Impacts of water depth and substrate type on Vallisneria natans at wave-exposed and sheltered sites in a eutrophic large lake

Water depth, substrate type and wave exposure are major environmental factors influencing the growth and distribution of submerged macrophytes. However, we lack knowledge on the ecological reactions of submerged macrophytes to those factors during an entire growth period in a eutrophic lake with a Secchi depth of less than 50cm. We investigated the interactive effects of water depth (60, 120 and 200cm), waves (exposed vs. sheltered) and substrate nutrient level (fertile sludge vs. brown clay) on the survival, growth and morphology of Vallisneria natans using a new type of rhizotron in a 252-day field experiment. Plant length, leaf number, ramet number, root length, root number and biomass generally decreased with increased water depth at both wave conditions and in both substrates. When exposed to wave, the biomass of V. natans in water at 200cm depth rapidly declined in both substrates. When sheltered to wave, aboveground biomass at 60cm water depth first declined to zero with many buried rhizomes remained. The tuber germination rate decreased with increasing water depth during the second year of germination. No plants sprouted at 200cm water depth except at the case of exposed wave and sludge substrate. Ramet number was influenced most by water depth, followed by aboveground biomass, leaf number, plant length, and survival rate. Wave shelter and brown clay enhanced the impacting strength of the water depth. Wave exposure exhibited no negative influence on survival percentage, whereas sludge had a positive influence on plant survival in deep water after overwintering. Waves negatively affected the rate of increase of plant growth in sludge but positively in clay. The positive joint effects of wave exposure and low-nutrient substrate were mainly on plant length and biomass. Water depth had a negative influence that predominated over substrate and waves regarding plant survival and growth. Moreover, this negative influence may be aggravated by an increasing risk of submersed macrophyte loss caused by decreased belowground growth under high water levels, high wave exposure and high-nutrient conditions. Decreasing the water depth could be an useful measure for submerged macrophyte restoration in freshwater habitats, even in turbid eutrophic water.