Bi-Cheng Dong

Effects of fragmentation on the survival and growth of the invasive, clonal plant Alternanthera philoxeroides

The capacity of small clonal fragments to survive and grow is a major factor in the spread of clonal plants, especially aquatic species. This study of the introduced, invasive, amphibious, stoloniferous herb Alternanthera philoxeroides in China tested the effects of fragment size, of loss of the stolon apex, and of the original position of a fragment within a clone before fragmentation on the survival and growth of fragments in a greenhouse experiment. The stolon internodes of plants consisting of one stolon with 5 ramets were severed in all 16 possible patterns, generating fragments of 1–5 ramets. These 16 fragmentation treatments were crossed with removal of the stolon apex. Fragmentation and apex removal did not affect total growth of plants, but more highly fragmented plants produced smaller, more numerous ramets. The survival of fragments consisting of one original ramet was about 85%, compared to 100% survival of fragments with 5 original ramets. Fragments consisting of the youngest ramets along the original stolon grew more than fragments of the same size that consisted of older ramets. These effects of fragmentation on growth provide new evidence for clonal integration of the rate at which connected ramets produce new ramets and novel evidence that fragments produced from different portions of clones have different potential for spread. They further suggest that detachment of small groups of distal ramets along stolons in the field might be very effective in spreading the species and that a trade-off between the number and the size of new ramets produced by fragments may represent an adaptive, plastic response to disturbance. Results do not recommend intentional fragmentation of clones as a means of controlling the rate of growth of this invasive species but suggest that incidental fragmentation may not increase its growth in mass, although it may increase its rate of spread.

Effects of soil nutrient heterogeneity on intraspecific competition in the invasive, clonal plant Alternanthera philoxeroides.

BACKGROUND AND AIMS Fine-scale, spatial heterogeneity in soil nutrient availability can increase the growth of individual plants, the productivity of plant communities and interspecific competition. If this is due to the ability of plants to concentrate their roots where nutrient levels are high, then nutrient heterogeneity should have little effect on intraspecific competition, especially when there are no genotypic differences between individuals in root plasticity. We tested this hypothesis in a widespread, clonal species in which individual plants are known to respond to nutrient heterogeneity. METHODS Plants derived from a single clone of Alternanthera philoxeroides were grown in the greenhouse at low or high density (four or 16 plants per 27·5 × 27·5-cm container) with homogeneous or heterogeneous availability of soil nutrients, keeping total nutrient availability per container constant. After 9 weeks, measurements of size, dry mass and morphology were taken. KEY RESULTS Plants grew more in the heterogeneous than in the homogeneous treatment, showing that heterogeneity promoted performance; they grew less in the high- than in the low-density treatment, showing that plants competed. There was no interactive effect of nutrient heterogeneity and plant density, supporting the hypothesis that heterogeneity does not affect intraspecific competition in the absence of genotypic differences in plasticity. Treatments did not affect morphological characteristics such as specific leaf area or root/shoot ratio. CONCLUSIONS Results indicate that fine-scale, spatial heterogeneity in the availability of soil nutrients does not increase competition when plants are genetically identical, consistent with the suggestion that effects of heterogeneity on competition depend upon differences in plasticity between individuals. Heterogeneity is only likely to increase the spread of monoclonal, invasive populations such as that of A. philoxeroides in China.

Burial Depth and Stolon Internode Length Independently Affect Survival of Small Clonal Fragments

Disturbance can fragment plant clones into different sizes and unstabilize soils to different degrees, so that clonal fragments of different sizes can be buried in soils at different depths. As a short-term storage organ, solon internode may help fragmented clones of stoloniferous plants to withstand deeper burial in soils. We address (1) whether burial in soils decreases survival and growth of small clonal fragments, and (2) whether increasing internode length increases survival and growth of small fragments under burial. We conducted an experiment with the stoloniferous, invasive herb Alternanthera philoxeroides, in which single-node fragments with stolon internode of 0, 2, 4 and 8 cm were buried in soils at 0, 2, 4 and 8 cm depth, respectively. Increasing burial depth significantly reduced survival of the A. philoxeroides plants and increased root to shoot ratio and total stolon length, but did not change growth measures. Increasing internode length significantly increased survival and growth measures, but there was no interaction effect with burial depth on any traits measured. These results indicate that reserves stored in stolon internodes can contribute to the fitness of the A. philoxeroides plants subject to disturbance. Although burial reduced the regeneration capacity of the A. philoxeroides plants, the species may maintain the fitness by changing biomass allocation and stolon length once it survived the burial. Such responses may play an important role for A. philoxeroides in establishment and invasiveness in frequently disturbed habitats.

Effects of Orientation on Survival and Growth of Small Fragments of the Invasive, Clonal Plant Alternanthera philoxeroides

Background The ability of small clonal fragments to establish and grow after disturbance is an important ecological advantage of clonal growth in plants and a major factor in the invasiveness of some introduced, clonal species. We hypothesized that orientation in the horizontal position (typical for stoloniferous plants) can increase the survival and growth of dispersed clonal fragments, and that this effect of orientation can be stronger when fragments are smaller and thus have fewer reserves to support initial growth. Methodology/Principal Findings To test these hypotheses, we compared performance of single-node pieces of stolon fragments of Alternanthera philoxeroides planted at angles of 0, 45 or 90° away from the horizontal position, with either the distal or the proximal end of the fragment up and with either 1 or 3 cm of stolon left attached both distal and proximal to the ramet. As expected, survival and growth were greatest when fragments were positioned horizontally. Contrary to expectations, some of these effects of orientation were stronger when attached stolons were longer. Orientation had smaller effects than stolon length on the performance of fragments; survival of fragments was about 60% with shorter stolons and 90% with longer stolons. Conclusions/Significance Results supported the hypothesis that orientation can affect establishment of small clonal fragments, suggested that effects of orientation can be stronger in larger rather than smaller fragments, and indicated that orientation may have less effect on establishment than amount of stored resources.

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 worlds 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.

Effects of Heterogeneous Competitor Distribution and Ramet Aggregation on the Growth and Size Structure of a Clonal Plant

Spatially heterogeneous distribution of interspecific competitors and intraspecific aggregation of offspring ramets may affect the growth and size structure of clonal plant populations, but these have been rarely studied. We conducted a greenhouse experiment in which we grew a population of eight offspring ramets (plants) of the stoloniferous clonal plant Hydrocotyle vulgaris aggregately or segregately in two homogeneous treatments with or without a competing grass Festuca elata and a heterogeneous treatment with a patchy distribution of the grass. In patchy grass treatments, H. vulgaris produced markedly more biomass, ramets and stolons in open patches (without grasses) than in grass patches, but displayed lower size variations as measured by coefficient of variation of biomass, ramets and stolons among the eight plants. In open areas, H. vulgaris produced statistically the same amounts of biomass and even more stolons and showed higher size variations in patchy grass treatments than in open (no grass) treatments. In grass areas, H. vulgaris grew much worse and displayed higher size variations in patchy grass treatments than in full grass treatments. Ramet aggregation decreased the growth of H. vulgaris in open treatments and in both open and grass patches in patchy grass treatments, but had little effect in full grass treatments. Ramet aggregation had little effect on size variations. Therefore, heterogeneous distribution of competitors can affect the growth and size structure of clonal plant populations, and ramet aggregation may decrease population growth when they grow in open environments or heterogeneous environments with a patchy distribution of interspecific competitors.

Patchy Distributions of Competitors Affect the Growth of a Clonal Plant When the Competitor Density Is High

Environments are patchy in not only abiotic factors but also biotic ones. Many studies have examined effects of spatial heterogeneity in abiotic factors such as light, water and nutrients on the growth of clonal plants, but few have tested those in biotic factors. We conducted a greenhouse experiment to examine how patchy distributions of competitors affect the growth of a rhizomatous wetland plant Bolboschoenus planiculmis and whether such effects depend on the density of the competitors. We grew one ramet of B. planiculmis in the center of each of the experimental boxes without competitors (Schoenoplectus triqueter), with a homogeneous distribution of the competitors of low or high density, and with a patchy distribution of the competitors of low or high density. The presence of competitors markedly decreased the growth (biomass, number of ramets, number of tubers and rhizome length) of the B. planiculmis clones. When the density of the competitors was low, the growth of B. planiculmis did not differ significantly between the competitor patches and competitor-free patches. However, when the density of the competitors was high, the growth of B. planiculmis was significantly higher in the competitor-free patches than in the competitor patches. Therefore, B. planiculmis can respond to patchy distributions of competitors by placing more ramets in competition-free patches when the density of competitors is high, but cannot do so when the density of competitors is low.

Effects of vegetative propagule pressure on the establishment of an introduced clonal plant, Hydrocotyle vulgaris

Some introduced clonal plants spread mainly by vegetative (clonal) propagules due to the absence of sexual reproduction in the introduced range. Propagule pressure (i.e. total number of propagules) may affect the establishment and thus invasion success of introduced clonal plants, and such effects may also depend on habitat conditions. A greenhouse experiment with an introduced plant, Hydrocotyle vulgaris was conducted to investigate the role of propagule pressure on its invasion process. High (five ramets) or low (one ramet) propagule pressure was established either in bare soil or in an experimental plant community consisting of four grassland species. H. vulgaris produced more total biomass under high than under low propagule pressure in both habitat conditions. Interestingly, the size of the H. vulgaris individuals was smaller under high than under low propagule pressure in bare soil, whereas it did not differ between the two propagule pressure treatments in the grassland community. The results indicated that high propagule pressure can ensure the successful invasion in either the grass community or bare soil, and the shift in the intraspecific interaction of H. vulgaris from competition in the bare soil to facilitation in the grassland community may be a potential mechanism.

Clonal integration increases tolerance of a phalanx clonal plant to defoliation

Defoliation by herbivores commonly imposes negative effects on plants, and physiological integration (resource sharing) can enhance the ability of guerilla clonal plants to tolerate stresses. Here we examined whether physiological integration can increase the ability of phalanx clonal plants to withstand defoliation. On a high mountain grassland in southwestern China, we subjected the phalanx clonal plant Iris delavayi within 10cm×10cm plots to three levels of defoliation intensity, i.e., control (no defoliation), moderate (50% shoot removal to simulate moderate herbivory) and heavy defoliation (100% shoot removal to simulate heavy herbivory), and kept rhizomes at the plot edges connected (allowing physiological integration) or disconnected (preventing integration) with intact ramets outside the plots. Defoliation significantly reduced leaf biomass, root biomass and ramet number of I. delavayi. Clonal integration did not affect the growth of I. delavayi under control, but significantly increased total biomass, rhizome and root biomass under heavy defoliation, and leaf biomass and ramet number under moderate defoliation. We conclude that clonal integration associated with resource reallocation plays an important role in maintaining the productivity of the alpine and subalpine grassland ecosystems in SW China where clonal plants are a dominant component of the grasslands and are commonly extensively managed with moderate grazing intensity. Our results also help to better understand the adaption and tolerance of phalanx clonal plants subjected to long-term grazing in the high mountain environment.

Effects of soil substrate heterogeneity and moisture on interspecific competition between Alternanthera philoxeroides and four native species

Aims Clonal plant species have the potential for high relative performance in heterogeneous environments, and this might increase the competitive ability and invasiveness of introduced clonal plant species. It was hypothesized that clonal species whose performance responds more to heterogeneity of a resource have higher competitive ability in habitats where this resource is more heterogeneous and that this relationship is stronger when other resources are less limiting. Methods To test these hypotheses, the perennial clonal herb Alternanthera philoxeroides, which is invasive in China, was grown alone or with each of four clonal perennial, co-occurring herbs native to China, i.e. Alternanthera sessilis, Cynodon dactylon, Hemarthria altissima and Wedelia chinensis. Plants were given homogeneous or heterogeneous soil substrate crossed with low and high levels of soil moisture. Important Findings Effects of heterogeneity on the accumulation of mass and ramets and on competitive effect and response of A. philoxeroides differed between native species and interacted with effects of soil moisture. A. philoxeroides reduced the final total mass or ramet number of the native species except A. sessilis, and the negative competitive effects on H. altissima and C. dactylon were more pronounced in heterogeneous than in homogeneous soil. Competitive response of A. philoxeroides was more negative to A. sessilis than to the other native species. Across native species, the competitive response of A. philoxeroides was more negative in heterogeneous than in homogeneous soils at low moisture level, but the reverse was true at high moisture level. Results do not consistently support either hypothesis, but do suggest that competitive ability can be partly explained by individual species traits such as size, and that some competitive effects and responses are emergent properties of interspecific interactions.