Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

Changes in phosphorus and nitrogen cycling following food web manipulations in a shallow Dutch lake

1. The flow of phosphorus and nitrogen through the food web of the shallow, eutrophic lake Wolderwijd was analysed for 2 different years before and for 1 year after food web manipulation.
2. After fish removal the water became clear and the growth of macrophytes began. Fish removal resulted in a significant reduction of the total nutrient pool in the water, but differences between the nutrient cycles before and after the experiment were mainly caused by a gradual change driven by a reduced phosphorus input.
3. The zooplankton biomass before and after food web manipulation did not change significantly. Unfavourable food conditions and predation by young fish limited zooplankton biomass after the food web manipulation.
4. After fish removal benthic algae, fish, zoobenthos and macrophytes form the largest pools of nutrients apart from the sediment top layer. However, they contribute only little to nutrient cycles in the water column.     

Changes in phosphorus and nitrogen cycling following food web manipulations in a shallow Dutch lake

1. The flow of phosphorus and nitrogen through the food web of the shallow, eutrophic lake Wolderwijd was analysed for 2 different years before and for 1 year after food web manipulation.
2. After fish removal the water became clear and the growth of macrophytes began. Fish removal resulted in a significant reduction of the total nutrient pool in the water, but differences between the nutrient cycles before and after the experiment were mainly caused by a gradual change driven by a reduced phosphorus input.
3. The zooplankton biomass before and after food web manipulation did not change significantly. Unfavourable food conditions and predation by young fish limited zooplankton biomass after the food web manipulation.
4. After fish removal benthic algae, fish, zoobenthos and macrophytes form the largest pools of nutrients apart from the sediment top layer. However, they contribute only little to nutrient cycles in the water column.     

Effects of food‐web structure on periphyton stoichiometry in eutrophic lakes: a mesocosm study

1. Aquatic herbivores typically have much higher concentrations of nutrients (e.g. N and P) in their tissues than there is in the food they eat. These stoichiometric differences can cause herbivores to be limited by the elemental quality of their food, which could affect, in turn, the structure of consumer communities and even alter key ecosystem processes. 2. In streams and in the littoral zone of shallow lakes, periphyton is an important food resource for benthic animals. Studying the elemental composition of periphyton may help us to understand food‐web structure, and any reciprocal effect of this structure on periphyton stoichiometry. 3. To understand how alterations in the food‐web structure affect the elemental composition of periphyton in a eutrophic lake, we carried out a long‐term experiment (14 months) in large‐scale mesocosms (40 m³), in which we manipulated food‐web structure, and which were dominated either by planktivorous fish (Rutilus rutilus) or herbivorous invertebrates (without fish). Periphyton was sampled monthly at three depths (0.5, 1.5 and 2.5 m) to determine its biomass and elemental composition (C/N/P ratio). Food‐web structure, physical and chemical parameters were monitored throughout the experiment. 4. Fish had indirect positive effect on periphyton biomass, leading to twofold higher levels than in herbivore‐dominated mesocosms. This result was probably due to control of benthic consumers by fish, suggesting a strong top–down control on periphyton by their consumers in fishless enclosures. 5. The elemental ratios C/P and C/N were lower in deep water in both treatments, mainly mediated by light availability, in accordance with the light/nutrient ratio hypothesis. These ratios were also lower in fishless treatments, probably due to increases in inorganic nutrient availability and grazing pressure in herbivore‐dominated systems. During winter, periphyton elemental composition was similar in both treatments, and was unrelated to inorganic nutrient availability. 6. These results indicate that any alteration of food‐web structure in lakes, such as in biomanipulation experiments, is likely to modify both the biomass and elemental quality of periphyton. Resultant effects on the consumers of periphyton and macrophytes could play a key role in the success of biomanipulations and should be taken into account in further studies.     

The influence of water level on macrophyte growth and trophic interactions in eutrophic Mediterranean shallow lakes: a mesocosm experiment with and without fish

1. Water‐level fluctuations are typical of lakes located in the semi‐arid Mediterranean region, which is characterised by warm rainy winters and hot dry summers. Ongoing climate change may exacerbate fluctuations and lead to more severe episodes of drought, so information on the effects of water level on the functioning of lake ecosystems in such regions is crucial. 2. In eutrophic Lake Eymir, Turkey, we conducted a 4‐month (summer) field experiment using cylindrical 0.8‐m‐ (low‐water‐level) and 1.6‐m‐deep (high‐water‐level) mesocosms (kept open to the sediment and atmosphere). Fish (tench, Tinca tinca, and bleak, Alburnus escherichii) were added to half of the mesocosms, while the rest were kept fishless. Ten shoots of Potamogeton pectinatus were transplanted to each mesocosm. 3. Sampling for physicochemical variables, chlorophyll a (chl‐a), zooplankton and per cent plant volume inhabited (PVI%) by macrophytes was conducted weekly during the first 5 weeks, and subsequently biweekly. Macrophytes were harvested on the last sampling date. During the course of the experiment, the water level decreased by 0.41 ± 0.06 m. 4. Throughout the experiment, fish affected zooplankton abundance (?), nutrient concentrations (+), chl‐a (+) and water clarity (?) most strongly in the low‐water‐level mesocosms and the zooplankton community shifted towards dominance of small‐sized forms. The fishless mesocosms had a higher zooplankton/phytoplankton ratio, suggesting higher grazing. 5. Greatest macrophyte growth was observed in the low‐water‐level fishless mesocosms. However, despite high nutrient concentrations and low water clarity, macrophytes were also abundant in the fish mesocosms and particularly increased following a water‐level decrease from midsummer onwards. Macrophyte growth was poor in the high‐water‐level mesocosms, even in the fishless ones with high water clarity. This was ascribed to extensive periphyton development reducing light availability for the macrophytes. 6. Our results indicate that a reduction in water level during summer may help maintain the growth of macrophytes in Mediterranean eutrophic shallow lakes, despite a strong negative effect of fish predation on water clarity. It is therefore probable that an expected negative effect of global climate change on water clarity because of eutrophication and enhanced top‐down control of fish may be, at least partly, counteracted by reduced water level, provided that physical disturbance is not severe.     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

Combined stable isotope and gut contents analysis of food webs in plant-dominated, shallow lakes

Summary 1. To determine feeding links between primary producers, invertebrates and fish, stable isotope analyses and gut content analyses of fish were conducted on the components of four shallow, eutrophic to hypertrophic, plant-dominated lakes.
2. Although separation of basal resources was possible, the diets of both fish and invertebrates were broad, comprising food from different compartments (planktonic, epiphytic/benthic), as well as from different trophic levels.
3. Mixing models were used to determine the extent to which periphyton production supported higher trophic levels. Only one species of invertebrate relied upon periphyton production exclusively.
4. Fish density affected the diets of invertebrates. The response was different for planktonic and epiphytic/benthic invertebrates. The proportion of periphyton production in the diets of zooplankton appeared to increase with fish density, whilst it decreased for other invertebrates.
5. As all zooplankton samples were collected in the open water at dusk, these results are further evidence for the diurnal horizontal migration of zooplankton. Although not conclusive, they are consistent with a behavioural response by invertebrates and zooplankton in the presence of fish.     

Phytoplankton food quality control of planktonic food web processes

We developed a mechanistic model of nutrient, phytoplankton, zooplankton and fish interactions to test the effects of phytoplankton food quality for herbivorous zooplankton on planktonic food web processes. When phytoplankton food quality is high strong trophic cascades suppress phytoplankton biomass, the zooplankton can withstand intense zooplanktivory, and energy is efficiently transferred through the food web sustaining higher trophic level production. Low food quality results in trophic decoupling at the plant-animal interface, with phytoplankton biomass determined primarily by nutrient availability, zooplankton easily eliminated by fish predation, and poor energy transfer through the food web. At a given nutrient availability, food quality and zooplanktivory interact to determine zooplankton biomass which in turn determines algal biomass. High food quality resulted in intense zooplankton grazing which favored fast-growing phytoplankton taxa, whereas fish predation favored slow-growing phytoplankton. These results suggest algal food quality for herbivorous zooplankton can strongly influence the nature of aquatic food web dynamics, and can have profound effects on water quality and fisheries production. Handling editor: D. Hamilton     

Littoral species diversity and biomass: concordance among organismal groups and the effects of environmental variables

Biomass and species diversity (richness and evenness) of littoral organisms were explored in 27 sites in three basins of the large Lake Saimaa system in eastern Finland. The basins differed in degree of nutrient loading and trophic status. Six organismal groups, i.e., phytoplankton, periphyton, macrophytes, crustacean zooplankton, benthic macroinvertebrates and fish were studied. Factors affecting the biomass and diversity of these groups were explored by multiple stepwise regression analysis. The biomass of different groups was explained by the same variables, mainly nutrients, while diversity was associated with different environmental factors among the studied groups. The biomass of periphyton, phytoplankton, zooplankton, and fish correlated significantly with each other. There was also an apparent association between the biomass of macrophytes and that of benthic invertebrates. However, no significant correlations were found among the diversity of the studied groups. In accordance with previous studies, our results did not support the existence of species-rich hotspots or the possibility of using any surrogate taxon to reveal overall biodiversity. Thus, for conservation planning, biological surveys should include extensive collection of taxonomic groups and organisms at all trophic levels.     

Food web structure in a tropical stream ecosystem

Abstract This study investigated the structure and properties of a tropical stream food web in a small spatial scale, characterizing its planktonic, epiphytic and benthic compartments. The study was carried out in the Potreirinho Creek, a second‐order stream located in the south‐east of Brazil. Some attributes of the three subwebs and of the conglomerate food web, composed by the trophic links of the three compartments plus the fish species, were determined. Among compartments, the food webs showed considerable variation in structure. The epiphytic food web was consistently more complex than the planktonic and benthic webs. The values of number of species, number of links and maximum food chain length were significantly higher in the epiphytic compartment than in the other two. Otherwise, the connectance was significantly lower in epiphyton. The significant differences of most food web parameters were determined by the increase in the number of trophic species, represented mainly by basal and intermediate species. High species richness, detritus‐based system and high degree of omnivory characterized the stream food web studied. The aquatic macrophytes probably provide a substratum more stable and structurally complex than the sediment. We suggest that the greater species richness and trophic complexity in the epiphytic subweb might be due to the higher degree of habitat complexity supported by macrophyte substrate. Despite differences observed in the structure of the three subwebs, they are highly connected by trophic interactions, mainly by fishes. The high degree of fish omnivory associated with their movements at different spatial scales suggests that these animals have a significant role in the food web dynamic of Potreirinho Creek. This interface between macrophytes and the interconnections resultant from fish foraging, diluted the compartmentalization of the Potreirinho food web.     

Long- and short-term dynamics of submerged macrophytes in two shallow eutrophk lakes

1. Periods with clear water and abundant submerged vegetation have alternated with periods of turbid water and sparse vegetation during recent decades in Lake Tåkern and Lake Krankesjön, two shallow, calcium-rich, moderately eutrophic lakes in southern Sweden, Between 1983 and 1991, submerged vegetation (predominant species: Chara tomentosa, Nitellopsis obtusa, Myriophyllum spicatum) covered about 50% of the open lake area in Lake Tåkern. In Lake Krankesjön, submerged vegetation was sparse during 1983–84, but increased continuously in the following years and covered about 50% of the open lake area by 1990 and 1991. Potamogeton pectinatus was the first species to expand in Lake Krankesjön, but was later replaced by C. tomentosa. 2. During 1983–84, turbidity was high in Lake Krankesjön, which indicated that submerged macrophytes were light-limited. During 1986–91, there was a negative correlation between the areal coverage of charophytes and angiosperms, indicating that competition for space had become an important limiting factor. The same negative correlation was found in Lake Tåkern for 1983–91. 3. Charophytes had much higher biomass per unit area than angiosperms in both lakes and reduced water movement considerably. This was probably one reason for the increase of water transparency in Lake Krankesjön during the spatial expansion of these plants. Charophytes also stored large amounts of phosphorus and nitrogen, Charophytes are probably superior competitors for both space and nutrients and thus have competitive advantage over angiosperms in this lake type. 4. In Lake Krankesjön, both P. pectinatus and C. tomentosa were negatively affected by high water level during the growing period. Total disappearance of submerged vegetation occurred in both lakes after catastrophic events (dry-out during summer or mechanical damage by ice) caused by extremely low water level. Changes in water level are thus one of the most important reasons for among-year fluctuations in areal coverage of submerged macrophytes in these lakes.     

Decline of charophytes during eutrophication: comparison with angiosperms

1. Charophytes have disappeared from several enriched lakes in Scania (southern Sweden) since the 1940s. Poor light conditions, rather than a toxic effect of phosphorus or negative impact of fish, are the most probable reason for this decline. 2. Small species of charophytes (shoot diameter 0.5–1.0 mm), which are able to form dense, low mats, still occur in eutrophic lakes with high phosphorus concentrations, but are restricted to areas of shallow water. In contrast, large species (shoot diameter 1–4 mm) have totally disappeared from the most turbid lakes. I suggest that these species are unable to grow in very shallow water because of damage by ice and wave action. 3. Maximum depth distribution (z_e) and Secchi depth (D) were measured in Scanian lakes for both charophytes and angiosperms and combined with data obtained from Chambers & Kalff (1985). According to the combined data, z_c and D are closely correlated with each other for both angiosperms and charophytes. 4. The z_c of charophytes is higher than z_c of angiosperms in clear lakes but lower in turbid lakes. Higher z_c of angiosperms in the most turbid lakes is explained by special adaptations of these species to poor light availability (shoot elongation, canopy formation, rapid growth during spring).     

基于稳定同位素技术的保安湖食物网结构特征研究

文章利用碳、氮稳定同位素技术对江湖阻隔典型湖泊-保安湖的食物网结构进行了研究。结果表明保安湖中鱼类消费者的主要营养级范围为2.1—3.3, 在调查到的16种鱼类中, 顶级肉食性鱼类种类很少, 杂食性鱼类的种类最多。保安湖食物网主要由两条营养传递途径构成, 即由POM、浮游植物为主要食物源的浮游牧食链与沉积物为主要食物源的底栖食物链。POM、浮游植物、浮游动物和底栖动物是保安湖水域食物网中鱼类的主要食物来源, 其次是沉积物中的碎屑和水生植物等。此外, 从基于理论食性数据的食物网与BIMM模型预测的食物网结构可以看出, 从POM、浮游植物、浮游动物到杂食性鱼类的浮游牧食链在整个食物网中具有主导性, 而从水生植物、沉积物和底栖动物到杂食性鱼类的底栖食物链相对重要性较低。     

Coexistence of Fish Species in a Large Lowland River: Food Niche Partitioning between Small-Sized Percids,Cyprinids and Sticklebacks in Submersed Macrophytes

In the spring and summer of each year, large patches of submersed aquatic macrophytes overgrow the bottom of the alluvial Warta River downstream of a large dam reservoir owing to water management practices. Environmental variables, macroinvertebrates (zoobenthos and epiphytic fauna, zooplankton) and fish abundance and biomass were assessed at this biologically productive habitat to learn intraseasonal dynamics of food types, and their occurrence in the gut contents of small-sized roach, dace, perch, ruffe and three-spined stickleback. Gut fullness coefficient, niche breadth and niche overlap indicated how the fishes coexist in the macrophytes. Chironomidae dominated in the diet of the percids. However, ruffe consumed mostly benthic chironomids, while perch epiphytic chironomids and zooplankton. The diet of dace resembled that in fast flowing water although this rheophilic species occurred at unusual density there. The generalist roach displayed the lowest gut fullness coefficient values and widest niche breadth; consequently, intraspecific rather than interspecific competition decided the fate of roach. Three-spined stickleback differed from the other fishes by consuming epiphytic simuliids and fish eggs. The diet overlap between fishes reaching higher gut fullness coefficient values was rather low when the food associated with the submersed aquatic macrophytes was most abundant; this is congruent with the niche overlap hypothesis that maximal tolerable niche overlap can be higher in less intensely competitive conditions.     

Impact of fish predation on cladoceran body weight distribution and zooplankton grazing in lakes during winter

1. It is well accepted that fish, if abundant, can have a major impact on the zooplankton community structure during summer, which, particularly in eutrophic lakes, may cascade to phytoplankton and ultimately influence water clarity. Fish predation affects mean size of cladocerans and the zooplankton grazing pressure on phytoplankton. Little is, however, known about the role of fish during winter. 2. We analysed data from 34 lakes studied for 8–9 years divided into three seasons: summer, autumn/spring and winter, and four lake classes: all lakes, shallow lakes without submerged plants, shallow lakes with submerged plants and deep lakes. We recorded how body weight of Daphnia and then cladocerans varied among the three seasons. For all lake types there was a significant positive correlation in the mean body weight of Daphnia and all cladocerans between the different seasons, and only in lakes with macrophytes did the slope differ significantly from one (winter versus summer for Daphnia). 3. These results suggest that the fish predation pressure during autumn/spring and winter is as high as during summer, and maybe even higher during winter in macrophyte‐rich lakes. It could be argued that the winter zooplankton community structure resembles that of the summer community because of low specimen turnover during winter mediated by low fecundity, which, in turn, reflects food shortage, low temperatures and low winter hatching from resting eggs. However, we found frequent major changes in mean body weight of Daphnia and cladocerans in three fish‐biomanipulated lakes during the winter season. 4. The seasonal pattern of zooplankton : phytoplankton biomass ratio showed no correlation between summer and winter for shallow lakes with abundant vegetation or for deep lakes. For the shallow lakes, the ratio was substantially higher during summer than in winter and autumn/spring, suggesting a higher zooplankton grazing potential during summer, while the ratio was often higher in winter in deep lakes. Direct and indirect effects of macrophytes, and internal P loading and mixing, all varying over the season, might weaken the fish signal on this ratio. 5. Overall, our data indicate that release of fish predation may have strong cascading effects on zooplankton grazing on phytoplankton and water clarity in temperate, coastal situated eutrophic lakes, not only during summer but also during winter.     

Submerged Macrophytes Mitigate Direct and Indirect Insecticide Effects in Freshwater Communities

Understanding how ecological interactions mitigate the impacts of perturbations such as pesticides in biological communities is an important basic and applied question for ecologists. In aquatic ecosystems, new evidence from microcosm experiments suggests that submerged macrophytes can buffer cladocerans from pulse exposures to the widely used insecticide malathion, and that mitigation increases with macrophyte density. However, whether these results scale up to more complex aquatic communities where ecological interactions such as competition can alter toxicity is unknown. Further, macrophyte abilities to mitigate different insecticide exposure scenarios (i.e. single versus repeated pulses) have never been tested. To address these gaps, we performed a factorial mesocosm experiment examining the influence of four macrophyte treatments (0, 10, 50, or 100 Elodea Canadensis shoots planted per mesocosm) crossed with three malathion exposure scenarios (no insecticide, single pulse, repeated pulses) on aquatic communities containing zooplankton, phytoplankton, periphyton, two snail species, and larval amphibians. In the absence of macrophytes, single malathion pulses caused short-term declines in cladoceran abundance followed by their rapid recovery, which precluded any indirect effects (i.e. trophic cascades). However, repeated malathion pulses caused cladoceran extinctions, resulting in persistent phytoplankton blooms and reduced abundance of one snail species. In contrast, with macrophytes present, even at low density, malathion had no effect on any taxa. We also discovered novel effects of macrophytes on the benthic food web. In the two highest macrophyte treatments, we observed trends of reduced periphyton biomass, decreased abundance of one snail species, and decreased amphibian time to and mass at metamorphosis. To our knowledge, this is the first evidence of negative submerged macrophyte effects on amphibians, a taxa of global conservation concern. Our findings suggest that facilitating macrophytes could be an important strategy for buffering freshwater communities from insecticides, though consideration of their impacts on animal species is necessary.     

Depth limits and minimum light requirements of freshwater macrophytes

1. Data for maximum colonization depth (Z_c) of five groups of submerged macrophytes and light attenuation were collected for forty-five Danish lakes and 108 non-Danish lakes. The macrophyte groups were bryophytes, charophytes, caulescent angiosperms, rosette-type angiosperms and Isoetes spp. 2. The data showed systematic differences among the groups in the relationship of Z_c to water transparency. In lakes with low transparency (Secchi disc transparency (Z_s) less than 7 m) caulescent angiosperms and charophytes penetrated deepest followed by bryophytes and Isoetes spp. In more transparent lakes bryophytes grew deepest, followed by charophytes, caulescent angiosperms and Isoetes spp. Rosette-type angiosperms had the lowest Z_c in all types of lakes. Charophytes and caulescent angiosperms had similar depth limits in lakes with Z_s < 4 m but charophytes grew deeper in more transparent lakes. The depth limits of both groups were independent of light penetration in lakes with very low transparency (Z_s < 1 m). The annual light exposure for the deepest growing macrophytes (bryophytes) was 20–95 mol photons m^–2. 3. The relationship between Z_c, macrophyte type and lake transparency could be explained by three distinct processes regulating Z_c. In lakes with low transparency (Z_s < 1 m), tall macrophytes (caulescent angiosperms and charophytes) compensate for light limitation by shoot growth towards the water surface and Z_c is therefore independent of transparency. In lakes with medium transparency (1 m < Z_s < 4 m) Z_c for angiosperms, charophytes and Isoetes spp. is constrained by light attenuation in the water column, corresponding to a linear relationship between Z_c and Z_s. This pattern also applies to bryophytes, despite lake transparency. In transparent lakes, the minimum light requirement at Z_c increased with increasing transparency for angiosperms, charophytes and Isoetes spp. 4. The minimum light requirements among submersed macrophytes (including marine macroalgae) depend on their plant-specific carbon value (plant biomass per unit of light-absorbing surface area) for the species/group, indicating that the light requirements of submersed plants are tightly coupled to the plants’ possibility to harvest light and hence to the growth form. 5. The light requirements increased on average 0.04% surface irradiance per degree increase in latitude corresponding to an average decrease in Z_c of 0.12 m per degree latitude.     

Top-down control in pelagic systems: a role for invertebrate predation

Limnologists have long recognized the importance of predation in freshwater communities. The majority of study of predator effects has involved vertebrate predators, with emphasis on planktivorous fish. Documented effects of planktivorous fish have been so dramatic that manipulations of their populations are seen by many as potential tools in lake management. However, the success of such manipulations is often less than desired due to the ubiquitous complexity of food webs and the pervasiveness of compensatory responses to food web manipulation. Recently, enormous effort has been applied to the Lake Kinneret pelagic food web in effort to reduced the abundance of the planktivorous Kinneret bleak Acanthobrama terraesanctae and thereby increase the biomass of herbivorous zooplankton in the hopes of increasing water clarity. We compared potential predation pressure on Lake Kinneret herbivorous zooplankton by bleak and the other major zooplankton predators in the lake, the cyclopoid copepods Mesocyclops ogunnus and Thermocyclops dybowskii. We found that, despite having much lower biomass, cyclopoid copepods accounted for a greater portion of the predation mortality on herbivorous zooplankton than bleak. Our results suggest that reductions in predation pressure by bleak will not yield subsequent increases in herbivorous zooplankton biomass. Rather, reductions in bleak predation pressure may allow for increases in cyclopoid copepod abundance and thereby a net increase in predation pressure on herbivorous zooplankton.     

Global ecological impacts of invasive species in aquatic ecosystems

The introduction of invasive species, which often differ functionally from the components of the recipient community, generates ecological impacts that propagate along the food web. This review aims to determine how consistent the impacts of aquatic invasions are across taxa and habitats. To that end, we present a global meta‐analysis from 151 publications (733 cases), covering a wide range of invaders (primary producers, filter collectors, omnivores and predators), resident aquatic community components (macrophytes, phytoplankton, zooplankton, benthic invertebrates and fish) and habitats (rivers, lakes and estuaries). Our synthesis suggests a strong negative influence of invasive species on the abundance of aquatic communities, particularly macrophytes, zooplankton and fish. In contrast, there was no general evidence for a decrease in species diversity in invaded habitats, suggesting a time lag between rapid abundance changes and local extinctions. Invaded habitats showed increased water turbidity, nitrogen and organic matter concentration, which are related to the capacity of invaders to transform habitats and increase eutrophication. The expansion of invasive macrophytes caused the largest decrease in fish abundance, the filtering activity of filter collectors depleted planktonic communities, omnivores (including both facultative and obligate herbivores) were responsible for the greatest decline in macrophyte abundance, and benthic invertebrates were most negatively affected by the introduction of new predators. These impacts were relatively consistent across habitats and experimental approaches. Based on our results, we propose a framework of positive and negative links between invasive species at four trophic positions and the five different components of recipient communities. This framework incorporates both direct biotic interactions (predation, competition, grazing) and indirect changes to the water physicochemical conditions mediated by invaders (habitat alteration). Considering the strong trophic links that characterize aquatic ecosystems, this framework is relevant to anticipate the far‐reaching consequences of biological invasions on the structure and functionality of aquatic ecosystems.     

Mechanisms regulating zooplankton populations in a high-mountain lake

SUMMARY 1. We studied the seasonal succession of phyto- and zooplankton and the potential impact of predation by salmonids on zooplankton population dynamics in a high-mountain Swiss lake.
2. A comparison of patterns in the abundance, body length, fecundity and age structure in the Daphnia galeata population strongly suggests that trout predation had little impact on the population and was not the cause for a decline in summer.
3. The dominance in the lake of adult trout that feed mainly on benthic prey may buffer the effect of predation on the larger zooplankton. Further, the relatively high amount of phytoplankton after spring thaw could be important for sustaining the Daphnia population under moderate fish predation.
4. Partial correlation analyses proved circumstantial evidence for both exploitative and interference competition between some zooplankton taxa. D. galeata depressed performance of other plankton species through exploitative competition.
5. Our study shows that the impact of fish on zooplankton in high-mountain lakes depends strongly on food web structure and trophic state of the lake. Where fish predation is weak, invertebrate predation combined with competition for food may be responsible for the dominance of large-bodied zooplankton species.