with environmental conditions such as soil nutrient and water availability. We conducted a greenhouse experiment with a widespread clonal plant Solidago canadensis. The experiment was a factorial design with four levels of soil microbial inoculation (fresh soil inocula from grasslands in northern and southern China that were expected to differ in soil microbial composition, a mixture of the two fresh soil inocula, and a sterilized mixed inoculum control), two levels of nutrient availability (low vs. high), and two levels of water supply (low vs. high, i.e., 1376 vs. 352 mm per year). Irrespective of water supply and nutrient availability, total, aboveground, and belowground mass of S. canadensis were generally higher when the plant grew in soil inoculated with a mixture of soil microbes from the south and north of China (in the mixed inoculum treatment) than when it grew in soil inoculated with soil microbes from only the north or the south or the sterilized control. Such effects of soil microbes on total and aboveground mass were stronger under high than under low nutrient availability and also under high than under low water supply. Our results suggest that interactions of different soil microbial communities can result in a synergistic effect on plant growth and such a synergistic effect depends on environmental conditions. The findings shed light on the importance of plant-microbe interactions during the spreading of some plant species in face of increased atmospheric nutrient deposition coupled with altered rainfall pattern due to global change.

Soil resource heterogeneity competitively favors an invasive clonal plant over a native one.

Soil resource heterogeneity can affect plant growth and competitive ability. However, little is known about how soil resource heterogeneity affects competitive interactions between invasive and native plants. We conducted an experiment with an invasive clonal plant Alternanthera philoxeroides and a coexisting native one Alternanthera sessilis. The experiment was a randomized design with three factors, i.e. two species (A. philoxeroides and A. sessilis), two interspecific competition treatments (with and without) and five soil treatments (three homogeneous treatments and two small-scale heterogeneous treatments consisting of two patches of 10 cm × 15 cm and with different initial planting positions). Irrespective of competition, increasing soil resource availability increased the growth of A. philoxeroides. Increasing soil resource availability also increased the growth of A. sessilis without competition, but had no impact with competition. Irrespective of competition, soil resource heterogeneity increased biomass and ramet production of A. philoxeroides, and such effects were independent of initial planting position. For A. sessilis, however, soil resource heterogeneity only increased ramet production when the initial plant was grown in the low-resource patch without competition. Our results suggest that both high soil resource availability and small-scale soil resource heterogeneity can increase the relative competitive ability of the invasive plant A. philoxeroides when grown with its native congener A. sessilis. These findings may partly explain the invasion success of this clonal species in area with high soil resource availability and heterogeneity caused by e.g. nitrogen deposition, fertilization and disturbance.

Remediation of cadmium-contaminated coastal saline-alkaline soil by Spartina alterniflora derived biochar.

Following oil extraction in the wetland of the Yellow River Delta, heavy metal contamination of coastal saline-alkaline soil, especially with cadmium (Cd), has become a serious environmental problem in some regions. Biochar application has been proposed to remedy Cd-contaminated soil, but the remediation effect is related to preparation conditions of biochar (e.g., pyrolysis temperature and raw material) and soil properties. The invasive plant, Spartina alterniflora, produces a high amount of biomass, making it suitable for biochar production in coastal China. We investigated the effect of S. alterniflora-derived biochar (SDB) pyrolyzed at four temperatures (350, 450, 550, and 650 °C) crossed with three addition ratios (1, 5, and 10%) and control on Cd contamination of coastal saline-alkaline soil. Pyrolysis temperature affected pH, surface area, and functional groups of SDB. SDB markedly improved soil pH and soil organic matter, but the degree of improvement was affected by pyrolysis temperature and addition ratio. SDB significantly altered available Cd content in soil, but reduced it only at low pyrolysis temperatures (350 and 450 °C). Available Cd content had a positive correlation with soil pH (R2 = 0.298, P < 0.01), but was not related to salinity and soil organic matter content. Thus, SDB pyrolyzed at 350 °C with 5% addition was optimal for passivating Cd in coastal saline-alkaline soil, since available Cd content in soil decreased mostly (by 26.9%). These findings act as a reference for the development of an application strategy for SDB to ameliorate Cd-contaminated coastal saline-alkaline soil.

Constraints on the evolution of phenotypic plasticity in the clonal plant Hydrocotyle vulgaris.

The evolution of phenotypic plasticity of plant traits may be constrained by costs and limits. However, the precise constraints are still unclear for many traits under different ecological contexts. In a glasshouse experiment, we grew ramets of 12 genotypes of a clonal plant Hydrocotyle vulgaris under the control (full light and no flood), shade and flood conditions and tested the potential costs and limits of plasticity in 13 morphological and physiological traits in response to light availability and flood variation. In particular, we used multiple regression and correlation analyses to evaluate potential plasticity costs, developmental instability costs and developmental range limits of each trait. We detected significant costs of plasticity in specific petiole length and specific leaf area in response to shade under the full light condition and developmental range limits in specific internode length and intercellular CO2 concentration in response to light availability variation. However, we did not observe significant costs or limits of plasticity in any of the 13 traits in response to flood variation. Our results suggest that the evolution of phenotypic plasticity in plant traits can be constrained by costs and limits, but such constraints may be infrequent and differ under different environmental contexts.

Invasive alien plants benefit more from clonal integration in heterogeneous environments than natives.

What confers invasive alien plants a competitive advantage over native plants remains open to debate. Many of the world's worst invasive alien plants are clonal and able to share resources within clones (clonal integration), particularly in heterogeneous environments. Here, we tested the hypothesis that clonal integration benefits invasive clonal plants more than natives and thus confers invasives a competitive advantage. We selected five congeneric and naturally co-occurring pairs of invasive alien and native clonal plants in China, and grew pairs of connected and disconnected ramets under heterogeneous light, soil nutrient and water conditions that are commonly encountered by alien plants during their invasion into new areas. Clonal integration increased biomass of all plants in all three heterogeneous resource environments. However, invasive plants benefited more from clonal integration than natives. Consequently, invasive plants produced more biomass than natives. Our results indicate that clonal integration may confer invasive alien clonal plants a competitive advantage over natives. Therefore, differences in the ability of clonal integration could potentially explain, at least partly, the invasion success of alien clonal plants in areas where resources are heterogeneously distributed.

Clonal integration increases relative competitive ability in an invasive aquatic plant.

PREMISE OF THE STUDY: Physiological integration between connected ramets is well known to increase performance of clonal plant species. However, no direct evidence appears to exist that integration can increase the ability of clonal species to compete with other species within mixed communities. We tested this hypothesis using two floating, invasive, aquatic species in which fragmentation-and thus extent of integration-is likely to vary between habitats and times. METHODS: Individual ramets of Pistia stratiotes and Eichhornia crassipes were grown in monoculture or in mixture, and new stolons bearing new offspring were severed or left intact. After 6 wk, the numbers of offspring and second-generation (2°) offspring produced by each original ramet, or parent, were counted; and the final dry mass of each parent, its stolons, its offspring, and its 2° offspring were measured. KEY RESULTS: Fragmentation decreased the relative competitive ability of Pistia, but not that of Eichhornia. This was mainly because Pistia accumulated ∼30% less dry mass of offspring when fragmented and grown with Eichhornia than in other treatments. Offspring of Pistia were smaller than those of Eichhornia in all treatments. CONCLUSIONS: Our results show that clonal integration can increase competitive ability in some clonal species. In this case, integration appeared to enable the small offspring of Pistia to compete more effectively with the large offspring of Eichhornia. Lower rates of fragmentation may select for production of more numerous, smaller vegetative offspring in clonal species.