Herbivory in omnivorous fishes: effect of plant secondary metabolites and prey stoichiometry

1. Many animals that consume freshwater macrophytes are omnivorous (i.e., they include both plant and animal matter in their diet). For invertebrate omnivorous consumers, selection of macrophyte species depends partly on the presence of secondary metabolites in plants, plant carbon/nutrient balances and/or physical structure of plants. However, little is known about the mechanisms influencing consumption of macrophytes in aquatic vertebrates. 2. For two fish species, the omnivorous rudd (Scardinius erythrophthalmus) and herbivorous grass carp (Ctenopharyngodon idella), feeding preferences were determined in three choice experiments. We tested (i) whether the presence of secondary metabolites and macrophyte stoichiometry affects macrophyte species selection by fish, (ii) the importance of macrophyte stoichiometry by manipulating the macrophytes experimentally and (iii) the rate of herbivory when the most palatable macrophyte is offered simultaneously with a common animal prey. 3. In a choice experiment with five species of submerged macrophytes (Callitriche sp., Chara globularis, Elodea nuttallii, Myriophyllum spicatum and Potamogeton pectinatus), Myriophyllum was clearly consumed least by both fishes, which strongly correlated with the highest phenolic concentration of this macrophyte. Additionally, a significant negative relationship was found between consumption and C : N ratio of the five macrophytes. The two most consumed macrophytes also had the lowest dry matter concentration (DMC). 4. In a second choice experiment, the C : N ratio of the least (Myriophyllum) and most (Potamogeton) palatable plants was manipulated by growing the macrophytes under fertilised and unfertilised conditions and subsequently feeding them to rudd. The avoidance of consumption of the chemically defended Myriophyllum by rudd was partly alleviated by the lowered C : N ratio. 5. The third choice experiment showed that both fishes preferred animal prey (the amphipod Gammarus pulex) over the most palatable macrophyte (Potamogeton) when offered simultaneously. The C : N ratio of the amphipods was about half that of the lowest C : N ratio measured in the macrophytes. Consumption by the fishes could not clearly be related to C : P or N : P ratios of prey items in any of the experiments. 6. We conclude that omnivorous fish avoid macrophytes that are chemically defended. However, when these defences are only minor, stoichiometry (C : N ratio) in combination with DMC may be a determining factor for consumption by vertebrate facultative herbivores.     

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.     

Selective herbivory by an invasive cyprinid,the rudd Scardinius erythrophthalmus

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Metal content in higher aquatic plants in a small siberian water reservoir

The dynamics of metal content in higher aquatic plants (macrophytes) in a small Bugach water reservoir in 1998–2006 was studied. A comparative estimation of the metal content in six macrophyte species (Typha latifolia L., Typha angustifolia L., Polygonium amphibium L., Potamogeton perfoliatus L., Potamogeton pectinatus L., Phragmites australis (Cav) Trin. Ex Steud.) showed that their metal concentrations do not generally exceed those known from the literature. Cluster analysis showed that the macrophyte species under study form two ecological groups with respect to the metal content, i.e., submerged plants (hydrophytes) and emergent aquatic plants (heliophytes).     

Habitat-specific predation susceptibilities of a littoral rotifer to two invertebrate predators

The rotifer Euchlanis dilatata lives associated with submerged vegetation in the littoral zone of freshwater lakes and ponds. I assessed habitat-specific predation susceptibilities for this rotifer in the presence of three aquatic macrophytes (Myriophyllum exalbescens, Elodea canadensis, and Ceratophyllum demersum) and two predators (damselfly nymphs — Enallagma carunculata; and cnidarians — Hydra). Rotifer survival was greatest on Myriophyllum in the presence of both predators. Conversely, the presence of the other macrophyte species actually increase rotifer suspectibility to predation by damselfly nymphs. I also manipulated plant structural complexity. As predicted, decreasing the relative complexity of each plant resulted in lower rotifer survival.     

Factors influencing macrophyte metrics in Estonian coastal lakes in the light of ecological status assessment

Macrophytes and factors affecting their distribution were studied in 19 coastal lakes of Estonia. The aim of the study was to determine the factors influencing the distribution of macrophytes in coastal lakes and to assess the suitability of valid macrophyte metrics. Our hypothesis was that in coastal lakes most of the macrophyte distribution patterns are caused by lake-specific variables. Morphological, physico-chemical and catchment area characteristics of the lakes varied greatly. Lakes were in different development stages—lakes nearest to the Baltic Sea were younger and more influenced by brackish water and the furthest lakes were older with more freshwater. All that variability was reflected in macrophyte parameters. Factor analysis of environmental indices divided them into three groups—catchment area, morphometric and water chemistry factors. The first factor may be considered as a pressure and the other two as lake-type-specific factors. Lake catchment area parameters had an influence on Bolboschoenus maritimus, Chara tomentosa and Typha latifolia abundance. Morphometric parameters had an influence on the depth distribution of macrophytes and water chemistry factors on the abundance of helophytes. Current indicator species showed more variability associated with lake-specific factors than with changes in status or the influence of pressures.     

Effects of contrasting omnivorous fish on submerged macrophyte biomass in temperate lakes: a mesocosm experiment

1. Freshwater fish can affect aquatic vegetation directly by consuming macrophytes or indirectly by changing water quality. However, most fish in the temperate climate zone have an omnivorous diet. The impact of fish as aquatic herbivores in temperate climates therefore remains unclear and depends on their dietary flexibility. 2. We tested the effects of a flexible omnivore and an herbivore on aquatic vegetation by comparing the effects of rudd (Scardinius erythrophthalmus, the most herbivorous fish in temperate climates) with grass carp (Ctenopharyngodon idella) in a mesocosm pond study. Exclosures distinguished herbivorous effects of fish on submerged macrophytes from indirect effects through changes in water quality, whereas stable isotope food‐web analysis provided information on fish diets. 3. We hypothesised that rudd, with its flexible diet and preference for animal food items, would only indirectly affect macrophytes, whereas grass carp, with its inflexible herbivorous diet, would directly affect macrophyte biomass. 4. Only grass carp significantly reduced macrophyte biomass through consumption. Rudd had no effect. Food‐web analysis indicated that rudd predominantly consumed animal prey, whereas grass carp included more plants in their diet, although they also consumed animal prey. Grass carp significantly affected water quality, resulting in lowered pH and increased N‐NH₄ concentrations, whereas more periphyton growth was observed in the presence of rudd. However, the indirect non‐herbivorous effects of both fish species had no effect on macrophyte biomass. 5. Both fish species should be considered as omnivores. Despite the fact that rudd is the most herbivorous fish in the western European climate zone, its effect on submerged macrophyte biomass is not substantial at natural densities and current temperatures.     

Macrophyte species composition,distribution and diversity in relation to some physicochemical factors in the littoral zone of Lake Ziway,Ethiopia

The species composition, distribution and diversity of macrophytes in Lake Ziway were determined at the end of dry and wet seasons along nine littoral sites during 2010 using a belt transect method. Some physicochemical parameters were also measured. Fourteen macrophyte species were identified with low species diversity for the lake (H' value of 1.805). The macrophyte species composition of the lake has undergone many changes during the last few decades. Arundo donax, which was never reported earlier, attained the highest relative density (30.7%) and frequency (30.5%) in this study, followed by Echinochloa colona, Potamogeton schweinfurthii, Cyperus articulatus, Typha latifolia and Cyperus papyrus. The average densities of A. donax, C. articulatus, E. colona and Pistia stratiotes were positively correlated with nutrients, whereas the density of the other species had negative correlation with physicochemical variables (CCA). Generally, the present macrophyte composition and abundance indicates healthy ecological condition of the lake. But if nutrient levels continue to increase and water levels continue to decline, we expect further changes in macrophyte composition and especially a shift towards invasive floating species.     

Nutrient removal in wetlands with different macrophyte structures in eastern Lake Taihu,China

A natural wetland of about 12 000 m² along the east coast of Lake Taihu was separated into five subzones with different macrophyte structures to investigate their nutrient removal dynamics. Wastewater was continuously pumped into the wetland from July 2008 to June 2009 at an average rate of 22 m³/h. Neighboring natural wetland with high density of macrophyte was chosen as a comparison site. The removal of TN, TDN, TP, and TDP in the experimental wetlands as a whole was about 79.3, 54.5, 4.5, and 3.4 kg, respectively. The decrease of nitrogen concentration was more pronounced in winter (January–March) 2009, representing a respective reduction of 46.4%, 48.0%, and 47.9% in TN, TDN, and NH₄–N concentration. Results reveal a higher nutrient removal potential in wetland dominated by Typha orientalis Presl, Zizania latifolia Turcz, and Hemarthria sibirica under high nutrient load. However, areas dominated by Zizania latifolia Turcz, Nelumbo nucifera Gaertn, and Ceratophyllum demersum L. had better purification performance when the above-water-surface macrophytes were harvested frequently. Dissolved oxygen, pH, and oxidation–reduction potential decreased with the increase of the percentage of Zizania latifolia Turcz-dominated macrophytes. High nutrient concentration in the comparison site and net increase of NH₄–N in Z1 indicate the possibility of water re-pollution by intense macrophyte decomposition. Furthermore, results suggest that harvesting macrophytes has potential ability in nitrogen, especially ammonium nitrogen removal, and hence could be considered in wetland construction for lake restoration.     

The effects of shoot age on colonization of an emergent macrophyte (Typha latifolia) by macroinvertebrates

1. The colonization and dynamics of epiphytic aquatic macroinvertebrates are described on first-year and second-year shoots of an emergent macrophyte (Typha latifolia) in a Swiss pond. Effects of shoot senescence on composition, richness, density and biomass of the macroinvertebrates are quantified. 2. There were two phases of colonization: a short-term colonization process which corresponded with that usually observed on inert substrates and a longer term colonization process related to the attractiveness of the shoots for the colonizers. In this second process, the older shoots showed a higher attractiveness for most invertebrate taxa. 3. Taxa colonized the shoots at different rates. Rapid colonizers included the limpet Ferrissia wautieri and the mayfly Cloeon dipterum. Conversely, Oligochaeta, Chironomidae and Ceratopogonidae were particularly slow colonizers. 4. The older shoots supported a higher mean annual richness, abundance and biomass of invertebrates. Compared with younger shoots, the older shoots demonstrated a higher carrying capacity for most invertebrate taxa, as is the case for many other freshwater macrophytes.     

Aquatic macrophyte breakdown in an Appalachian river

Weight loss from Podostemum ceratophyllum, Elodea canadensis, Potamogeton crispus, Justicia americana, and Typha latifolia was measured by exposing air dried leaf material in 15 cm² nylon mesh bags (3 mm octagonal openings). Breakdown rates for these species were 0.037, 0.026, 0.02l, 0.016, and 0.007 day^–1, respectively. In general, these rates are much faster than reported rates of tree leaf breakdown in streams. Shredders accounted for 35% of the macroinvertebrates found on the leaf bags. However, macroinvertebrate densities on the aquatic macrophyte tissue were well below densities commonly found on leaf bags in small streams. The rapid breakdown of aquatic macrophytes in the New River suggests that organic matter from this source may constitute a significant pulse in the annual energy dynamics of the river.     

A Method to Estimate Practical Radial Oxygen Loss of Wetland Plant Roots

The estimation of practical radial oxygen loss (ROL) of wetland plant roots was attempted in this study. We have devised a new method to measure ROL of wetland plant roots. The whole root system was bathed in an anoxic nutrient solution. Oxygen released from the root was removed immediately by introducing oxygen-free nitrogen gas (O₂ < 4 nmol L⁻¹) to mimic natural habitats where released oxygen is consumed rapidly due to chemical and biological oxidation processes. Oxygen removed from the root-bathing chamber was simultaneously detected colorimetrically by use of the highly oxygen-sensitive anthraquinone radical anion (AQ·⁻) in a cell outside the root-bathing chamber, which decolorized by a rapid reaction with oxygen. An emergent macrophyte Typha latifolia L. was incubated, and its ROL was measured by both the new method and one of the conventional methods, the closed chamber/electrode method, by which the ROL of Typha latifolia L. had not yet been measured. The new method succeeded in detecting the ROL, whereas the conventional method was not able to detect oxygen, due to the level being below the detection limit of the oxygen electrode. The oxygen supply via the seedlings of Typha latifolia L. was ca. 10 times higher compared with control measurements without plant. Light illumination significantly enhanced the ROL of Typha latifolia L. (0.33 nmol O₂ g⁻¹ root dry weight s⁻¹ under light and 0.18 nmol O₂ g⁻¹ root dry weight s⁻¹ in the dark). Theses values fall between those previously reported by the closed chamber/titanium citrate method and the open chamber/electrode method.     

Loss of Submerged Macrophytes in Shallow Lakes in North‐Eastern Germany

During the 1950s, the submerged vegetation of shallow lakes in north‐eastern Germany was dominated by nutrient tolerant species, with Ceratophyllum demersum and Myriophyllum sp. being most common. Almost one third of 300 investigated lakes had already lost their submerged macrophytes at that time. Very shallow lakes showed either high or low macrophyte abundance. Increasing depth resulted in medium macrophyte abundances, which may contribute to the stabilisation of local or temporary clearwater states. Forty years later, the percentage of lakes without macrophytes had dramatically increased. Between 55 and 85% of the investigated lakes showed a low abundance. The decline was most pronounced in very shallow lakes. The majority of the investigated lakes showed summer TP concentrations below 100 μg L^–1, but no colonisation by submerged macrophytes, which indicates a resilience against re‐colonisation.     

Interactive effects of road salt and sediment disturbance on the productivity of seven common aquatic macrophytes

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Interspecific differences in radial oxygen loss from the roots of three <Emphasis Type="Italic">Typha</Emphasis> species

A comparison was made of the radial oxygen loss (ROL) from the roots of three Typha species, Typha latifolia L., Typha orientalis Presl and Typha angustifolia L., which resemble each other in morphology. ROLs were evaluated in the laboratory for seedlings of T. orientalis and T. angustifolia in order to compare them with the ROL value for T. latifolia obtained in our previous study. Measurements were conducted using the highly oxygen-sensitive anthraquinone radical anion as an oxygen indicator, which enabled us to simulate the natural conditions in which the oxygen released from the root is immediately consumed by the soil. Among the three Typha species, the ROL was the highest in T. angustifolia, followed by T. latifolia and T. orientalis. Illumination significantly enhances the ROL of T. orientalis, and this effect was also observed for T. latifolia in our previous study, whereas it did not affect the ROL of T. angustifolia. These results indicate that ROL differs significantly between species, even among members of the same genus that are similar in morphology.     

Implications of rudd (Scardinius erythrophthalmus) herbivory on submerged macrophytes in a shallow eutrophic lake

The role of rudd (Scardinius erythrophthalmus) herbivory was studied in Kirkkojärvi, a shallow and turbid basin in Lake Hiidenvesi, Finland. The submerged species dominating in the rudd diets were Potamogeton obtusifolius, Ranunculus circinatus, Sparganium emersum, bryophytes, and filamentous algae. Plant consumption estimated with bioenergetics modelling increased with fish age, being highest in late summer concomitant with the macrophyte biomass peak. Depending on the age structure, a rudd biomass of 20 kg ha^?1 consumes 18–23 kg of macrophytes ha^?1 a^?1, while a rudd biomass of 100 kg ha^?1 results in plant consumption of 92–115 kg ha^?1 a^?1. Although, rudd seemed to feed rather unselectively on suitable-sized and edible plants, some species abundant in the littoral, such as Myriophyllum verticillatum and pleustophytic Ceratophyllum demersum, were not found in rudd guts, indicating selective plant consumption. In Kirkkojärvi, selective grazing by rudd and increased turbidity and high nutrient levels partly caused by bottom dwelling cyprinid fish, may promote the inedible and pleustophytic macrophytes, which have increased in Kirkkojärvi during the past decades.

     

Effects of macrophyte heterogeneity and food availability on structural parameters of the macroinvertebrate community in a Pampean stream

Environmental heterogeneity in natural ecosystems influences several parameters at the population and community levels. In freshwater ecosystems, habitat heterogeneity can be provided by macrophyte species with different structural shapes. Previous studies suggest that aquatic plants with more complex architectures will support higher number, biomass, and taxon richness of macroinvertebrates than plants with simpler shape. We investigated the influence of macrophyte structural heterogeneity (quantified by fractal dimension) and food availability (represented by epiphytic biomass) on several parameters (number of individuals, biomass, body size distribution, taxon richness, and diversity) of the macroinvertebrate community in a Pampean stream. Four submerged macrophyte species (Egeria densa, Elodea ernstae, Ceratophyllum demersum, and Stuckenia striata) and associated macroinvertebrates were sampled in late spring, summer, and autumn. Plants were photographed and fractal dimension was estimated from the images by the box-counting method. Fractal dimension was independent of plant surface area per unit of macrophyte biomass and differed significantly among species. Mean fractal dimension varied between 1.29 and 1.62, and increased following the sequence E. densa → S. striata → E. ernstae → C. demersum. Macrophyte species with higher fractal dimension supported a greater abundance of macroinvertebrates, especially those of small body size (500–1,000 μm); but fractal dimension was unrelated to macroinvertebrate biomass, richness, and diversity. However, overall animal biomass was significantly associated to the epiphytic abundance. Consequently, macrophyte heterogeneity influences macroinvertebrate density and body size distribution, while animal biomass depends on epiphytic food resources provided by plants.     

Plant community dynamics in a chain of lakes: principal factors in the decline of rooted macrophytes with eutrophication

Shoe Lake and East Graham Lake, part of a small chain of lakes in southeastern Michigan, USA, differ in nutrient loading and in the structure and productivity of their aquatic plant communities. A comparative study of species frequency and biomass distributions, nutrient contents, and responses to experimental nutrient enrichment and shading, was conducted to determine the principal factors controlling the macrophyte dynamics. A central objective was to address the question of why rooted macrophyte growth declines with eutrophication, and to test existing models designed to explain this phenomenon. In the more eutrophic Shoe Lake, diversity and productivity of rooted macrophytes were relatively low, restricted primarily by combined shading of phytoplankton, periphyton, and non-rooted macrophytes (principally Ceratophyllum demersum, along with Utricularia vulgaris and Cladophora fracta). In the less eutrophic East Graham Lake, lower nitrogen availability restricted the growth of all of these shading components, resulting in clearer water and higher productivity and diversity of rooted macrophytes. The macrophytes did not allelopathically suppress the phytoplankton in East Graham Lake. The results supported a direct relationship between nutrient loading, increasing growth of phytoplankton, periphyton and non-rooted macrophytes, and decline of rooted macrophytes.     

Chemical oxygen demand,nitrogen and phosphorus removal by subsurface wetlands with Phragmites vegetation in different models

To improve the removal efficiency of subsurface wetlands vegetated mainly by Phragmites, pilot‐scale gravel‐based wetlands were used to treat sewage characterized by chemical oxygen demand (COD), total nitrogen (TN) and total phosphorus (TP) pollution. For Phragmites vegetation, COD, TP and TN removal loads of wetland vegetation with Phragmites australis–Typha angustata–Scirpus validus as main species reached 0.517 g m^?2 d^?1, 0.277 g P m^?2 d^?1 and 0.023 g N m^?2 d^?1. The COD removal loads in pilot‐scale and medium‐scale (260 m² in area) wetlands with Phragmites‐monoculture vegetation were 0.62–0.64 g m^?2 d^?1, while that of P. australis–T. angustata–S. validus wetland reached 0.974 g m^?2 d^?1. Thus, the preferable poly‐culture model for Phragmites wetland vegetation was P. australis, T. angustata, S. validus and Zizania latifolia with stem density ratio of 8:1:5:1. After harvest, nitrogen and phosphorus standing stocks of wetland vegetations ranged only 2.2–9.93 g N m^?2 and 5.39–13.5 g P m^?2, respectively, as both the above ground biomass and the nitrogen and phosphorus contents of the wetland vegetation harvested in late autumn were low.     

A Century of Change in Macrophyte Abundance and Composition in Response to Agricultural Eutrophication

Clear Lake, Iowa, USA is a shallow, agriculturally eutrophic lake that has changed drastically over the past century. Eight macrophyte surveys since 1896 were pooled and examined to characterize long-term impacts of eutrophication on macrophyte community composition and relative abundance. Surveys in 1981 and 2000 revealed few submergent and floating-leaved species and a dominance in emergent species (Scirpus, Typha). Over the past century, however, species richness has declined from a high of 30 species in 1951 to 12 found today, while the community composition has shifted from submergent-(99%) to emergent-dominated floras (84%). Potamogeton praelongus was the first emergent species to disappear but was followed by several other clear water Potamogeton species. Several floating leaved and emergent genera increased in relative abundance with eutrophication, notably Nuphar, Nymphaea, Phragmites, Polygonum, Sagittaria, Scirpus, and Typha. P. pectinatus was present over the entire century due to its tolerance of eutrophic conditions. Macrophyte growth was generally light-limited, with 93% of the variance in relative abundance of submergent species explained by changes in water transparency. Clear Lake exhibits signs of alternative stable states, oscillating between clear and turbid water, coupled with high and low submerged species relative abundance. The maximum macrophyte richness occurred as the lake oscillated between submergent- and emergent-dominated states. Changes in the water level have also impacted macrophyte growth since the area of the lake occupied by emergent macrophytes was negatively correlated with water level. Strongest correlations indicated that macrophytes respond to water level variations with a 2-year time-lag.