Wei-Ming He

Novel weapons and invasion: biogeographic differences in the competitive effects of Centaurea maculosa and its root exudate (±)-catechin

Recent studies suggest that the invasive success of Centaurea maculosa may be related to its stronger allelopathic effects on native North American species than on related European species, one component of the “novel weapons” hypothesis. Other research indicates that C. maculosa plants from the invasive range in North America have evolved to be larger and better competitors than conspecifics from the native range in Europe, a component of the “evolution of increased competitive ability” hypothesis. These hypotheses are not mutually exclusive, but this evidence sets the stage for comparing the relative importance of evolved competitive ability to inherent competitive traits. In a competition experiment with a large number of C. maculosa populations, we found no difference in the competitive effects of C. maculosa plants from North America and Europe on other species. However, both North American and European C. maculosa were much better competitors against plants native to North America than congeners native to Romania, collected in areas where C. maculosa is also native. These results are consistent with the novel weapons hypothesis. But, in a second experiment using just one population from North America and Europe, and where North American and European species were collected from a broader range of sites, competitive interactions were weaker overall, and the competitive effects of C. maculosa were slightly stronger against European species than against North American species. Also consistent with the novel weapons hypothesis, (±)-catechin had stronger effects on native North American species than on native European species in two experiments. Our results suggest that the regional composition of the plant communities being invaded by C. maculosa may be more important for invasive success than the evolution of increased size and competitive ability.

Clonal Integration Affects Growth, Photosynthetic Efficiency and Biomass Allocation, but not the Competitive Ability, of the Alien Invasive Alternanthera philoxeroides under Severe Stress

BACKGROUND AND AIMS Many notorious alien invasive plants are clonal, but little is known about some roles and aspects of clonal integration. Here, the hypothesis is tested that clonal integration affects growth, photosynthetic efficiency, biomass allocation and competitive ability of the exotic invasive weed Alternanthera philoxeroides (Amaranthaceae). METHODS The apical parts of Alternanthera were grown either with or without the lawn grass Schedonorus phoenix (tall fescue) and their stolon connections to the basal parts grown without competitors were either severed or left intact. KEY RESULTS Competition greatly reduced the maximum quantum yield of photosystem II (F(v)/F(m)) and growth (biomass, number of ramets and leaves, total stolon length and total leaf area) of the apical Alternanthera, but not the biomass of S. phoenix. Stolon connections significantly increased F(v)/F(m) and growth of Alternanthera. However, such effects on growth were smaller with than without competition and stolon connections did not alter the relative neighbour effect of Alternanthera. Stolon connections increased Alternantheras biomass allocation to roots without competition, but decreased it with competition. CONCLUSIONS Clonal integration contributed little to Alternantheras competitive ability, but was very important for Alternanthera to explore open space. The results suggest that the invasiveness of Alternanthera may be closely related to clonal integration.

Clonal integration supports the expansion from terrestrial to aquatic environments of the amphibious stoloniferous herb Alternanthera philoxeroides

Effects of clonal integration on land plants have been extensively studied, but little is known about the role in amphibious plants that expand from terrestrial to aquatic conditions. We simulated expansion from terrestrial to aquatic habitats in the amphibious stoloniferous alien invasive alligator weed (Alternanthera philoxeroides) by growing basal ramets of clonal fragments in soils connected (allowing integration) or disconnected (preventing integration) to the apical ramets of the same fragments submerged in water to a depth of 0, 5, 10 or 15 cm. Clonal integration significantly increased growth and clonal reproduction of the apical ramets, but decreased both of these characteristics in basal ramets. Consequently, integration did not affect the performance of whole clonal fragments. We propose that alligator weed possesses a double-edged mechanism during population expansion: apical ramets in aquatic habitats can increase growth through connected basal parts in terrestrial habitats; however, once stolon connections with apical ramets are lost by external disturbance, the basal ramets in terrestrial habitats increase stolon and ramet production for rapid spreading. This may contribute greatly to the invasiveness of alligator weed and also make it very adaptable to habitats with heavy disturbance and/or highly heterogeneous resource supply.

Adaptation of Rhizome Connections in Drylands: Increasing Tolerance of Clones to Wind Erosion

BACKGROUND AND AIMS Wind erosion is a severe stress for plants in drylands, but the mechanisms by which plants withstand erosion remain largely unknown. Here, the hypothesis is tested that maintaining rhizome connections helps plants to tolerate erosion. METHODS Five transects were established across an inland dune in Inner Mongolia, China, and measurements were made of leaf number, biomass per ramet and rhizome depth of Psammochloa villosa in 45 plots. In 40 x 40 cm plots of P. villosa on another dune, the top 15 or 30 cm of sand was removed for 1.5 or 3 months to simulate short- and long-term moderate and severe erosion, respectively, with untreated plots as controls, and the rhizomes at the edges of half of the plots were severed to mimic loss of rhizome connections. KEY RESULTS Leaf number and biomass per ramet showed quadric relationships with rhizome depth; when rhizomes were exposed to the air, the associated ramets either died or became very weak. Ramet number, leaf number and biomass per plot decreased with increasing erosion severity. Rhizome connections did not affect these traits under control or short-term erosion, but increased them under long-term erosion. CONCLUSIONS Rhizome connections alleviated the negative effects of erosion on P. villosa, very likely because the erosion-stressed ramets received water and/or photosynthates translocated from those connected ramets that were not subject to erosion. This study provides the first evidence that maintaining rhizome connections helps plants to tolerate erosion in drylands.

Physiological integration in an introduced, invasive plant increases its spread into experimental communities and modifies their structure

What determines the invasiveness of introduced plants is still poorly known. Many of the most invasive plant species are clonal, and physiological integration between connected individuals (ramets) of clonal plants may contribute to their ability to spread into communities and reduce performance of existing species. This contribution of integration to the invasiveness of clonal plants may be greater in denser communities. A greenhouse study was conducted to test these two hypotheses. High- and low-density communities were created by sowing seeds of eight grassland species. Each community was planted with three ramets of the stoloniferous, introduced plant Alternanthera philoxeroides that were disconnected from or left connected to ramets growing on bare soil. Connection increased the spread of Alternanthera within a community, but did not reduce community biomass. Alternanthera grew less in high-density communities, but connection did not improve its growth more than in low-density communities. Low-density communities had higher evenness when Alternanthera was connected than when it was disconnected because shoot mass was lower in the more abundant species in the community and higher in the less abundant ones. These results partly supported the first hypothesis, but not the second. The effect of integration on community structure could be due to higher resource import by the ramets of Alternanthera closer to the dominant species. Integration therefore can increase the initial spread of new clonal plant species into communities and modify the effects of this spread on community structure.

Differential belowground allelopathic effects of leaf and root of Mikania micrantha

This study evaluates the relative contributions of leaves and roots to the belowground allelopathic effects of Mikania micrantha. The hypothesis that leaves contribute more to the allelopathic effect than roots was experimentally tested. We assessed the allelopathic effects of aqueous extracts from Mikania leaves and roots on the seed germination and seedling growth of two co-occurring woody plants in southern China, Lagerstroemia indica L. and Robinia pseudoacacia L. The results showed that the aqueous extracts from Mikania leaves and roots had inhibitory effects on the woody species. Allelopathic activity depended on the concentration of the extracts, target species, and the extract sources (i.e., leaves vs. roots of Mikania). Leaf extract showed stronger allelopathic effects than root extract on germination percentage, initial germination time, speed of germination, and shoot height; while root extract had greater allelopathic effects on roots than leaf extract. The latter phenomenon might greatly promote the invasion success of Mikania due to more direct and effective allelopathy of root. Our results suggest that allelopathy of root extract on belowground biomass might be greater than that of leaf extract for some species in contrast allelopathy of leaf extract on belowground biomass might also be greater than that of root extract for other species, at least for their effects on root growth of the target species.

Evidence for Enhanced Mutualism Hypothesis: Solidago canadensis Plants from Regular Soils Perform Better

The important roles of plant-soil microbe interactions have been documented in exotic plant invasion, but we know very little about how soil mutualists enhance this process (i.e. enhanced mutualism hypothesis). To test this hypothesis we conducted two greenhouse experiments with Solidago canadensis (hereafter Solidago), an invasive forb from North America, and Stipa bungeana (hereafter Stipa), a native Chinese grass. In a germination experiment, we found soil microbes from the rhizospheres of Solidago and Stipa exhibited much stronger facilitative effects on emergence of Solidago than that of Stipa. In a growth and competition experiment, we found that soil microbes strongly facilitated Solidago to outgrow Stipa, and greatly increased the competitive effects of Solidago on Stipa but decreased the competitive effects of Stipa on Solidago. These findings from two experiments suggest that in situ soil microbes enhance the recruitment potential of Solidago and its ability to outcompete native plants, thereby providing strong evidence for the enhanced mutualism hypothesis. On the other hand, to some extent this outperformance of Solidago in the presence of soil microbes seems to be unbeneficial to control its rapid expansion, particularly in some ranges where this enhanced mutualism dominates over other mechanisms.

Cuscuta australis restrains three exotic invasive plants and benefits native species

In this study we conducted field investigations to examine the effects of native Cuscuta australis on three exotic invasive plants (i.e. Ipomoea cairica, Mikania micrantha, and Wedelia trilobata) and on the invaded native communities. The results showed that C. australis produced high infection rates on the exotic invasive hosts but low ones on the native species. Furthermore, the results showed that C. australis exhibited vigorous growth and high reproduction when it grew on M. micrantha and W. trilobata, indicating that these exotic invasive plants are more rewarding hosts than are native plants for C. australis. C. australis infection was positively related to the growth traits (e.g. biomass, cover, and total leaf area) and nutrient contents (e.g. N, P, and K) of the exotic invasive plants. The infections of C. australis significantly decreased the growth and nutrient contents of exotic invasive hosts, and the host–parasite interactions benefited the native species with increased species richness and biodiversity, facilitating the recovery of invaded native communities. This study provides a model for a native agent to both resist exotic invasive plants and benefit other native species. Furthermore, it indicates that certain native agents in invaded regions can be an effective and environmentally benign alternative to traditional biological control.

Is there a risk to living large? Large size correlates with reduced growth when stressed for knapweed populations

A central hypothesis in ecology is that plant life history evolution is constrained by fundamental “compromises between the conflicting selection pressures resulting from particular combinations of competition, stress, and disturbance”, with stress being defined as abiotic conditions that restrict production. Biogeographic differences among native and non-native ranges of invasive plants may provide unique opportunities for tests of this theory. We conducted a greenhouse experiment with Centaurea stoebe plants from North American and European populations. We compared the total biomass and phenotypic plasticity indices for plants from the native and non-native ranges under stressed and non-stressed conditions. The average size of Centaurea stoebe plants from 13 North American populations was greater than that of plants from 18 European populations regardless of stress treatment. However, when plants from the same populations were exposed to lower resources the differences in biomass between plants from North American and European populations were significantly less, suggesting that large plants were poorer stress tolerators. For all 31 populations the regression slope for the relationship between mean mass for populations in non-stressful conditions and mean mass in stressful conditions was less than 1.0, indicating that populations that produced large plants in good conditions also produced plants that grew disproportionately less in stressful conditions. These findings suggest that Centaurea stoebe may be evolving towards being a good “competitor” (sensu Grime, 1977) in its invasive range, but at the cost of being a good “stress tolerator”.

Are clonal plants more tolerant to grazing than co-occurring non-clonal plants in inland dunes?

Clonal traits such as clonal integration and storage functions of rhizomes or stolons may provide clonal plants with additional advantages against grazing over non-clonal plants. Here, we hypothesize that clonal species have a larger capacity for compensatory growth than co-occurring non-clonal species. In inland dunes in northern China, individual plants of two rhizomatous clonal species (Bromus ircutensis and Psammochloa villosa) and two non-clonal ones (Artemisia intramongolica and Astragalus melilotoides) were subjected to 0% (control), 50% (moderate) and 90% (heavy) shoot removal. Compared with control, heavy clipping greatly increased the relative growth rate in Bromus and Psammochloa, but decreased that in Artemisia and Astragalus. Heavy clipping affected above-ground dry weight and the number of modules more negatively in Artemisia and Astragalus than in Bromus and Psammochloa. These results support the hypothesis and suggest that clonal species are more tolerant to grazing than co-occurring non-clonal species in inland dunes.