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In shallow lake ecosystems, flooding is a key disturbance factor of aquatic vegetation. Aquatic plants, especially submerged plants, play key roles in water ecosystems. Liangzi Lake experienced severe flooding in July 2010, and the elevated water levels lasted for 3 months. In this study, 10 transects with 120 monitoring points were set up for monthly monitoring during the 3-year period, encompassing the period before and after the flooding (2009–2011). The numbers, biomass, and diversity of the submerged plants, as well as the physical and chemical characteristics of the lake water, were surveyed. There were 12 species belonging to 7 families and 7 genera in Liangzi Lake. Eleven of the submerged plant species were found in 2009, but, after the flood, that number decreased to five in 2011. The total biomass differed significantly over the three years (P < 0.05), with the largest biomass in 2009 and smallest in 2011. In 2009 and 2010, Potamogeton maackianus was the dominant species, but its dominant position weakened in 2011. After the flood, water transparency decreased, and the water depth, turbidity, total nitrogen, and total phosphorus increased. A redundancy analysis between the submerged plants and environmental factors found that the water transparency, turbidity, and water depth were the key environmental factors affecting the plants. These results suggest that the long-lasting severe flooding of Liangzi Lake in 2010 led to the degradation of both the submerged plant community and water quality.

When exotic species are introduced into new areas, establishment is a vital step in their invasion process. Therefore, overwintering strategies determine whether an exotic species from low latitudes can successfully invade middle- and high-latitude areas. In this study, we investigated the effects of nutrient and water drawdown on overwintering in an exotic aquatic plant from the tropical zone, Eichhornia crassipes, at the northern margin of its distribution in China. The population density, size of individuals, and the size and nitrogen concentration of overwintering organs (stem base) of E. crassipes that grew in high-nutrition water were greater than those that grew in low-nutrient water before winter. The overwinter survival rate of E. crassipes was significantly affected by the water level and nutrient. The thick and dense floating mat of E. crassipes can increase the temperature of water bodies; therefore, the overwinter survival rate of E. crassipes was higher in constant-water-level and high-nutrient treatment. In contrast, due to the loss of heat preservation provided by the floating mats and the low nitrogen concentration in the stem base, all individuals of E. crassipes died in constant-water-level and low-nutrient treatment. In the water-drawdown treatments, the stem base of E. crassipes was directly exposed to low-temperature air; therefore, the overwinter survival rate of E. crassipes was lower. Our results reveal that eutrophication can not only improve the competitiveness of E. crassipes but can also improve the survival rate of overwintering plants in temperate regions. Our study also suggests that removing nutrients from the water and regulating the water level can limit the invasion of E. crassipes in temperate and subtropical regions.

Spatial heterogeneity in soil nutrient availability is considered to play an important role in promoting plant invasion success and can affect interspecific competition. Although some clonal plants have been demonstrated to be correlated with resource heterogeneity in terrestrial systems, little is known about how soil nutrient heterogeneity affects the growth of invasive aquatic plants or their population structure. A greenhouse experiment was therefore conducted to study the response of the invasive aquatic plant Myriophyllum aquaticum to the spatial heterogeneity of soil nutrients under three plant densities (one, four, or twelve plants per container) with a constant amount of soil nutrients per container. The results showed that soil nutrient heterogeneity significantly increased the number of shoots in the single-plant density treatment. However, heterogeneous soil nutrient treatment significantly increased the number of shoots at the expense of total biomass and aboveground biomass in the twelve-plant density treatment. The heterogeneous soil nutrient treatment had few effects on other growth traits and intraspecific competition under different plant density treatments. These results indicate that spatial heterogeneity in soil nutrient availability may facilitate the spread of M. aquaticum.

The strategy of producing rapid initial growth and establishing early in the growing season is important, and it is employed by invasive macrophytes. Elodea nuttallii and Egeria densa, two Hydrocharitaceae species, became weeds after invading many countries in recent years. Comparative studies on their invasive traits in relation to native species during winter and spring are limited. In the present study, we compared the growth performance of these two exotic species with a perennial native species, Potamogeton maackianus, in different water depths (1, 2, and 3 m) during winter (January and February) and spring (March and April). Three morphological traits (shoot number, root number and shoot length), total biomass, relative growth rate (RGR) and two physiological photosynthetic traits (total chlorophyll content and the maximum quantum yield of PSII [Fv/Fm]) were measured for each macrophyte. All three species could overwinter as entirely leafy plants. Biomass, RGR, morphological traits and physiological traits were all different among species. However, water depths had a significant effect only on morphological traits. At all water depths, E. nuttallii had significantly higher values for morphological traits, total biomass and RGR than P. maackianus, while E. densa had significantly fewer roots and a lower total chlorophyll content than P. maackianus. Except for Fv/Fm at a 3 m water depth, morphological and physiological photosynthetic traits, biomass and RGR of E. nuttallii were significantly higher than those of E. densa. In addition, a large number of adventitious roots developed from E. nuttallii but not from the other two species. These results indicate that the advantages of E. nuttallii to grow in winter and spring may make it more prone to expansion than E. densa in China.

Submerged macrophytes and epiphytic algae play significant roles in the functioning of aquatic ecosystems. Submerged macrophytes can influence the epiphytic algal community by directly or indirectly modifying environmental conditions (nutrients, light, etc.). From December to June of the following year, we investigated the dynamics of the dominant winter species Potamogeton crispus, its epiphytic algae, and water quality parameters in the shallow Liangzi Lake in China. The richness of epiphytic algae had a trend similar to that of P. crispus coverage, which increased in the first four months and then decreased in the following three months. The structural equation model (SEM) showed that P. crispus affected the richness of epiphytic algae by reducing nutrient concentrations (reduction in total organic carbon, total nitrogen and chemical oxygen demand) and enhancing water transparency (reduction in turbidity and total suspend solids) to enhance the richness of epiphytic algae. The results indicated that high amounts of submerged macrophyte cover can increase the richness of the epiphytic algal community by changing water quality.

Clonal plants may face various types of resource heterogeneity in their natural habitats; as such, spatial or temporal resource heterogeneity can affect the growth of clonal plants. Clonal plants can concentrate their organs in a smaller area where resources are high would cause heterogeneity to increase competition between plants. Most studies on resource heterogeneity have investigated the response of plants under a single density or by manipulating a single resource. Few studies have tested the effects of the heterogeneous distribution of two covariable resources on plant growth and intraspecific competition. A greenhouse experiment was therefore conducted to study plant responses to the spatial and temporal heterogeneity of the soil and water supply under a variety of plant densities (one, two, four, or six plants per container). The perennial clonal herb Bolboschoenus yagara was grown under different combinations of water supply patterns, soil nutrient distribution types and plant densities while maintaining the total water and soil nutrient availability per container constant. Compared with that at a relatively high plant density, soil nutrient heterogeneity resulted in significantly less total plant biomass and less-modified morphological traits when the plant density is relative low. At the highest plant density, compared with the homogeneous soil treatments, the heterogeneous soil treatments significantly increased the total biomass and R/S ratio. Water supply patterns also clearly affected plant morphological traits at the highest plant density. Furthermore, soil heterogeneity significantly increased intraspecific competition intensity at low plant densities, but did not significantly affect intraspecific competition intensity at higher plant densities. Water heterogeneity had little impact on intraspecific competition. These results suggest that the growth performance and intraspecific competition of B. yagara are more strongly affected by soil nutrient distribution rather than by water supply patterns and that competition for soil nutrients may increase plant sensitivity to soil heterogeneity.

The strategy of producing rapid initial growth and establishing early in the growing season is important, and it is employed by invasive macrophytes. Elodea nuttallii and Egeria densa, two Hydrocharitaceae species, became weeds after invading many countries in recent years. Comparative studies on their invasive traits in relation to native species during winter and spring are limited. In the present study, we compared the growth performance of these two exotic species with a perennial native species, Potamogeton maackianus, in different water depths (1, 2, and 3 m) during winter (January and February) and spring (March and April). Three morphological traits (shoot number, root number and shoot length), total biomass, relative growth rate (RGR) and two physiological photosynthetic traits (total chlorophyll content and the maximum quantum yield of PSII [Fv/Fm]) were measured for each macrophyte. All three species could overwinter as entirely leafy plants. Biomass, RGR, morphological traits and physiological traits were all different among species. However, water depths had a significant effect only on morphological traits. At all water depths, E. nuttallii had significantly higher values for morphological traits, total biomass and RGR than P. maackianus, while E. densa had significantly fewer roots and a lower total chlorophyll content than P. maackianus. Except for Fv/Fm at a 3 m water depth, morphological and physiological photosynthetic traits, biomass and RGR of E. nuttallii were significantly higher than those of E. densa. In addition, a large number of adventitious roots developed from E. nuttallii but not from the other two species. These results indicate that the advantages of E. nuttallii to grow in winter and spring may make it more prone to expansion than E. densa in China.

Understanding the biotic and abiotic factors that influence the susceptibility of a community to invasion is beneficial for the prediction and management of invasive species and the conservation of native biodiversity. However, the relationships between factors and invasibility of a community have not been fully confirmed, and the factors most associated with the susceptibility of a community to invasion have rarely been identified. In this study, we investigated the species richness patterns in aquatic exotic and native plants and the relationships of exotic species richness with habitat and water environment factors in 262 aquatic plant communities in China. A total of 11 exotic plant species were recorded in our field survey, and we found neither a negative nor a positive relationship between aquatic exotic and native plant species richness. The aquatic exotic plant species richness is negatively correlated with the relative coverage and biomass of native plants but positively correlated with the total nitrogen (TN), total phosphorus (TP), and chemical oxygen demand (COD) concentrations in the water. The native plant species richness, native species' relative coverage, and native species' biomass were positively related to each other, whereas the TP, TN, and COD were also positively related to each other. The native plant species richness, native species' relative coverage, and native species biomass were each negatively correlated with the TP, TN, and COD. In addition, biotic rather than abiotic predictors accounted for most of the variation in exotic plant richness. Our results suggest that improving the vegetation coverage and the biodiversity of native plants is the most effective approach for preventing alien plant invasions and minimizing their impacts on freshwater ecosystems.