Interactive effects of light and snail herbivory rather than nutrient loading determine early establishment of submerged macrophytes

Submerged macrophytes play a key role in maintaining a clear‐water phase and promoting biodiversity in shallow aquatic ecosystems. Since their abundance has declined globally due to anthropogenic activities, it is important to include them in aquatic ecosystem restoration programs. Macrophytes establishment in early spring is crucial for the subsequent growth of other warm‐adapted macrophytes. However, factors affecting this early establishment of submerged macrophytes have not been fully explored yet. Here, we conducted an outdoor experiment from winter to early spring using the submerged macrophytes Potamogeton crispus and Vallisneria spinulosa to study the effects of shading, nutrient loading, snail herbivory (Radix swinhoei), and their interactions on the early growth and stoichiometric characteristics of macrophytes. The results show that the effects strongly depend on macrophyte species. Biomass and number of shoots of P. crispus decreased, and internode length increased during low light conditions, but were not affected by nutrient loading. P. crispus shoot biomass and number showed hump‐shaped responses to increased snail biomass under full light. In contrast, the biomass of the plant linearly decreased with snail biomass under low light. This indicates an interaction of light with snail herbivory. Since snails prefer grazing on periphyton over macrophytes, a low density of snails promoted growth of P. crispus by removing periphyton competition, while herbivory on the macrophyte increased during a high density of snails. The growth of V. spinulosa was not affected by any of the factors, probably because of growth limitation by low temperature. Our study demonstrates that the interaction of light with snail herbivory may affect establishment and growth of submerged macrophytes in early spring. Macrophyte restoration projects may thus benefit from lowering water levels to increase light availability and making smart use of cold‐adapted herbivores to reduce light competition with periphyton.     

螺类与着生藻类的相互作用及其对沉水植物的影响

浅水湖泊的富营养化常导致水生植被的退化与浮游藻类的爆发［１０,１８,２９］。可利用光通常是决定沉水植物分布、生物量和生产力的最重要因子,因此,伴随高营养负荷的浮游藻类繁殖,极大地削弱了沉水植物的光合能力［２０］。然而,Ｐｈｉｌｉｐｓ等人［２８］认为,...     

INTERACTIONS BETWEEN EPIPHYTES, MACROPHYTES AND FRESHWATER SNAILS: A REVIEW

Epiphyton-feeding snails are often a conspicuous feature ofthe invertebrate fauna associated with submerged freshwatermacrophytes. In this paper I review the different interactionstaking place between snails, epiphyton and macrophytes. Studies on grazing by freshwater snails show that snails havea great impact on the biomass, productivity and species compositionof epiphytic communities. Direct effects of grazing on livingmacrophytes are probably of minor importance, but snails havea significant indirect effect on macrophytes by reducing thedetrimental impact of epiphyton (e.g. shading and competitionfor nutrients). Predators of snails can have a mediating effecton snail-epiphyton-macrophyte interactions, both through a directpredatorprey relationship (reducing the density of snails) andby inducing a habitat displacement of the snails. In a studyon the effects of predation by the pumpkinseed sunfish (a specializedsnail predator) it was found that predation indirectly affectsthe biomass and species composition of epiphytic algae by regulatingthe density of snails.     

Rapid adaptive responses of rosette‐type macrophyte Vallisneria natans juveniles to varying water depths: The role of leaf trait plasticity

Rosette‐type submerged macrophytes are widely distributed across a range of water depths in shallow lakes and play a key role in maintaining ecosystem structures and functions. However, little is known about the rapid adaptive responses of such macrophytes to variations in water depth, especially at the juvenile stage. Here, we conducted a short‐term in situ mesocosm experiment, in which the juveniles of Vallisneria natans were exposed to a water depth gradient ranging from 20 to 360 cm. Twenty‐two leaf‐related traits were examined after 4 weeks of growth in a shallow lake. Most (18) traits of V. natans generally showed high plasticity in relation to water depth. Specifically, juveniles allocated more biomass to leaves and had higher specific leaf area, leaf length‐to‐width ratio, chlorophyll content, and carotenoids content in deep waters, displaying trait syndrome associated with high resource acquisition. In contrast, V. natans juveniles in shallow waters had higher leaf dry matter content, leaf soluble carbohydrate content, carotenoids per unit chlorophyll, and peroxidase activity, pertaining to resource conservation. Notably, underwater light intensity was found to be the key factor explaining the trait plasticity along the water depth gradient, and 1.30 mol photons m⁻² d⁻¹ (at 270 cm) could be the optimal irradiance level based on the total biomass of V. natans juveniles. The present study highlights the significance of leaf trait plasticity for rosette‐type macrophytes in response to variations in water depth and sheds new light on the differences between trade‐offs in deep‐ and shallow‐water areas.     

Low‐pH seawater alters indirect interactions in rocky‐shore tidepools

Ocean acidification is expected to degrade marine ecosystems, yet most studies focus on organismal‐level impacts rather than ecological perturbations. Field studies are especially sparse, particularly ones examining shifts in direct and indirect consumer interactions. Here we address such connections within tidepool communities of rocky shores, focusing on a three‐level food web involving the keystone sea star predator, Pisaster ochraceus, a common herbivorous snail, Tegula funebralis, and a macroalgal basal resource, Macrocystis pyrifera. We demonstrate that during nighttime low tides, experimentally manipulated declines in seawater pH suppress the anti‐predator behavior of snails, bolstering their grazing, and diminishing the top‐down influence of predators on basal resources. This attenuation of top‐down control is absent in pools maintained experimentally at higher pH. These findings suggest that as ocean acidification proceeds, shifts of behaviorally mediated links in food webs could change how cascading effects of predators manifest within marine communities.     

Effects of Spirogyra arcta on biomass and structure of submerged macrophyte communities

The presence of algae can greatly reduce the amount of light that reaches submerged macrophytes, but few experimental studies have been conducted to examine the effects of algae on biomass and structure of submerged macrophyte communities. We constructed communities with four submerged macrophytes (Hydrilla verticillata, Egeria densa, Ceratophyllum demersum, and Chara vulgaris) in three environments in which 0 (control), 50 and 100% of the water surface was covered by Spirogyra arcta. Compared to the control treatment, the 100% spirogyra treatment decreased biomass of the submerged macrophyte communities and of all the four macrophytes except C. demersum. Compared to the control and 50% treatments, the 100% treatment significantly increased relative abundance of C. demersum and decreased that of E. densa. Therefore, the presence of S. arcta can greatly affect the productivity and alter the structure of submerged macrophyte communities. To restore submerged macrophyte communities in conditions with abundant algae, assembling communities consisting of C. demersum or similar species may be a good practice.     

Effects of eutrophication and snails on Eurasian watermilfoil (Myriophyllum spicatum) invasion

Exotic species can invade and establish new habitats both as a result of their own traits, and as a result of the characteristics of the environment they invade. Here, we show that the abundance of the invasive submerged aquatic plant, Myriophyllum spicatum (Eurasian watermilfoil) is highly dependent on the conditions of the environment in a mesocosm experiment. M. spicatum is allelopathic towards epiphytic algae, and in the absence of algivorous snails, we found that the abundance of both algae and M. spicatum significantly increased with experimentally increased nutrient loading, while the abundance of native submerged macrophytes declined. However, when snails were present, snail biomass increased with increasing nutrient loading, and M. spicatum biomass was consistently low while native submerged macrophyte biomass was consistently high. Our results stress the importance of the interaction between species traits and environmental conditions when considering the invasiveness of certain exotic species and the invasibility of certain environments.     

不同初始螺类密度对沉水植物刺苦草Vallisneria spinulosa Yan及其附着藻类的影响

大型水生植物及其附着藻类是浅水湖泊中的重要初级生产者。淡水螺类作为重要的初级消费者,其密度对沉水植物及其附着藻类的影响存在争议。本研究设置4种初始螺类密度（0、40、80、240 ind·m^-2）,研究淡水螺类（椭圆萝卜螺Radix swinhoei H.Adams）对刺苦草（Vallisneria spinulosa Yan）及其附着藻类的直接牧食作用和螺类种群的变化。结果显示,在添加螺处理中,刺苦草和人工基质表面附着藻类的生物量显著降低,同时沉水植物的生长显著增加,在较高初始密度螺类处理中刺苦草产生更多的分株。到实验结束时,螺类的死亡率较高,但3个有螺处理间螺类鲜重无显著差异,而高初始密度螺类条件下的最终密度仍较高,同时个体重量（均重）也较小。在中富营养条件下淡水螺类可以直接牧食沉水植物叶片,但对植物生长的抑制作用不显著,有可能是因为沉水植物并不能作为唯一的食物来源维持螺类种群,同时螺类的种群结构受到水体营养水平等因素的制约。     

Impact of invasive apple snails in Hong Kong on wetland macrophytes,nutrients, phytoplankton and filamentous algae

1. Grazing by invasive species can affect many aspects of an aquatic system, but most studies have focused on the direct effects on plants. We conducted mesocosm and laboratory experiments to examine the impact of the invasive apple snail Pomacea canaliculata on macrophytes, filamentous algae, nutrients and phytoplankton. 2. In a freshwater pond, we confined 500 g of Myriophyllum aquaticum or Eichhornia crassipes with 0, 2, 4 or 8 apple snails in 1 m × 1 m × 1 m enclosures for approximately 1 month. Apple snails grazed heavily on both species of macrophytes, with higher overall weight losses at higher snail densities. The damage patterns differed between the two macrophytes. In M. aquaticum, both leaves and stems suffered from substantial herbivory, whereas in E. crassipes, only the roots suffered significant weight reduction. 3. In addition to grazing on macrophytes, apple snails appeared to have controlled the growth of filamentous algae, as these did not develop in the snail treatments. The ability of P. canaliculata to control filamentous algae was supported by a laboratory experiment where the consumption was as high as 0.25 g g⁻¹ snail DW d⁻¹. Because of a lack of native herbivorous snails in the pond, the growth of filamentous algae (mainly Spirogyra sp.) reached 80.3 g m⁻², forming a spongy pond scum in the no‐apple snail control. Together with previous reports that apple snails could eat the juveniles and eggs of other freshwater snails, our results indicated that P. canaliculata could have out‐competed native herbivorous snails from the pond by predation on their juveniles or eggs. Alternatively, P. canaliculata might have out‐competed them by monopolisation of food resources. 4. Nitrogen and phosphorous concentrations remained low throughout both experiments and were not correlated with apple snail density. The treatment effects on chlorophyll a (Chl a) and phytoplankton composition varied in the two experiments. In the M. aquaticum experiment, with increasing snail density, Chl a increased, and the phytoplankton community became dominated by Cryptophyceae. In the E. crassipes experiment, Chl a level was independent of snail density, but with increasing snail density, the phytoplankton community became co‐dominated by Cryptophyceae, Chlorophyceae and Bacillariophyceae. 5. Given the multiple effects of P. canaliculata on wetland biodiversity and function, management strategies should be developed to prevent its further spread. In invaded wetlands, strategies should be developed to eradicate the apple snail and re‐introduce native snails which can control the development of filamentous algae.     

Factors influencing the stability of nutrient-enriched freshwater macrophyte communities: the role of sticklebacks Pungitius pungitius and freshwater snails

1. Enclosures, with the following characteristics, were used to investigate the effects of increased nutrient (nitrate and phosphate) loading on freshwater communities: (a) controls with normal densities of invertebrates, and including macrophytes, molluscs and fish (sticklebacks), and communities similar to the controls but with (b) much reduced snail densities and (c) increased fish densities. 2. The addition of nutrients resulted in increases in the biomass of phytoplankton and epiphyton, as well as zooplankton counts, in all the experimental enclosures. 3. A reduction in the snail population density resulted in a significant decrease in phytoplankton density and an increase, compared to controls, in the density of epiphytic algae growing on the leaves of the submerged macrophytes, Potamogeton crispus and Groenlandia densa. It also resulted in a significant increase (3.5 times that in the controls) in chlorophyll-a values of epilithic algae growing on glass slides, an increase in senescent tissue in macrophytes and a decline in the mass of Groenlandia densa by the end of the experiment. 4. An increase in the fish population resulted in a significant decline, compared to the controls, in the densities of zooplankton (including Chydorus sphaericus and copepods), Lymnaea peregra (from May to mid-June), Asellus meridianus, Crangonyx pseudagracilis, Culex (P.) molestus and Physa fontinalis. It also resulted in a change in the composition of the phytoplankton, with an increase in the large colonial Eudorina elegans and a decline in smaller planktonic algae, including Ankistrodesmus and Cryptomonas spp. 5. The possible mechanisms responsible for the effects are evaluated and their relevance to conservation and management of freshwater macrophyte communities are discussed.     

Predatory cues drive colony size reduction in marine diatoms

1. Colony formation is a common feature among nonmotile marine phytoplankton. Several theories exist around the potential benefits of larger colonies.
2. Here, we test the hypothesis that predation is one of the drivers behind colony formation and chain length plasticity. We exposed cultures of Thalassiosira rotula, Chaetoceros curvisetus, and Chaetoceros affinis to copepodamides, a chemical alarm cue released by copepods and perceived as an indicator of predation threat by their prey. This was coupled with a grazing experiment, which compared copepod grazing rates on different chain lengths.
3. Our results show that T. rotula and C. curvisetus decreased their chain lengths by 79% and 49%, respectively, in response to copepodamides. Single cells and short chains were grazed at lower rates compared with long chains, and the copepodamide‐driven size shift led to 30% and 12% lower grazing in T. rotula and C. curvisetus, respectively. In contrast, C. affinis showed a slight increased chain length in response to copepodamides although nonsignificant.
4. We found that 2 of 3 studied species reduce their chain length in response to the presence of copepod grazers. Altered size structure has implications for the route of carbon in the marine food webs and carbon export to deeper strata.

     

An evaluation of the interactions between freshwater pulmonate snail hosts of human schistosomes and macrophytes

An account is given of a laboratory investigation designed to evaluate the extent to which the freshwater pulmonate snail Biomphalaria glabrata (Say) can utilize various species of aquatic plants, mainly macrophytes, when presented in the following forms over different time scales: normal plants; dried plant material; homogenized plant material in calcium alginate matrices; water-soluble filtrates of plant homogenates in the medium. The following propositions, derived from the theory of phased coevolution of components of the module consisting of the epiphytic bacteria, algae, snails and macrophytes, are evaluated on the basis of the present results and others including those obtained in this laboratory. That as the snails had become specialized to exploit surface communities of epiphytic algae, decaying plant material and dissolved organic matter (DOM) early in their evolutionary history they would continue to exploit these resources when they later become associated with aquatic macrophytes. That pulmonate snails would tend to be feeding generalists capable of adapting to food of varying chemical composition, given sufficient time, provided it was sufficiently small or flaccid. That although macrophytes and snails show a strong positive relationship, the living macrophyte tissue would be little used by the snails. That the hard outer envelope, inherited from their terrestrial ancestors, would remain as the major defence mechanism of aquatic macrophytes against attack by snails and other aquatic invertebrates. That aquatic macrophytes would invest little in the nutrient deficiency strategy to reduce attack by invertebrates such as snails. That truly aquatic submerged macrophytes would not possess secondary plant compounds (SPC) that would be molluscicidal. Emergent parts of subaquatic or aquatic plants might be expected to be better sources of SPC with molluscicidal factors than submerged aquatic plants. Species of epiphytic or planktonic algae might be better sources of SPC with molluscicidal effects than aquatic macrophytes. That the strategies developed by pulmonate snails for obtaining their energy supplies would not be conducive to rapid speciation. The analysis of the present and other related results supports these propositions. Predictions based on the theory of mutualism involving the snails, macrophytes and other components of the module also receive some support from an analysis of the present results. The additional empirical work that could be undertaken to test this theory is briefly discussed.(ABSTRACT TRUNCATED AT 400 WORDS)     

Do Amplitudes of Water Level Fluctuations Affect the Growth and Community Structure of Submerged Macrophytes?

Submerged macrophytes are subjected to potential mechanical stresses associated with fluctuating water levels in natural conditions. However, few experimental studies have been conducted to further understand the effects of water level fluctuating amplitude on submerged macrophyte species and their assemblages or communities. We designed a controlled experiment to investigate the responses of three submerged macrophyte species (Hydrilla verticillata, Ceratophyllum demersum and Elodea nuttallii) and their combinations in communities to three amplitudes (static, ± 30 cm, ± 60 cm) of water level fluctuations. Results showed that water level fluctuating amplitude had little effects on the community performance and the three tested species responded differently. H. verticillata exhibited more growth in static water and it was negatively affected by either of the water level fluctuations amplitude, however, growth parameters of H. verticillata in two fluctuating water level treatments (i.e., ± 30 cm, ± 60 cm) were not significantly different. On the other hand, the growth of C. demersum was not significantly correlated with different amplitude treatments. However, it became more abundant when water levels fluctuated. E. nuttallii was inhibited by the two fluctuating water level treatments, and was less in growth parameters compared to the other species especially in water level fluctuating conditions. The inherent differences in the adaptive capabilities of the tested species indicate that C. demersum or other species with similar responses may be dominant species to restore submerged macrophyte communities with great fluctuating water levels. Otherwise, H. verticillata, E. nuttallii or other species with similar responses could be considered for constructing the community in static water conditions.     

Epiphyte shading: Its role in resulting depth distribution of submerged aquatic macrophytes

The biomass of epiphytic algae increased more than the biomass of phytoplankton with increased nutrient availability (N, P, Si) in four lakes. Large epiphyte biomasses on the submerged macrophyte,Littorella uniflora, reduce light attenuation more than phytoplankton did. High values of shading by epiphytes correspond with a reduction in the maximum depth penetration ofLittorella by more than 1 meter.     

Epiphytic algae and macroinvertebrates on submerged and floating-leaved macrophytes in an Italian lake

1. The presence of contiguous beds of submerged (Myriophyllum spicatum, Ceratophyllum demersum and Najas marina) and floating-leaved (Trapa natans) vegetation in a north Italian lake allowed us to test the effect of the different host architecture on epiphytic algae and invertebrates and to predict the consequences for the lake of changes in the predominant vegetation. 2. Epiphyton development, measured as carbon, nitrogen, phosphorus, chlorophyll a (Chl a), phaeophytin and as algal and macroinvertebrate density, was significantly higher on submerged plants than on T. natans. The C : Chl a ratio, a proxy of the ratio of heterotrophs to autotrophs, was higher on the floating-leaved plants. The elemental (C : N : P) and pigment (Chl a : phaeophytin) ratios were not significantly different between the two vegetation types. 3. The taxonomic composition of epiphytic algae and invertebrates was similar on the different plants. The more varied morphology of the floating-leaved T. natans resulted in a higher diversity of epiphytic algae, however, but not of macroinvertebrates. 4. There was a significant inverse relationship between epiphyton biomass and the standing crop of the host plant, suggesting a key role for light and water exchange in epiphyton development. 5. Replacement of floating-leaved by submerged plants would increase the total biomass of epiphytic algae and invertebrates.     

Impacts of sediment type on the performance and composition of submerged macrophyte communities

To restore deteriorated lake ecosystems, it is important to identify environmental factors that influence submerged macrophyte communities. While sediment is a critical environmental factor for submerged macrophytes and many studies have examined effects of sediment type on the growth of individual submerged macrophytes, very few have tested how sediment type affects the growth and species composition of submerged macrophyte communities. We constructed submerged macrophyte communities containing four co-occurring submerged macrophytes (Hydrilla verticillata, Myriophyllum spicatum, Ceratophyllum demersum and Chara fragilis) and subjected them to three sediment treatments, i.e., clay, a mixture of clay and quartz sand at a volume ratio of 1:1 and a mixture at a volume ratio of 1:4. Compared to the clay, the 1:1 mixture treatment greatly increased overall biomass, number of shoot nodes and shoot length of the community, but decreased its diversity. This was because it substantially promoted the growth of H. verticillata within the community, making it the most abundant species in the mixture sediment, but decreased that of M. spicatum and C. demersum. The sediment type had no significant effects on the growth of C. fragilis. As a primary nutrient source for plant growth, sediment type can have differential effects on various submerged macrophyte species and 1:1 mixture treatment could enhance the performance of the communities, increasing the overall biomass, number of shoot nodes and shoot length by 39.03%, 150.13% and 9.94%, respectively, compared to the clay treatment. Thus, measures should be taken to mediate the sediment condition to restore submerged macrophyte communities with different dominant species.     

Biological control of phytoplankton by the subtropical submerged macrophytes Egeria densa and Potamogeton illinoensis: a mesocosm study

1. In temperate regions, submerged macrophytes can hamper phytoplankton blooms. Such an effect could arise directly, for instance via allelopathy, or indirectly, via competition for nutrients or the positive interaction between submerged macrophytes and zooplankton grazing. However, there is some evidence that the positive interaction between submerged macrophytes and zooplankton grazing is less marked in warmer regions, where the interaction is less well studied, and that negative effects of higher water plants on phytoplankton biomass are weaker. 2. We carried out two consecutive mesocosm experiments in Uruguay (subtropical South America) to study the effects of two common submerged macrophytes from this region (Egeria densa and Potamogeton illinoensis) on phytoplankton biomass, in the absence of zooplankton grazing. We compared phytoplankton development between different macrophyte treatments (no macrophytes, artificial macrophytes, real Egeria and real Potamogeton). We used artificial macrophytes to differentiate between physical effects (i.e. shading, sedimentation and competition with periphyton) and biological effects (i.e. nutrient competition and allelopathy). 3. In Experiment 1, we found no evidence for physical effects of macrophytes on phytoplankton biomass, but both macrophyte species seemed to exert strong biological effects on phytoplankton biomass. Only Egeria affected phytoplankton community structure, particularly tempering the dominance of Scenedesmus. Nutrient addition assays revealed that only Egeria suppressed phytoplankton through nutrient competition. 4. We performed a second mesocosm experiment with the same design, but applying saturating nutrient conditions as a way of excluding the effects of competition for nutrients. This experiment showed that both macrophytes were still able to suppress phytoplankton through biological mechanisms, providing evidence for allelopathic effects. Our results indicate that both common macrophytes are able to keep phytoplankton biomass low, even in the absence of zooplankton grazing.     

Allelopathic inhibition of epiphytes by submerged macrophytes

The hypothesis that epiphytes are more vulnerable to allelochemicals released by submerged macrophytes than phytoplankton was tested by measuring growth and photosystem (PS) II activity of three common epiphytic algae and cyanobacteria in coexistence with Myriophyllum spicatum using dialysis tubes. Results were compared with earlier experiments on planktonic species. Contrary to the planktonic species, the tested epiphytes, the green algae Stigeoclonium tenue, the diatom Gomphonema parvulum and the cyanobacterium Oscillatoria limosa, were not significantly inhibited by M. spicatum. Growth and PS II activity of O. limosa were even significantly enhanced by M. spicatum, but this effect disappeared under phosphorus-deficiency due to the allelopathically induced inhibition of the alkaline phosphatase activity or phosphorus leakage by the macrophytes. My findings of a lower vulnerability of epiphytes against allelopathic substances of submerged macrophytes are supported by results of a literature survey.     

Snail herbivory on submerged macrophytes and nutrient release: Implications for macrophyte management

Radix swinhoei (H. Adams) is a freshwater snail commonly found in shallow regions of Lake Taihu. This research estimated, based on experiments, the consumption rates of R. swinhoei on three young submerged plants (Vallisneria spiralis, Hydrilla verticillata and Potamogeton malaianus) and its rates of nutrient release. Results showed that the snails consumed V. spiralis at the highest rate (23.34 mg g⁻¹ d⁻¹), P. malaianus at a lower rate (11.97 mg g⁻¹ d⁻¹), and H. verticillata at the lowest rate (7.04 mg g⁻¹ d⁻¹). The consumption rates on V. spiralis varied significantly, with snail size, ranging from 13.63 mg g⁻¹ d⁻¹ for large-size snails to 143.42 mg g⁻¹ d⁻¹ for small-size ones.The average nutrient release rates of snails grazing on different macrophytes were 45.93 μg PO₄-P and 0.58 mg NH₄-N g⁻¹ d⁻¹. The food species had a significant effect on NH₄-N release rates but not on PO₄-P. However, the snail size had a significant effect on PO₄-P release rates and not on NH₄-N. The present study indicates that through selective grazing and nutrient release, snails may impose a significant impact on the macrophyte community, which should be considered in managing the macrophytes of a lake.     

Responses of four submerged macrophytes to freshwater snail density (Radix swinhoei) under clear‐water conditions: A mesocosm study

Macrophytes play a key role in stabilizing clear‐water conditions in shallow freshwater ecosystems. Their populations are maintained by a balance between plant grazing and plant growth. As a freshwater snail commonly found in shallow lakes, Radix swinhoei can affect the growth of submerged macrophytes by removing epiphyton from the surface of aquatic plants and by grazing directly on macrophyte organs. Thus, we conducted a long‐term (11‐month) experiment to explore the effects of snail density on macrophytes with distinctive structures in an outdoor clear‐water mesocosm system (with relatively low total nitrogen (TN, 0.66 ± 0.27 mg/L) and total phosphorus (TP, 36 ± 20 μg/L) and a phytoplankton chlorophyll a (Chla) range of 14.8 ± 4.9 μg/L) based on two different snail densities (low and high) and four macrophyte species treatments (Myriophyllum spicatum, Potamogeton wrightii, P. crispus, and P. oxyphyllus). In the high‐density treatment, snail biomass and abundance (36.5 ± 16.5 g/m² and 169 ± 92 ind/m², respectively) were approximately twice that observed in the low‐density treatment, resulting in lower total and aboveground biomass and ramet number in the macrophytes. In addition, plant height and plant volume inhabited (PVI) showed species‐specific responses to snail densities, that is, the height of P. oxyphyllus and PVI of M. spicatum were both higher under low‐density treatment. Thus, compared with low‐density treatment, the inhibitory effects of long‐term high snail density on macrophytes by direct feeding may be greater than the positive effects resulting from epiphyton clearance when under clear‐water conditions with low epiphyton biomass. Thus, under clear‐water conditions, the growth and community composition of submerged macrophytes could be potentially modified by the manual addition of invertebrates (i.e., snails) to lakes if the inhibitory effects from predatory fish are minor.