Fish play a key role in the trophic dynamics of lakes, not least in shallow systems. With climate warming, complex changes in fish community structure may be expected owing to the direct and indirect effects of temperature, and indirect effects of eutrophication, water-level changes and salinisation on fish metabolism, biotic interactions and geographical distribution. We review published and new data supporting the hypotheses that, with a warming climate, there will be changes in: fish community structure (e.g. higher or lower richness depending on local conditions); life history traits (e.g. smaller body size, shorter life span, earlier and less synchronised reproduction); feeding mode (i.e. increased omnivory and herbivory); behaviour (i.e. stronger association with littoral areas and a greater proportion of benthivores); and winter survival. All these changes imply higher predation on zooplankton and macroinvertebrates with increasing temperatures, suggesting that the changes in the fish communities partly resemble, and may intensify, the effects triggered by eutrophication. Modulating factors identified in cold and temperate systems, such as the presence of submerged plants and winter ice cover, seem to be weaker or non-existent in warm(ing) lakes. Consequently, in the future lower nutrient thresholds may be needed to obtain clear-water conditions and good ecological status in the future in currently cold or temperate lakes. Although examples are still scarce and more research is needed, we foresee biomanipulation to be a less successful restoration tool in warm(ing) lakes without a strong reduction of the nutrient load.
Shallow lakes, the most abundant lake type in the world, are very sensitive to climatic changes. The structure and functioning of shallow lakes are greatly impacted by submerged plants, and these may be affected by climate warming in various, contrasting, ways.Following a space-for-time substitution approach, we aimed to analyse the role of aquatic (submerged and free-floating) plants in shallow lakes under warm climates. We introduced artificial submerged and free-floating plant beds in five comparable lakes located in the temperate zone (Denmark, 55-571N) and in the subtropical zone (Uruguay, 30-351S), with the aim to study the structure and dynamics of the main associated communities.Regardless of differences in environmental variables, such as area, water transparency and nutrient status, we found consistent patterns in littoral community dynamics and structure (i.e. densities and composition of fish, zooplankton, macroinvertebrates, and periphyton) within, but substantial differences between, the two regions. Subtropical fish communities within the macrophyte beds exhibited higher diversity, higher density, smaller size, lower relative abundance of potentially piscivores, and a preference for submerged plants, compared with otherwise similar temperate lakes. By contrast, macroinvertebrates and cladocerans had higher taxon richness and densities, and periphyton higher biomass, in the temperate lakes. Several indicators suggest that the fish predation pressure was much stronger among the plants in the subtropical lakes. The antipredator behaviour of cladocerans also differed significantly between climate zones. Submerged and free-floating plants exerted different effects on the spatial distribution of the main communities, the effects differing between the climate zones. In the temperate lakes, submerged plants promoted trophic interactions with potentially positive cascading effects on water transparency, in contrast to the free-floating plants, and in strong contrast to the findings in the subtropical lakes.The higher impact of fish may result in higher sensitivity of warm lakes to external changes (e.g. increase in nutrient loading or water level changes). The current process of warming, particularly in temperate lakes, may entail an increased sensitivity to eutrophication, and a threat to the high diversity, clear water state.
1. Structural complexity may stabilise predator-prey interactions and affect the outcome of trophic cascades by providing prey refuges. In deep lakes, vulnerable zooplankton move vertically to avoid fish predation. In contrast, submerged plants often provide a diel refuge against fish predation for large-bodied zooplankton in shallow temperate lakes, with consequences for the whole ecosystem. 2. To test the extent to which macrophytes serve as refuges for zooplankton in temperate and subtropical lakes, we introduced artificial plant beds into the littoral area of five pairs of shallow lakes in Uruguay (30°-35°S) and Denmark (55°-57°N). We used plants of different architecture (submerged and free-floating) along a gradient of turbidity over which the lakes were paired. 3. We found remarkable differences in the structure (taxon-richness at the genus level, composition and density) of the zooplankton communities in the littoral area between climate zones. Richer communities of larger-bodied taxa (frequently including Daphnia spp.) occurred in the temperate lakes, whereas small-bodied taxa characterised the subtropical lakes. More genera and a higher density of benthic/plant-associated cladocerans also occurred in the temperate lakes. The density of all crustaceans, except calanoid copepods, was significantly higher in the temperate lakes (c. 5.5-fold higher). 4. Fish and shrimps (genus Palaemonetes) seemed to exert a stronger predation pressure on zooplankton in the plant beds in the subtropical lakes, while the pelagic invertebrate Chaoborus sp. was slightly more abundant than in the temperate lakes. In contrast, plantassociated predatory macroinvertebrates were eight times more abundant in the temperate than in the subtropical lakes. 5. The artificial submerged plants hosted significantly more cladocerans than the freefloating plants, which were particularly avoided in the subtropical lakes. Patterns indicating diel horizontal migration were frequently observed for both overall zooplankton density and individual taxa in the temperate, but not the subtropical, lakes. In contrast, patterns of diel vertical migration prevailed for both the overall zooplankton and for most individual taxa in the subtropics, irrespective of water turbidity. 1009 6. Higher fish predation probably shapes the general structure and dynamics of cladoceran communities in the subtropical lakes. Our results support the hypothesis that horizontal migration is less prevalent in the subtropics than in temperate lakes, and that no predatoravoidance behaviour effectively counteracts predation pressure in the subtropics. Positive effects of aquatic plants on water transparency, via their acting as a refuge for zooplankton, may be generally weak or rare in warm lakes.
An experiment in >1000 river and riparian sites found spatial patterns and controls of carbon processing at the global scale.
Summary 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.
Several studies have demonstrated a latitudinal gradient in the proportion of omnivorous fish species (that is, consumers of both vegetal and animal material) in marine ecosystems. To establish if this global macroecological pattern also exists in fresh and brackish waters, we compared the relative richness of omnivorous fish in freshwater, estuarine, and marine ecosystems at contrasting latitudes. Furthermore, we sought to determine the main environmental correlates of change in fish omnivory. We conducted a meta-analysis of published data focusing on change in the relative richness of omnivorous fishes in native fish communities along a broad global latitudinal gradient, ranging from 41°S to 81.5 N°including all continents except for Antarctica. Data from streams, rivers, lakes, reservoirs, estuaries, and open marine waters (ca. 90 papers covering 269 systems) were analyzed. Additionally, the relationship between the observed richness in omnivory and key factors influencing trophic structure were explored. For all ecosystems, we found a consistent increasing trend in the relative richness of omnivores with decreasing latitude. Furthermore, omnivore richness was higher in freshwaters than in marine ecosystems. Our results suggest that the observed latitudinal gradient in fish omnivory is a global ecological pattern occurring in both freshwater and marine ecosystems. We hypothesize that this macroecological pattern in fish trophic structure is, in part, explained by the higher total fish diversity at lower latitudes and by the effect of temperature on individual food intake rates; both factors ultimately increasing animal food limitation as the systems get warmer.
Summary 1. Small cladocerans, copepod nauplii and rotifers often dominate the zooplankton community in tropical and subtropical lakes. This is probably because of high predation pressure by small omnivorous–planktivorous fish, but experimental evidence is scarce. 2. This study used two approaches to test the effect of the small omnivorous–planktivorous fish species Jenynsia multidentata, which is frequently abundant in (sub)tropical eutrophic lakes in South America, on the size distribution of zooplankton. In Lake Blanca (Uruguay), which lacks any piscivores, we sampled seasonally for both fish and zooplankton. We also conducted an outdoor mesocosm experiment with treatments containing or lacking J. multidentata. 3. Together, the empirical and experimental data suggest that J. multidentata predation plays an important role in modulating the size structure of the zooplankton community in subtropical lakes. In the absence of J. multidentata, stocked large‐sized zooplankters like Daphnia obtusa were abundant in the experiments, while small‐sized zooplankton dominated in the presence of fish, as they did in the lake itself from spring to the end of the season.
Phytoplankton abundance and biomass can be explained as a result of spatial and temporal changes in physical and biological variables, and also by the externally imposed or self-generated spatial segregation. In the present study, we analyzed contrasting-season changes in the phytoplankton communities of five subtropical shallow lakes, covering a nutrient gradient from oligo-mesotrophy to hypereutrophy, using a morphologically based functional approach to cluster the species. Six environmental variables accounted for 46% of the total phytoplankton morphological groups variance, i.e., turbidity (Secchi disk), conductivity, total phosphorus, total nitrogen, total zooplankton abundance, and herbivorous meso:microzooplankton density ratio. The differences in resource availability and zooplankton abundance among the systems were related with important changes in phytoplankton composition and structure. Within phytoplankton assemblages, adaptations to improve both light and phosphorus/nitrogen uptake were important in nutrient-poor systems; while grazing-avoidance mechanisms, such as colonial forms or bigger individuals, seemed relatively important only in eutrophic Lake Blanca, where light was not a limiting factor. However, this was not observed in the nutrient-rich Lake Cisne, where low light availability (due to clay resuspension and dark water color) was identified as the main structuring factor. Our results suggest that the composition of phytoplankton morphologically based functional groups appear to reliably describe the trophic sate of the lakes. However, other factors, such as nonbiological turbid condition, or zooplankton composition, may interact rendering interpretations difficult, and therefore, deserve further studies and evaluation.
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