Snail communities increase submerged macrophyte growth by grazing epiphytic algae and phytoplankton in a mesocosm experiment

The relationships between producers (e.g., macrophytes, phytoplankton and epiphytic algae) and snails play an important role in maintaining the function and stability of shallow ecosystems. Complex relationships exist among macrophytes, epiphytic algae, phytoplankton, and snails. We studied the effects of snail communities (consisting of Radix swinhoei, Hippeutis cantori, Bellamya aeruginosa, and Parafossarulus striatulus) on the biomass of phytoplankton and epiphytic algae as well as on the growth of three species of submerged macrophytes (Hydrilla verticillata, Vallisneria natans, and one exotic submerged plant, Elodea nuttallii) in a 90‐day outdoor mesocosm experiment conducted on the shore of subtropical Lake Liangzihu, China. A structural equation model showed that the snail communities affected the submerged macrophytes by grazing phytoplankton and epiphytic algae (reduction in phytoplankton Chl‐a and epiphytic algal abundance), enhancing the biomass of submerged macrophytes. Highly branched macrophytes with high surfaces and morphologies and many microhabitats supported the most snails and epiphytic algae (the biomass of the snail communities and epiphytic algae on H. verticillata was greater than that on V. natans), and snails preferred to feed on native plants. Competition drove the snails to change their grazing preferences to achieve coexistence.     

Implications of rapid changes in chlorophyll-<Emphasis Type="Italic">a</Emphasis> of plankton,epipelon, and epiphyton in a Pampean shallow lake: an interpretation in terms of a conceptual model

Biomass assessments of algae in wetlands usually include only the phytoplankton community without considering the contribution of other algal associations to total algal biomass. This omission prevents an accurate evaluation of the phytoplankton community as an integral part of the total ecosystem. In the present work, the biomass contributions (expressed as chlorophyll-a content per m² of lake) of phytoplankton, epiphyton on both submerged and emergent macrophytes, and epipelon were measured in Lacombe Lake, Argentina, for the purpose of (1) establishing the relative importance of the phytoplankton and (2) evaluating the entire contribution of algal biomass within the context of the Goldsborough & Robinson conceptual model. Our sampling was carried out monthly for a year in sites representative of different conditions with respect to water depth and type of macrophytes. Physicochemical analyses of water were performed following standard methods. Plankton was collected in a five-level profile at deeper stations and in subsurface samples at the shallow one. Samples of sediment obtained with corers were collected for epipelon sampling and segments of plants were cut at different levels, so as to obtain the epiphytes by scraping. Pigment was extracted with aqueous acetone and calculations were made by means of the Lorenzen equation. According to the Goldsborough & Robinson model, a Lake State developed here during the winter (phytoplankton maxima: 150 mg chlorophyll-a per m²). Then, through the subsequent growth of the submerged macrophytes, an Open State was observed, characterized by a maximum epiphyton biomass (at 3,502 mg chlorophyll-a per m²) along with lower levels of phytoplankton biomass. The epiphytic algae on the emergent macrophytes were always present but attained only relatively low biomass values (maximum: 120 mg of chlorophyll-a per m² in February). The epipelon biomass varied between 50 and 252 mg chlorophyll-a per m², registering a considerable contribution of settled algae from the water column (phytoplankton). This study contributes to our knowledge of wetland dynamics through its assessment of the rapid changes in the relative contributions of both planktonic and attached algae to the total algal biomass within the context of specific environmental factors. Guest editors: U. M. Azeiteiro, I. Jenkinson & M. J. Pereira Plankton Studies     

Wave exposure related growth of epiphyton: implications for the distribution of submerged macrophytes in eutrophic lakes

The distribution of submerged macrophytes in eutrophic lakes has been found to be skewed towards sites with intermediate exposure to waves. Low submerged macrophyte biomass at exposed sites has been explained by, for instance, physical damage from waves. The aim of this study was to investigate if lower biomass at sheltered sites compared to sites with intermediate exposure to waves can be caused by competition from epiphyton.Investigations were performed in eutrophic lakes in southern Sweden. Samples of submerged macrophytes and epiphytic algae on the macrophytes were taken along a wave exposure gradient. The amount of epiphyton (AFDW) per macrophyte biomass decreased with increased exposure. Biomass of submerged macrophytes, on the other hand, increased with increased exposure until a relatively abrupt disappearance of submerged vegetation occurred at high exposures. Production of epiphytic algae was monitored on artificial substrates from June to September at a sheltered and an exposed site in three lakes. It was higher at sheltered sites compared with exposed sites.We suggest that epiphytic algae may be an important factor in limiting the distribution of submerged macrophytes at sheltered sites in eutrophic lakes.     

Structural and functional characteristics of epiphyton and epipelon in relation to their distribution in Lake Vechten

Epiphyton and epipelon were quantitatively collected, respectively, from the submerged macrophytes and the sandy lake bottom of Lake Vechten (The Netherlands). On a weight basis, epiphyton was maximal in autumn and epipelon in summer. In winter the chemical composition of epiphyton and epipelon was similar. In summer the epiphyton had on a unit weight basis more organic matter and carbonate, and had per unit organic matter a higher algal number, nitrogen and energy content than the epipelon. Algae predominating the epiphyton were filamentous greens and pennate diatoms; those in the epipelon were pennate diatoms and blue-green algae. In both cases, species known to frequent the phytoplankton were abundant. The diatoms were quantified using paper chromatographic pigment analyses. Both the epiphyton and the epipelon exhibited maximal photosynthesis in mid summer. That light was generally the limiting factor was evident from periphyton developed on artificial substrates. This periphyton differed widely in its composition from that on the natural substrates, mainly because the latter collected much more sedimenting matter.In dense Ceratophyllum stands light was severely attenuated and the significant gradients in oxygen and pH were caused by the differences with depth in the proportions of photosynthesis and respiration. The oxygen content and pH at the bottom decreased owing to epipelic respiration. The epiphytic composition depended greatly on the degree of light attenuation. The epiphytic and epipelic respiration, except during part of the early summer, exceeded photosynthesis on a 24 h basis; this included the macrophytic photosynthesis during the time the vegetation was maximally developed. During the growing season import of organic matter, i.e. deposited seston, greatly exceeded that due to the photosynthetic production. After the summer maximum, the epipelon decreased faster than predicted from its oxygen exchange. It was concluded that sedimentation and resuspension determined mainly the changes in epiphyton and epipelon. Especially when covered with vegetation, the lower littoral of Lake Vechten plays a large part in the aerobic decomposition of sestonic organic matter.     

Universal chlorophyll equations for estimating chlorophylls <Emphasis Type="Italic">a,b, c</Emphasis>, and <Emphasis Type="Italic">d</Emphasis> and total chlorophylls in natural assemblages of photosynthetic organisms using acetone,methanol, or ethanol solvents

A universal set of equations for determining chlorophyll (Chl) a, accessory Chl b, c, and d, and total Chl have been developed for 90 % acetone, 100 % methanol, and ethanol solvents suitable for estimating Chl in extracts from natural assemblages of algae. The presence of phaeophytin (Ph) a not only interferes with estimates of Chl a but also with Chl b and c determinations. The universal algorithms can hence be misleading if used on natural collections containing large amounts of Ph. The methanol algorithms are severely affected by the presence of Ph and so are not recommended. The algorithms were tested on representative mixtures of Chls prepared from extracts of algae with known Chl composition. The limits of detection (and inherent error, ±95 % confidence limit) for all the Chl equations were less than 0.03 g m⁻³. The algorithms are both accurate and precise for Chl a and d but less accurate for Chl b and c. With caution the algorithms can be used to calculate a Chl profile of natural assemblages of algae. The relative error of measurements of Chls increases hyperbolically in diluted extracts. For safety reasons, efficient extraction of Chls and the convenience of being able to use polystyrene cuvettes, the algorithms for ethanol are recommended for routine assays of Chls in natural assemblages of aquatic plants.     

Effects of different macrophyte growth forms on sediment and P resuspension in a shallow lake

The effects of floating-leaved, submerged and emergent macrophytes on sediment resuspension and internal phosphorus loading were studied in the shallow Kirkkojärvi basin by placing sedimentation traps among different plant beds and adjacent open water and by sediment and water samples. All the three life forms considerably reduced sediment resuspension compared with non-vegetated areas. Both among submerged (Ceratophyllum demersum, Potamogeton obtusifolius, Ranunculus circinatus) and emergent (Typha angustifolia) plants, resuspension rate was on average 43% of that in the adjacent open water, while within floating-leaved plants (Nuphar lutea) the corresponding value was 87%. The effects of submerged and emergent vegetation increased in the course of the growing season together with increasing plant density. Among floating-leaved vegetation, such seasonal trend in resuspension effects was not observed. Compared with the non-vegetated area, floating-leaved, submerged and emergent plants reduced internal phosphorus loading on average by 21, 12 and 26 mg m⁻² d⁻¹, respectively. The effects of floating-leaved plants on resuspension-mediated internal phosphosrus loading were thus comparable to the effects of the other two life forms.     

Temporal changes and vertical distribution of macrophytes in Lake Kawaguchi

 The distribution of macrophytes in Lake Kawaguchi, Japan, was surveyed in August 1999 using a sampling anchor from a boat. The survey revealed that the present aquatic vegetation was composed of 17 submerged species, and no floating-leaved plants were present. The diversity of submerged plants and their vertical growth limits decreased progressively from the west to the east end of the lake. At the time of the survey, the dominant species was Elodea nuttallii (Planch.) St. John, which had once grown explosively in the mid-1980s in the lake, and except for Vallisneria asiatica Miki, all of the species commonly observed in the lake seem to have diminished their growth greatly. Among them, Potamogeton compressus L., which was the most dominant species in the lake before the invasion of Elodea nuttallii, had decreased most severely. It is suggested that the present state of aquatic vegetation in this lake was much influenced by the persistent growth of Elodea nuttallii as well as by environmental factors. Received: August 7, 2001 / Accepted: February 9, 2002     

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.     

不同光照强度下物种组合对沉水植物苦草种间关系的影响

为了解种间关系对沉水植物群落结构的影响，在不同光照(20%自然光和50%自然光)和不同物种组合下，研究了长江中下游常见优势沉水植物苦草(Vallisneria natans)与黑藻(Hydrilla verticillata)和穗状狐尾藻(Myriophyllum spicatum)的相互作用。结果表明，在低光下，苦草与穗状狐尾藻混种时，苦草生物量、株高和叶数均没有明显变化，当穗状狐尾藻的比例较高时，苦草根长生长受到抑制，根叶比呈下降趋势；在高光下，穗状狐尾藻比例的增加会促进苦草单株生物量和叶生物量的增加，而对苦草株高、根长和叶数无显著影响；与黑藻混种相比，苦草与穗状狐尾藻混种时，苦草的株高、根长和叶数均无显著差异，而苦草的单株生物量和叶生物量均呈降低趋势。因此，物种组合和混种比例均会影响苦草与其他物种的相互作用关系，进而影响沉水植被的群落动态。     

Sediment type and the clonal size greatly affect the asexual reproduction,productivity, and nutrient absorption of Vallisneria natans

Most aquatic vegetation restorations involve the transplantation of submerged macrophytes. Sediment type and the clonal size are of great significance as they determine the fate of submerged macrophytes. In order to ensure successful restoration, a simulation experiment was conducted using aquarium mesocosms to investigate the response of stolon propagation capacity, the morphological features and productivity of Vallisneria natans for four types of sediment (lake mud [L], lake mud + sand [L + S, 50:50, v/v mixture], sand [S], clay [C]), and three types of clonal sizes. Results showed that sediment types significantly affected V. natans biomass accumulation, stolon propagation ability, ramet morphological characteristics, and productivity, where the asexual reproduction ability and productivity ranked as L > L + S > S > C in four sediment types. Total biomass, maximum net production, number of ramets, root diameter, number of stolons, and stolon propagation rate were all highest in L. In L and L + S, the plant chlorophyll content was higher than in S and C. The root diameter and the ratio of aboveground/underground biomass in S were the smallest among the four sediments. Moreover, when more V. natans seedlings were linked, more ramets and biomass were produced. The stolon propagation rate was ranked as the stolon with single seedling greater than the stolon with two‐linked seedlings greater than the stolon with three‐linked seedlings in L and L + S. The concentration of total nitrogen, total phosphorus, and NO₃^?‐N in water was remarkably reduced in four aquariums. Findings provide a scientific basis for restoring submerged macrophytes in different sediment settings.     

Influences of habitat complexity on the diversity and abundance of epiphytic invertebrates on plants

SUMMARY 1. The compound influence of habitat complexity and patch size on stream invertebrate assemblages associated with submerged macrophytes was investigated through field sampling of two natural macrophyte species with contrasting leaf morphologies (complex, Ranunculus yezoensis; simple, Sparganium emersum) and an experiment with two artificial plants with different levels of morphological complexity. 2. The artificial plant experiment was designed to separate the effects of habitat area (patch size) and habitat complexity, thus enabling a more rigorous assessment of complexity per se than in previous studies where only a single patch size was used. Simple and complex artificial plants were established with five different patch sizes corresponding to the range found in natural plants. 3. Invertebrates occurred on both complex and simple forms of natural and artificial plants at similar abundances with dipterans and ephemeropterans being predominant. Taxon richness was higher on structurally complex Ranunculus than on simple Sparganium and was similarly higher on the complex artificial plant than on the simple one, over the entire range of habitat patch sizes. Thus, architectural complexity affected the taxon richness of epiphytic invertebrates, independently of habitat scale. 4. On the natural plants there was no difference in the abundance (both number of individuals and biomass) of invertebrates between simple and complex forms, while on artificial plants more invertebrates occurred on complex than on simple forms. The amount of particulate organic matter, >225 μm (POM) and chlorophyll a showed mixed patterns on natural and artificial plants, suggesting that the availability of these resources is not an overriding proximate factor controlling invertebrate abundance on plants. The difficulty of extrapolating from experimental results involving use of artificial plants is discussed, especially when considering the relationship between habitat structure and the occurrence of epiphytic invertebrates on natural plants.     

A study on annual changes in surface cover of floating-leaved plants in a lake using aerial photography

A simple method of aerial photography was used to investigate annual and between-year changes in surface cover of the floating-leaved plants, Nymphoides peltata, Nelumbo nucifera and Trapa natans, in natural stands in Lake Kasumigaura. Oblique photographs of vegetation were taken with a 35-mm camera with real color film from airplanes between 1983 and 1989. Vegetation maps were traced on the positions after projective transformation to vertical photography. The distribution of the stand of Nymphoides peltata seemed to be restricted to a region of less than 1.5 m water depth. A rise of 1.0 m in the water level after a typhoon in 1986 caused a subsequent decrease in vegetation area of Nelumbo nucifera. The vegetation recovered in shallower regions after one year in 1987. The expansion rate of Nymphoides peltata and Nelumbo nucifera in one direction was in the range 3.8–10.0 m yr^-1 and 4.1–20.8 m yr^-1, respectively. Nelumbo nucifera, Nymphoides peltata and Trapa natans, having high leaf biomass to total biomass may therefore be more vulnerable to large flood and waves. Fluctuation in water level and spatial difference in water depth may also be important factors determining the vegetation area and survival of Nymphoides peltata, Nelumbo nucifera and Trapa natans.     

Changes in algal assemblages along observed and experimental phosphorus gradients in a subtropical wetland, U.S.A.

• 1 We wanted to determine if changes in algae in the Everglades were due to increased phosphorus (P) loading. Epiphytic algae, water chemistry, and surface sediment chemistry were characterized from 32 sloughs along a P gradient in the Everglades and changes in the algal assemblages along the P gradient were compared with those along an experimental P gradient of in situ mesocosms. The sloughs are the wettest open water habitats characterized by floating and submerged aquatic plants in the Everglades.
• 2 Algal species composition was much more sensitive to P concentration than algal biomass. The diatom species variance among sloughs, captured by 1st ordination axis, was more highly correlated with total P (TP) in surface sediments (r = ‐ 0.79), than soluble reactive P (SRP) (r = ‐ 0.08) and TP (r = ‐ 0.48) in the water column. Algal biomass (µg chl a cm^‐2) was not significantly correlated with P (SRP: r = 0.22, TP: r = 0.19, sediment TP: r = 0.07) along the P gradient in the Everglades. Cluster analysis classified diatom species assemblages in 32 sloughs into three groups (TWIN I, II, III), which corresponded to three zones along the P gradient. Dominant diatom species shifted from Mastogloia smithii (40.3%), Cymbella scotica (22.3%), and Fragilaria synegrotesca (21.8%) in TWIN I to Nitzschia amphibia (22.4%) and C. microcephala (12.4%) in TWIN III. TP in surface sediments and TP in epiphyton assemblages increased 4‐ and 5‐fold from TWIN I to TWIN III, respectively.
• 3 Patterns in epiphytic assemblages along the experimental P gradient in the mesocosms were very similar to those along the Everglades P gradient. Shannon diversity indices and species richness significantly increased along both P gradients. TN : TP ratio in epiphyton assemblages significantly decreased as sediment TP increased along both P gradient. Ordination analysis showed that diatom assemblages in the impacted zone (TWIN III) were ordinated closely to the assemblages from the highest P treatments in the mesocosms. The assemblages from the less impacted zone (TWIN I) were ordinated closely to the assemblages from controls in the mesocosms.
• 4 Concurrence between results of our survey and experiments suggest that changes in epiphytic assemblages along the P gradient in the Everglades are caused by increases in P concentrations.

     

Seasonal succession of lotic epiphyton: effects of macrophyte cover and grazing by chironomid larvae

The epiphyton is a community of periphyton on the aquatic plants. The seasonal dynamics of biomass, production and community structure of epiphytic algae on Potamogeton perfoliatus was studied at the Moscow River reach with low flow velocity from May to October 2000. The relative importance of physical-chemical environmental variables and herbivorous pressure in structuring the epiphyton community was accessed using multivariate redundancy analysis. Algal composition was mainly determined by the macrophyte cover, suggestion the overriding importance of the underwater light availability in seasonal development of epiphyton. There was a significant positive relationship between biomass of attached diatom Cocconeis placentula and abundance of chironomid larvae (r = 0.57; p < 0.01). This indicates that increase in herbivorous pressure caused the shift in epiphyton community structure. However, the intensity of herbivorous impact on epiphyton was ultimately determined by light availability in macrophyte stands. At low irradiance level and consequent slow growth of algae the upper layer of epiphyton was seriously destroyed by chironomids that resulted in dominance of C. placentula. As irradiance level and epiphyton productivity increased chironomids were unable to prevent biomass accumulation of the upper layer thus inhibiting the growth of C. placentula. The high concentrations of nutrients and relatively stable discharge determined the crucial role of light regime in the regulation of trophic interactions between epiphyton and chironomid larvae.     

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.     

Epiphytes from a deep-water characean meadow in an oligotrophic New Zealand lake: species composition,biomass and photosynthesis

1. The epiphytic flora of a characean meadow in Lake Coleridge, a deep, oligotrophic lake on the South Island of New Zealand, was dominated by diatoms, particularly Eunotia pectinalis and Achnanthes minutissima. The meadows occupied a depth range from 5 to 30 m. Adnate taxa predominated at all depths below 5 m, while increased taxonomic diversity at 5 m resulted from an increased abundance of erect taxa, including chlorophytes and stalked diatoms. 2. Seasonal changes in epiphyte biomass were followed using artificial substrata and by estimating epiphyte chlorophyll a concentration on host plants. The latter required development of a novel technique utilizing the consistent relationship between fucoxanthin and chlorophyll a concentrations in the epiphyton. Epiphyte chlorophyll a on host plants varied with depth and host species between 0.1 and 0.3 mg g^–1 dry weight. Maximum epiphyte biomass was at 10–15 m depth. At depths of 15 m and less, epiphyte chlorophyll a reached a maximum of ≈ 200–300 mg m^–2 in mid-summer, while at greater depths maximum biomass was less and coincided with a period of clear water in spring. 3. Photosynthetic carbon fixation was estimated from photosynthesis–radiation curves and estimates of radiation flux at sampling depths. At depths greater than 10 m, variability of the vertical extinction coefficient of lake water rather than seasonal fluctuations in incident radiation were responsible for determining the temporal pattern of production. Chlorophyll a-specific photosynthesis was estimated to peak in summer at 5 m (8 mg mg^–1 day^–1), and in spring at all other depths. 4. Annual epiphyte production was estimated as 27 g C m^–2 year^–1 at 5 m depth, falling to 15 g C m^–2 year^–1 at 15 m and 1 g C m^–2 year^–1 at 30 m. Areal biomass changes tended to be temporally but not quantitatively coupled to estimated in situ photosynthesis, and we hypothesize that epiphyte biomass may have been controlled by grazing gastropod snails.     

Dry weight loss and colonization of plant litter by macroinvertebrates: plant species and lake types compared

Rates of dry weight loss and of colonization of litter samples of different plants by macro-invertebrates were estimated using litter bags submerged in different types of lake. Samples of Carex rostrata litter were submerged for 3 months, in summer, in two lakes, one with an acidic and nutrient-poor water and one with near-neutral and nutrient-rich water. More plant litter remained (75%) in the former lake than in the latter (51–62%, varying with the mesh size of the litter bags used). The biomass of invertebrates in the litter samples from the acid lake was about 10 times less than in those from the near-neutral. Oligochaeta and Chironomidae were the dominant taxa found in the litter bags from the acidic lake, whereas Hirudinea and Isopoda predominated in those from the near-neutral lake.Litter samples from four different plants (Carex rostrata, Typha latifolia, Triticum sp. and Phleum pratense) were immersed in litter bags in another near-neutral lake further south for 12 months. When recovered, about 45% of the Triticum litter remained compared with only 7% of the Phleum. The values for Carex and Typha fell between these two extremes. When expressed as per g dry weight of residual plant litter, the greatest biomass of invertebrates was recorded in the Phleum litter samples and the least in the Typha samples. The faunal composition of the invertebrates in the different types of litter was approximately the same, with Isopoda and Hirudinea predominating.     

Floating-leaved and emergent vegetation as habitat for fishes in a eutrophic temperate lake without submerged vegetation

Although submerged vegetation is considered to be the most suitable refuge against predators and form of foraging habitat for small fishes, submerged plants are often scarce or lacking in turbid eutrophic lakes. To evaluate emergent (Zizania latifolia) and floating-leaved (Nelumbo nucifera) vegetation as refuge areas against predators and as foraging habitats for small fishes, we investigated the fauna, abundance, and size distribution of the fish community as well as the abundance of possible prey for small fishes in beds of each vegetation type in a eutrophic shallow lake: Lake Teganuma in Japan. The leaves and stems of N. nucifera occupied an area 4.2 times larger than that of Z. latifolia. The high coverage of the water surface with plants most likely induced the hypoxia found in the N. nucifera bed. The diversity of small fishes was greater in the Z. latifolia bed with piscivorous fish than in the N. nucifera bed without piscivorous fish. The diversity of fish species in the vegetation was enhanced when there was an increased diversity of possible food sources rather than an absence of predators. Some aquatic insects of the same species had a much lower δ¹³C signature at hypoxic locations than at less hypoxic locations in the N. nucifera bed. Such site differences within a bed were not observed in the organisms caught in the Z. latifolia bed. The insects in hypoxic zones with a δ¹³C signature lower than ?30 ‰ were more depleted in ¹³C than the surface sediment or attached algae, suggesting that the larvae in the hypoxic zones incorporated the organic materials generated by methane-oxidizing bacteria. We can therefore conclude that floating-leaved vegetation, especially a N. nucifera bed, is not suitable as a replacement for submerged vegetation because of its potential to induce hypoxia, which can decrease the diversity of the fish fauna.     

Floating meadow epiphyton: biological and chemical features of epiphytic material in an Amazon floodplain lake

SUMMARY. 1. Quantities and the chemical composition of epiphyton on the roots of floating aquatic macrophytes were measured in Lake Calado, an Amazon floodplain lake. Growth of epiphytic algae following physical disturbance and losses of epiphyton due to grazing and storms were investigated.
2. Deposition of silt from invading river water decreased chlorophyll and nutrient content (%C, %N, %P) of epiphyton during rising water. N:P ratios of epiphyton indicated that proximity to the river increased supplies of phosphorus. Attached algal biomass per unit root tissue was higher overall during the falling water period, when light was greater, storms less frequent, and new host plant tissue produced more slowly.
3. Epiphytic algal biomass at the margins of floating meadows exceeded that of the phytoplankton in the open water on a per unit area basis. Increases in attached algal chlorophyll ranged from two- to ten-fold over 1 week. Artificial denudation of roots was followed by rapid regrowth of attached algae, leading, after I week, to four-fold increases in chlorophyll over the pre-denuded state.
4. Wind-blown macrophytes experienced an episodic loss of 70% of epiphytic material in less than 1 h. Particulate material lost from roots grazed by snails included root tissue and contained significantly more carbon than material lost from ungrazed roots.     

Effects of macrophyte growth forms on invertebrate communities in saline lakes of the Wyoming High Plains

In saline lakes, areal cover and both species and structural diversity of macrophytes often decline as salinity increases. To assess effects of the loss of certain macrophyte growth forms, we characterized benthic and epiphytic invertebrates in three growth forms (thin-stemmed emergents, erect aquatics, and low macroalgae) in oligosaline lakes (0.8–4.2 mS cm⁻¹) of the Wyoming High Plains, USA. We also measured the biomass and taxonomic composition of epiphytic and benthic invertebrates in two erect aquatics with very similar structure that are found in both oligosaline (Potamogeton pectinatus) and mesosaline (9.3–23.5 mS cm⁻¹) (Ruppia maritima) lakes. Although total biomass of epiphytic invertebrates varied among oligosaline lakes, the relative distribution of biomass among growth forms was similar. For epiphytic invertebrates, biomass per unit area of lake was lowest in emergents and equivalent in erect aquatics and low macroalgae; biomass per unit volume of macrophyte habitat was greatest in low macroalgae. For benthic invertebrates, biomass was less beneath low macroalgae than other growth forms. Taxonomic composition did not differ appreciably between growth forms for either benthic or epiphytic invertebrates, except that epiphytic gastropods were more abundant in erect aquatics. Total biomass of epiphytic and benthic invertebrates for the same growth form (erect aquatic) did not differ between oligosaline (Potamogeton pectinatus) and mesosaline (Ruppia maritima) lakes, but taxonomic composition did change. In the oligosaline to mesosaline range, direct toxic effects of salinity appeared important for some major taxa such as gastropods and amphipods. However, indirect effects of salinity, such as loss of macrophyte cover and typically higher nutrient levels at greater salinities, probably have larger impacts on total invertebrate biomass lake-wide.