Homogenization of fish assemblages in different lake depth strata at local and regional scales

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Lake size,macrophytes, and omnivory contribute to food web linkage in temperate shallow eutrophic lakes

Physical, chemical and biological processes facilitate cross-habitat connections in lakes, prompting food webs to be supported by different subsidies. We tested the hypothesis that the pelagic food web is subsidized by littoral resources and fish foraging behaviour plays a major role in carbon flux and on food web structure in shallow hypereutrophic lakes. We performed a fish diet and carbon and nitrogen isotope analyses to predict the linkage between littoral and pelagic habitats in three shallow hypereutrophic lakes. Lakes differed in morphology, fetch, macrophyte composition and width of the littoral zone. δ¹³C signals of seston differed among lakes, but were similar to other producers. Macroinvertebrates and fish carbon signatures were more enriched in the lake co-dominated by emergent and submerged vegetation. Fish foraging behaviour indicates that more than the 80% of the carbon that sustain adult fish was channelled from the littoral. In conclusion, littoral carbon were relevant and sustain, in part, food web in these shallow lakes. Factors like the extension of the littoral zone, lake morphometry, and the dominance of multi-chain omnivorous fish facilitate the connection among lake compartments and the transference of littoral carbon to lake food web.     

Stable isotopes indicate that zebra mussels (Dreissena polymorpha) increase dependence of lake food webs on littoral energy sources

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Invasive macrophyte effects on littoral trophic structure and carbon sources

Limited data from terrestrial ecosystems suggest that invasive species can affect energy flow and nutrient cycling in invaded systems. This is likely also true for aquatic ecosystems, yet little information is available on food web effects of invasive macrophytes. This study examined the effects of dominant invasive Eurasian watermilfoil on lake trophic structure and energy flow. Stable isotopes of carbon and nitrogen were used to compare trophic structure in invaded and uninvaded lakes and macrophyte stands. Contribution of native and invasive macrophytes, their epiphyton and detritus to the upper trophic level of lacustrine food webs was partitioned using mixing models. Carbon isotope values of macroinvertebrate consumers were similar to macrophyte-associated production in stands from which they were collected. However, contribution of Eurasian watermilfoil and its epiphyton to higher trophic level was negligible, and littoral fish derived most of their energy from sources associated with native macrophytes, despite their lower abundance. This means that littoral fish may depend on the remaining patches of native macrophytes in lakes invaded by non-native plants. Considering previous findings, these results show that the assessment of ecosystem-level processes is needed to understand the entire range of impacts of invasive species.     

Response of a shallow Mediterranean lake to nutrient diversion: does it follow similar patterns as in northern shallow lakes?

1. In view of the paucity of data on the response of warm shallow lakes to reductions in nutrient loading, this paper presents a long‐term limnological data set to document changes in the food‐web of a shallow Mediterranean lake (Lake Albufera, Valencia, Spain) that has experienced reductions in phosphorus (P) (77%) and nitrogen (N) (24%) loading following sewage diversion. 2. Nine years after sewage diversion, P concentration in the lake was reduced by 30% but remained high (TP = 0.34 mg L^?1), although the mean water retention time in the lake was only 0.1 years. Nitrate concentrations did not significantly change, probably because the lake continued to receive untreated effluents from ricefields. 3. Chlorophyll a concentration was reduced by half (annual mean of 180 μg L^?1). Cyanobacteria abundance remained high but its composition changed towards smaller species, both filamentous and chroococcal forms. 4. Cladocera abundance increased and reached peaks twice a year (December to March and July to September). After nutrient reduction, short‐term clear‐water phases (up to 5 weeks) occurred during February to March in several years, concomitant with annual flushing of the lake and lower fish densities. The abundance of Cladocera in winter contrasted with the spring peaks observed in northern restored shallow lakes. The zooplankton to phytoplankton biomass ratio remained lower than in northern temperate shallow lakes, probably because of fish predation on zooplankton. 5. Improvement of the water quality of Lake Albufera remained insufficient to counteract littoral reed regression or improve underwater light allowing submerged plants re‐colonise the lake. 6. Sewage diversion from Lake Albufera impacted the food web through the plankton, but higher trophic levels, such as fish and waterfowl, were affected to a lesser degree. Although the fish species present in the lake are mainly omnivorous, long‐term data on commercial fish captures indicated that fish communities changed in response to nutrient level and trophic structure as has been observed in restored shallow lakes at northern latitudes. 7. Phosphorus concentrations produced similar phytoplankton biomass in Lake Albufera as in more northern shallow lakes with abundant planktivorous fish and small zooplankton. However, in Lake Albufera, high average concentrations were maintained throughout the year. Overall, results suggest that nutrient control may be a greater priority in eutrophicated warm shallow lakes than in similar lakes at higher latitudes.     

Substantial differences in littoral fish community structure and dynamics in subtropical and temperate shallow lakes

1. Fish play a key role in the functioning of temperate shallow lakes by affecting nutrient exchange among habitats as well as lake trophic structure and dynamics. These processes are, in turn, strongly influenced by the abundance of submerged macrophytes, because piscivorous fish are often abundant at high macrophyte density. Whether this applies to warmer climates as well is virtually unknown. 2. To compare fish community structure and dynamics in plant beds between subtropical and temperate shallow lakes we conducted experiments with artificial submerged and free‐floating plant beds in a set of 10 shallow lakes in Uruguay (30°–35°S) and Denmark (55°–57°N), paired along a gradient of limnological characteristics. 3. The differences between regions were more pronounced than differences attributable to trophic state. The subtropical littoral fish communities were characterised by higher species richness, higher densities, higher biomass, higher trophic diversity (with predominance of omnivores and lack of true piscivores) and smaller body size than in the comparable temperate lakes. On average, fish densities were 93 ind. m⁻² (±10 SE) in the subtropical and 10 ind. m⁻² (±2 SE) in the temperate lakes. We found a twofold higher total fish biomass per unit of total phosphorus in the subtropical than in the temperate lakes, and as fish size is smaller in the former, the implication is that more energy reaches the littoral zone fish community of the warmer lakes. 4. Plant architecture affected the spatial distribution of fish within each climate zone. Thus, in the temperate zone fish exhibited higher densities among the artificial free‐floating plants while subtropical fish were denser in the artificial submerged plant beds. These patterns appeared in most lakes, regardless of water turbidity or trophic state. 5. The subtropical littoral fish communities resembled the fish communities typically occurring in temperate eutrophic and hypertrophic lakes. Our results add to the growing evidence that climate warming may lead to more complex and omnivory‐dominated food webs and higher density and dominance of smaller‐sized fish. This type of community structure may lead to a weakening of the trophic cascading effects commonly observed in temperate shallow lakes and a higher risk of eutrophication.     

Do environmental differences between lakes in northwestern Argentinean Patagonia affect the infection of Philureter trigoniopsis (monogenea) in Galaxias maculatus (osmeriformes)?

Philureter trigoniopsis is the only parasite found in the ureters and urinary bladder of Galaxias maculatus in Patagonian Andean lakes. The dynamics of this endoparasitic monogenean were studied in Lake Gutiérrez, a body of water with scarce shoreline vegetation, where the host has an annual cycle of migration to the deep pelagic zone of the lake. To compare variations of the infection related to differences between lakes, G. maculatus specimens were sampled monthly with baited traps from September 1998 to November 1999 in Lake Moreno, which is an oligotrophic body of water with emergent shoreline vegetation and where the fish do not migrate to the deep pelagic areas of the lake. In addition, data for summer infections of P. trigoniopsis from 10 Andean Patagonian lakes that differ in aquatic vegetation, depth, and area were compared. In Lake Moreno prevalence of P. trigoniopsis showed a seasonal pattern, with 1-yr-old fish exhibiting the highest values of prevalence and mean intensity. Negative correlations between water temperature and prevalence and between age of fish and abundance were found. Our results suggest that age of fish may be the main factor structuring the distribution of P. trigoniosis in populations of G. maculatus. At the regional level the relationship between the infection and the characteristics of the lakes was also observed, with prevalence and mean intensity of P. trigoniopsis in G. maculatus higher in large deep lakes without macrophytes.     

Some like it deep: Intraspecific niche segregation in ruffe (Gymnocephalus cernua)

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OPINION Manipulating lake community structure: where do we go from here?

SUMMARY. 1 More than 10 years experience with whole lake pelagic manipulation has suggested some general trends applicable to all freshwater pelagic communities and some specific trends related to lake depth.
2 Among the general trends is the observation that the trophic cascade is strongly damped. This means that changes in phytoplankton biomass can be assured only when the fish community is strongly manipulated.
3 Among the depth related trends is the observation that in shallow lakes, changes in fish community structure are more likely to have cascading impacts on phytoplankton than are changes in deep lakes.
4 In shallow lakes, fish removal frequently results in decreased turbidity which is associated with the development of dense macrophyte populations and significant reductions of algal standing stocks. The mechanisms involve: increased grazing by zooplankton, the removal of fish induced bioturbation and nutrient recycling, and direct and indirect macrophyte effects (shading, zooplankton refuges and competition for nutrients).
5 In shallow lakes, where planktivore biomass can be regulated and macrophyte development is acceptable, fish biomanipulalions are likely to result in reduced algal populations and improved water quality.
6 In deep lakes, where macrophytes are not as important, long-term effects of fish manipulations are strongly dependent upon the probability of non-grazable algal bloom development. This is determined by many factors (chemical, physical and grazer related) which modify the impact that grazers have on phytoplankton biomass.
7 In deep lakes, successful fish biomanipulations may only be effective when chemical and physical factors are altered to produce algal species compositions that permit strong top-down control of prey by predators.     

Diatom community succession in the recent history of a eutrophic Yunnan Plateau lake, Lake Dianchi, in subtropical China

Fish introduction, eutrophication and disappearance of aquatic vegetation are important disturbances of aquatic ecosystems, especially in plateau lakes, which are generally considered to be very vulnerable. Fish were introduced to Lake Dianchi, a eutrophic plateau lake in southwest China, in the late 1950s and 1970s. After the introduction, invasive fish became the dominant species, and the total fish yield increased. Meanwhile, the trophic level of Lake Dianchi had a tendency to increase in the past decades because of the increases in human activities in the watershed area. In addition, the area of aquatic vegetation decreased from more than 90 to 1.8% of the lake area from the 1950s to 2000. This study investigated the effects of fish introduction, eutrophication and aquatic vegetation on the diatom community of Lake Dianchi by examining the changes of microfossil diatom assemblage and abundance. Results showed that the absolute abundance and diatom assemblages changed after fish were introduced. The endemic species, Cyclotella rohomboideo-elliptica, disappeared with the introduction of fish and increasing trophic levels after 1958. Fragilaria crotonensis entered into the lake with the introduction of fish and gradually thrived in the lake after 1958. Diatom species numbers also decreased gradually from 21 to 9 from the past to present. Epiphytic diatoms disappeared with the decrease of aquatic vegetation after 1985. Our study indicated that eutrophication was the most important process determining diatom abundance, and fish introduction was a secondary process determining diatom abundance, while aquatic vegetation had a more important role in structuring the diatom community in this eutrophic plateau lake.     

滇中高原湖泊鱼类多样性的研究

依据历年馆藏的鱼类标本和资料,采用多元逐步回归与多项式逐步回归的分析方法,系统分析了滇中6个高原湖泊鱼类的物种丰度、特有种以及属的数目与7个主要湖泊环境因子之间的关系。分析结果显示,控制湖泊鱼类物种丰度、特有种数和属数的关键湖泊环境因子是湖泊面积和湖岸线长度;其中与湖泊面积呈显著正相关,而与湖岸线长度呈负相关。分析不停留于这一常规的横向比较,而是从历史生物地理学的角度,更深入一步揭示了滇中6个湖泊鱼类多样性的演化实质上是受湖泊的发育阶段所控制的。     

Depth gradients in food‐web processes linking habitats in large lakes: Lake Superior as an exemplar ecosystem

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Water‐level fluctuations regulate the structure and functioning of natural lakes

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Horizontal distribution of cladocerans in arctic Greenland lakes – impact of macrophytes and fish

Submerged macrophytes may play an important role as a refuge for zooplankton against predators. However, a recent study suggests that their importance depends on the trophic state of the lake. We studied the impact of fish and macrophytes on the horizontal distribution of pelagic cladocerans in 56 oligotrophic arctic Greenland lakes. In north-east and western Greenland, zooplankton was sampled in the near-shore (littoral) and central (pelagial) part of all lakes and fish were sampled with multiple mesh-sized gill nets. Macrophytes were visually estimated in the littoral. In north-east Greenland, 5 taxa of cladocerans were found, while 14 taxa were recorded in western Greenland. Daphnia pulex occurred only in fishless lakes in both northeast and western Greenland and avoided the near-shore areas in the shallow and deep lakes. Bosmina spp. and Holopedium gibberum were evenly distributed between the littoral and the pelagial in the deep and shallow fishless lakes. However, their near-shore density was lowest in the presence of fish. Macrophyte-related and benthic cladocerans concentrated either in the littoral or were evenly distributed between the littoral and the pelagial, irrespective of depth and fish presence or absence. Macrophytes had no impact on the horizontal distribution of pelagic cladocerans. Thus, it is concluded that horizontal heterogeneity of Bosmina spp. and Holopedium gibberum might be affected by the presence of fish.     

Relationships between fish feeding guild and trophic structure in English lowland shallow lakes subject to anthropogenic influence: implications for lake restoration

The shallow lakes of Eastern England have been subject to intense anthropogenic pressures including nutrient enrichment and fish stocking. We sought to determine the relationships between fish community structure and other ecosystem characteristics in 28 of these lakes through classification of fish species into piscivorous, zooplanktivorous and benthivorous feeding guilds according to the literature. Canonical correspondence analysis produced clear associations between fish and ecosystem characteristics that generally agreed with other theoretical (e.g. the alternative stable states hypothesis) and empirical studies, but with some important differences. There was a striking lack of relationships between nutrients and other variables, indicating the importance of top-down rather than bottom-up processes as a structuring force in the generally eutrophic study lakes. The presence of submerged (and shoreline) vegetation was associated with a diverse assemblage of apparently co-existing piscivorous (principally pike Esox lucius) and zooplanktivorous species. Perch Perca fluviatilis, a significant predator in other studies, was unimportant and argued to be limited by water quality in the extremely shallow lakes. In contrast, the benthivorous fish guild (principally carp Cyprinus carpio, bream Abramis brama and tench Tinca tinca) essentially represented the inverse of the potential pelagic associations between piscivores/zooplanktivores and vegetation. The introduction of large benthivores to many study lakes could have precipitated a loss of submerged vegetation through direct uprooting during foraging, with the effect of simplifying the fish community being most acute where littoral vegetation was limited by other anthropogenic factors. It is implied that attempts to promote or restore submerged vegetation in these lakes would best target benthivorous species.     

Behavioral and trophic plasticity of juvenileTilapia mossambica in utilization of the unstable littoral habitat

Synopsis JuvenileTilapia mossambica Peters (2.5–10.0 cm standard length) in Lake Sibaya, South Africa, move daily from deep offshore waters to shallow (<0.5 m) littoral areas where they feed for several hours and then return to deep water. The timing of these movements varies in response to changing physical and biological features of the littoral environment. During this study (December 1973 – February 1976), lake level rose more than 1 m. At the start of the study, littoral areas visited by juvenileT. mossambica were free of vegetation and debris, but at higher lake levels trees and brush from the eroding shoreline were common in nearshore littoral waters. When the littoral zone was free of debris, juvenileT. mossambica visited nearshore waters only during daylight hours. After a 0.65 m rise in lake level and considerable accumulation of debris, these fish were abundant in the littoral zone only at night. This reversal can be attributed to efforts by juvenileT. mossambica to avoid their principal predator, the catfishClarias gariepinus. At low lake levelC. gariepinus ventured into the littoral only at night but at higher lake levels this predator was found within flooded vegetation during the day. When further increases in lake level flooded marginal grasslands, juvenileT. mossambica abandoned the littoral zone in favor of this newly created eulittoral habitat where potential fish predators did not occur. With the change in habitat, the diet changed from benthic detrital aggregate to periphyton. These observations suggest the importance of behavioral and throphic plasticity in the ability of tilapia to utilize unstable habitats.     

Movement of plankton through lake-stream systems

1. River plankton are often assumed to come from upstream lakes, but the factors controlling the movement of plankton between lakes and rivers into outflow streams are unclear. We tested the possibility that the physical structure of the littoral zone near the lake outlet (depth, presence of macrophytes) and diurnal differences in plankton composition at the lake surface influence the movement of plankton from the lake into the stream and determine their persistence downstream. 2. Zooplankton and phytoplankton biomass, community composition and mean body size were compared between two deep lakes without macrophytes at the lake edge and two shallow lakes with macrophytes at the lake edge. Samples were collected day and night on three dates, in the lake centre, in the littoral zone adjacent to the lake outlet, at the outlet and at two sites downstream in Algonquin Park, Ontario, Canada. 3. The morphology of lake edges clearly affects the movement of lake zooplankton into outlet streams. Outlets draining deeper littoral zones had higher zooplankton biomass than shallow littoral outlets (P < 0.0001), but these differences disappeared within 50 m downstream of the lake. There was no difference in mean zooplankton body size among lake outlets or between littoral and outlet samples. However, shallow littoral zones were dominated by cyclopoid copepods and deeper littoral zones were dominated by Bosmina longirostris. In contrast, phytoplankton biomass entering the outlet was similar to that found within the lake and did not vary with lake outlet morphology. These effects were consistent across several sampling weeks and were not affected by surface zooplankton biomass changes associated with diurnal vertical migration in the lake centre. 4. A comparison with published river zooplankton data suggests that zooplankton are rapidly eliminated from shallow outlet streams (≤1 m deep) but persist in most deeper outlet rivers (≥2 m deep). Because the depth of an outlet river determines downstream zooplankton community development, the contribution of lakes to river plankton communities may be influenced by the location of each lake within the drainage basin. These findings suggest that lake and outflow physical structure influences connection strength between spatially successive habitats.     

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.