Extrinsic and intrinsic controls of zooplankton diversity in lakes

Pelagic crustacean zooplankton were collected from 336 Norwegian lakes covering a wide range of latitude, altitude, lake area, mean depth, production (as chlorophyll a), and fish community structure. Mean zooplankton species richness during the ice-free season was generally low at high latitudes and altitudes. Further, lower species richness was recorded in western lakes, possibly reflecting constraints on migration and dispersal. However, despite obvious spatial limitations, geographic boundaries were only weak predictors of mean zooplankton richness. Similarly, lake surface area did not contribute positively to mean richness such as seen in other ecosystem surveys. Rather, intrinsic factors such as primary production and fish community (planktivore) structure were identified by regression analysis as the major predictors of zooplankton diversity, while a positive correlation was observed between species richness and total zooplankton biomass. However, in spite of a large number of variables included in this study, the predictive power of multiple regression models was modest (<50% variance explained), pointing to a major role for within-lake properties, as yet unidentified intrinsic forces, stochasticity, or dispersal as constraints on zooplankton diversity in these lakes.     

Zooplankton biodiversity and lake trophic state: explanations invoking resource abundance and distribution

While empirical studies linking biodiversity to local environmental gradients have emphasized the importance of lake trophic status (related to primary productivity), theoretical studies have implicated resource spatial heterogeneity and resource relative ratios as mechanisms behind these biodiversity patterns. To test the feasibility of these mechanisms in natural aquatic systems, the biodiversity of crustacean zooplankton communities along gradients of total phosphorus (TP) as well as the vertical heterogeneity and relative abundance of their phytoplankton resources were assessed in 18 lakes in Quebec, Canada. Zooplankton community richness was regressed against TP, the spatial distribution of phytoplankton spectral groups, and the relative biomass of spectral groups. Since species richness does not adequately capture ecological function and life history of different taxa, features which are important for mechanistic theories, relationships between zooplankton functional diversity (FD) and resource conditions were examined. Zooplankton species richness showed the previously established tendency to a unimodal relationship with TP, but functional diversity declined linearly over the same gradient. Changes in zooplankton functional diversity could be attributed to changes in both the spatial distribution and type of phytoplankton resource. In the studied lakes, spatial heterogeneity of phytoplankton groups declined with TP, even while biomass of all groups increased. Zooplankton functional diversity was positively related to increased heterogeneity in cyanobacteria spatial distribution. However, a smaller amount of variation in functional diversity was also positively related to the ratio of biomass in diatoms/chrysophytes to cyanobacteria. In all observed relationships, a greater variation of functional diversity than species richness measures was explained by measured factors, suggesting that functional measures of zooplankton communities will benefit ecological research attempting to identify mechanisms behind environmental gradients affecting diversity.     

Biogeography of bacterioplankton in lakes and streams of an Arctic tundra catchment

Bacterioplankton community composition was compared across 10 lakes and 14 streams within the catchment of Toolik Lake, a tundra lake in Arctic Alaska, during seven surveys conducted over three years using denaturing gradient gel electrophoresis (DGGE) of PCR-amplified rDNA. Bacterioplankton communities in streams draining tundra were very different than those in streams draining lakes. Communities in streams draining lakes were similar to communities in lakes. In a connected series of lakes and streams, the stream communities changed with distance from the upstream lake and with changes in water chemistry, suggesting inoculation and dilution with bacteria from soil waters or hyporheic zones. In the same system, lakes shared similar bacterioplankton communities (78% similar) that shifted gradually down the catchment. In contrast, unconnected lakes contained somewhat different communities (67% similar). We found evidence that dispersal influences bacterioplankton communities via advection and dilution (mass effects) in streams, and via inoculation and subsequent growth in lakes. The spatial pattern of bacterioplankton community composition was strongly influenced by interactions among soil water, stream, and lake environments. Our results reveal large differences in lake-specific and stream-specific bacterial community composition over restricted spatial scales (<10 km) and suggest that geographic distance and connectivity influence the distribution of bacterioplankton communities across a landscape.     

Independent gradients of producer, consumer, and microbial diversity in lake plankton

Interactions between trophic levels during food web assembly can drive positive correlations in diversity between producers, consumers, and decomposers. However, the contribution of trophic interactions relative to local environmental factors in promoting species diversity is poorly understood, with many studies only considering two trophic levels. Here we examine correlations in diversity among zooplankton, phytoplankton, and bacteria in the pelagic zone of 31 lakes in British Columbia, Canada. We sampled species diversity of zooplankton and phytoplankton through morphological identification, and bacterial genetic diversity was estimated by denaturing gradient gel electrophoresis (DGGE) of 16S rDNA polymorphisms. We looked for correlations in diversity that were independent of the abiotic environment by statistically controlling for 18 limnological variables. No significant correlations were found between the diversity of zooplankton, phytoplankton, and bacteria. In addition, the physical factors that were associated with species composition in one trophic level were independent of those that were important for another. Our results provide no support for the importance of direct feedbacks between producers, consumers, and decomposers in maintaining diversity. Zooplankton, phytoplankton, and bacterial diversity and composition are regulated independently from one another and respond to different environmental variables. These results suggest that species of lake plankton show loose trophic associations with one another due to broad diets in consumers and decomposers.     

Metacommunity assembly and sorting in newly formed lake communities

new habitats are created, community assembly may follow independent trajectories, since the relative importance of dispersal limitation, priority effects, species interactions, and environmental gradients can vary as assembly proceeds. Unfortunately, tracking community colonization and composition across decades is challenging. We compiled a multiyear community composition data set and reconstructed past communities with remains from sediment cores to investigate cladoceran assembly dynamics in six older (1920s) and two more recently formed (1950s) lakes. We found that current communities cluster along a gradient of thermal stratification that is known to influence predation intensity. Assembling communities showed evidence for a greater influence of species sorting and a reduced influence of spatial structure since the first colonizations. However, lake community trajectories varied considerably, reflecting different colonization sequences among lakes. In the older lakes, small-bodied cladocerans often arrived much earlier than large-bodied cladocerans, while the two younger lakes were colonized much more rapidly, and one was quickly dominated by a large-bodied species. Thus, by combining contemporary community data with paleoecological records, we show that assembly history influences natural community structure for decades while patterns of ecological sorting develop.     

Species dispersal rates alter diversity and ecosystem stability in pond metacommunities

Metacommunity theory suggests that relationships between diversity and ecosystem stability can be determined by the rate of species dispersal among local communities. The predicted relationships, however, may depend upon the relative strength of local environmental processes and disturbance. Here we evaluate the role of dispersal frequency and local predation perturbations in affecting patterns of diversity and stability in pond plankton metacommunities. Pond metacommunities were composed of three mesocosm communities: one of the three communities maintained constant "press" predation from a selective predator, bluegill sunfish (Lepomis macrochirus); the second community maintained "press" conditions without predation; and the third community experienced recurrent "pulsed" predation from bluegill sunfish. The triads of pond communities were connected at either no, low (0.7%/d), or high (20%/d) planktonic dispersal. Richness and composition of zooplankton and stability of plankton biomass and ecosystem productivity were measured at local and regional spatial scales. Dispersal significantly affected diversity such that local and regional biotas at the low dispersal rate maintained the greatest number of species. The unimodal local dispersal-diversity relationship was predator-dependent, however, as selective press predation excluded species regardless of dispersal. Further, there was no effect of dispersal on beta diversity because predation generated local conditions that selected for distinct community assemblages. Spatial and temporal ecosystem stability responded to dispersal frequency but not predation. Low dispersal destabilized the spatial stability of producer biomass but stabilized temporal ecosystem productivity. The results indicate that selective predation can prevent species augmentation from mass effects but has no apparent influence on stability. Dispersal rates, in contrast, can have significant effects on both species diversity and ecosystem stability at multiple spatial scales in metacommunities.     

Native fish diversity alters the effects of an invasive species on food webs

Aquatic communities have been altered by invasive species, with impacts on native biodiversity and ecosystem function. At the same time, native biodiversity may mitigate the effects of an invader. Common carp (Cyprinus carpio) is a ubiquitous, invasive fish species that strongly influences community and ecosystem processes. We used common carp to test whether the potential effects of an invasive species are altered across a range of species diversity in native communities. In mesocosms, treatments of zero, one, three, and six native fish species were used to represent the nested subset patterns observed in fish communities of lakes in Illinois, USA. The effect of the invader was tested across fish richness treatments by adding common carp to the native community and substituting native biomass with common carp. Native species and intraspecific effects reduced invader growth. The invader reduced native fish growth; however, the negative effect was minimized with increasing native richness. The zooplankton grazer community was modified by a top-down effect from the invader that increased the amount of phytoplankton. Neither the invader nor richness treatments influenced total phosphorus or community metabolism. Overall, the invader reduced resources for native species; and the effect scaled with how the invader was incorporated into the community. Higher native diversity mitigated the impact of the invader, confirming the need to consider biodiversity when predicting the impacts of invasive species.     

Spectral analysis unmasks synchronous and compensatory dynamics in plankton communities

Community biomass is often less variable than the biomasses of populations within the community, yet attempts to implicate compensatory dynamics between populations as a cause of this relationship often fail. In part, this may be due to the lack of appropriate metrics for variability, but there is also great potential for large-scale processes such as seasonality or longer-term environmental change to obscure important dynamics at other temporal scales. In this study, we apply a scale-resolving method to long-term plankton data, to identify the specific temporal scales at which community-level variability is influenced by synchrony or compensatory dynamics at the population level. We show that variability at both the population and community level is influenced strongly by a few distinct temporal scales: in phytoplankton, ciliate, rotifer, and crustacean communities, synchronous dynamics are predominant at most temporal scales. However, in phytoplankton and crustacean communities, compensatory dynamics occur at a sub-annual scale (and at the annual scale in crustaceans) leading to substantial reductions in community-level variability. Aggregate measures of population and community variability do not detect compensatory dynamics in these communities; thus, resolving their scale dependence unmasks dynamics that are important for community stability in this system. The methods and results presented herein will ultimately lead to a better understanding of how stability is achieved in communities.     

A vertical-compressed three-dimensional ecological model in Lake Taihu, China

A three-dimensional ecological model on the basis of the analyses of environmental characteristics is set up for Lake Taihu, one of the largest shallow lakes in China. The hydrodynamic processes, nutrient cycling, chemical processes and biological processes are integrated in the model. Model state variables include: water current, surface displacement, nutrients of nitrogen and phosphorus, as well as their different forms such as ammonia nitrogen, nitrate nitrogen, phosphate phosphorus, etc., biomasses of macroplankton, phytoplankton, zooplankton and fish, and also the nutrient levels of macroplankton and phytoplankton. A nutrient budget and sediment transformation are also coupled in the model. The data from January 17, 1997 to January 18, 1998 are use to calibrate the model. The model results have shown good agreement with the observations. It implies that the model could be used for the lake environmental management and research for examining the processes and determining the water quality. The reasons of deviations between the modelled results and the observed values are also discussed. There are six factors that explain the deviations of the modelled results from the observed values and they can be grouped into two sets. One set of problems is associated with the standard deviation introduced by sampling and analyses. The second set of problems can be solved by introduction of processes lacking in the present model (resuspension, phytoplankton transportation mode under the wind with low speed, shifts in species composition and varied size of phytoplankton and zooplankton). The latter two processes should be included in the model at a later stage by integration of a structurally dynamic approach into the three-dimensional model.     

Fish reintroductions reveal smooth transitions between lake community states

Whether communities respond smoothly or discontinuously to changing environmental conditions has important consequences for the preservation and restoration of ecosystems. Theory shows that communities may exhibit a variety of responses to environmental change, including abrupt transitions due to the existence of alternate states. However, there have been few opportunities to look for such transitions in nature. Here, we examine the impact of a two-orders-of-magnitude decrease and then increase in planktivore abundance in Wintergreen lake (Michigan, USA), caused by the extinction and reintroduction of two dominant fish species (largemouth bass, Micropterus salmoides, and bluegill, Lepomis macrochirus). Over a 16 + yr period of slow change from high planktivory to low planktivory back to high planktivory, the zooplankton community changed smoothly and predictably between states. In years of low planktivory, the zooplankton assemblage was dominated by a single, large, cladoceran species, Daphnia pulicaria, whereas in years of high planktivory, D. pulicaria disappeared and was replaced by a suite of small-bodied cladocerans. We quantified the multivariate change in zooplankton community dissimilarity and found that community state smoothly tracked changes in planktivore density in both a forward and backward direction. Thus, there was little evidence of discontinuity in this system where transitions are strongly driven by planktivory.     

Trajectories of zooplankton recovery in the Little Rock Lake whole-lake acidification experiment.

Understanding the factors that affect biological recovery from environmental stressors such as acidification is an important challenge in ecology. Here we report on zooplankton community recovery following the experimental acidification of Little Rock Lake, Wisconsin, USA. One decade following cessation of acid additions to the northern basin of Little Rock Lake (LRL), recovery of the zooplankton community was complete. Approximately 40% of zooplankton species in the lake exhibited a recovery lag in which biological recovery to reference basin levels was delayed by 1-6 yr after pH recovered to the level at which the species originally responded. Delays in recovery such as those we observed in LRL may be attributable to "biological resistance" wherein establishment of viable populations of key acid-sensitive species following water quality improvements is prevented by other components of the community that thrived during acidification. Indeed, we observed that the recovery of species that thrived during acidification tended to precede recovery of species that declined during acidification. In addition, correspondence analysis indicated that the zooplankton community followed different pathways during acidification and recovery, suggesting that there is substantial hysteresis in zooplankton recovery from acidification. By providing an example of a relatively rapid recovery from short-term acidification, zooplankton community recovery from experimental acidification in LRL generally reinforces the positive outlook for recovery reported for other acidified lakes.     

Large-scale biodiversity patterns in freshwater phytoplankton

Our planet shows striking gradients in the species richness of plants and animals, from high biodiversity in the tropics to low biodiversity in polar and high-mountain regions. Recently, similar patterns have been described for some groups of microorganisms, but the large-scale biogeographical distribution of freshwater phytoplankton diversity is still largely unknown. We examined the species diversity of freshwater phytoplankton sampled from 540 lakes and reservoirs distributed across the continental United States and found strong latitudinal, longitudinal, and altitudinal gradients in phytoplankton biodiversity, demonstrating that microorganisms can show substantial geographic variation in biodiversity. Detailed analysis using structural equation models indicated that these large-scale biodiversity gradients in freshwater phytoplankton diversity were mainly driven by local environmental factors, although there were residual direct effects of latitude, longitude, and altitude as well. Specifically, we found that phytoplankton species richness was an increasing saturating function of lake chlorophyll a concentration, increased with lake surface area and possibly increased with water temperature, resembling effects of productivity, habitat area, and temperature on diversity patterns commonly observed for macroorganisms. In turn, these local environmental factors varied along latitudinal, longitudinal, and altitudinal gradients. These results imply that changes in land use or climate that affect these local environmental factors are likely to have major impacts on large-scale biodiversity patterns of freshwater phytoplankton.     

The interplay between environmental conditions and allee effects during the recovery of stressed zooplankton communities

Many important ecological phenomena depend on the success or failure of small introduced populations. Several factors are thought to influence the fate of small populations, including resource and habitat availability, dispersal levels, interspecific interactions, mate limitation, and demographic stochasticity. Recent field studies suggest that Allee effects resulting from mate limitation can prevent the reestablishment of sexual zooplankton species following a disturbance. In this study, we explore the interplay between Allee effects and local environmental conditions in determining the population growth and establishment of two acid-sensitive zooplankton species that have been impacted by regional anthropogenic acidification. We conducted a factorial design field experiment to test the impact of pH and initial organism densities on the per capita population growth (r) of the sexual copepod Epischura lacustris and the seasonally parthenogenetic cladoceran Daphnia mendotae. In addition, we conducted computer simulations using r values obtained from our experiments to determine the probability of extinction for small populations of acid-sensitive colonists that are in the process of colonizing recovering lakes. The results of our field experiment demonstrated that local environmental conditions can moderate the impacts of Allee effects for E. lacustris: Populations introduced at low densities had a significantly lower r at pH 6 than at pH 7. In contrast, r did not differ between pH 6 and 7 environments when E. lacustris populations were introduced at high densities. D. mendotae was affected by pH levels, but not by initial organism densities. Results from our population growth simulations indicated that E. lacustris populations introduced at low densities to pH 6 conditions had a higher probability of extinction than those introduced at low densities to a pH 7 environment. Our study indicates that environmental conditions and mate limitation can interact to determine the fate of small populations of sexually reproducing zooplankton species. If a more rapid recovery of acid-damaged zooplankton communities is desired, augmentation of dispersal levels may be needed during the early phases of pH recovery in order to increase the probability of establishment for mate-limited zooplankton species.     

Intraspecific variation in a predator affects community structure and cascading trophic interactions

Intraspecific phenotypic variation in ecologically important traits is widespread and important for evolutionary processes, but its effects on community and ecosystem processes are poorly understood. We use life history differences among populations of alewives, Alosa pseudoharengus, to test the effects of intraspecific phenotypic variation in a predator on pelagic zooplankton community structure and the strength of cascading trophic interactions. We focus on the effects of differences in (1) the duration of residence in fresh water (either seasonal or year-round) and (2) differences in foraging morphology, both of which may strongly influence interactions between alewives and their prey. We measured zooplankton community structure, algal biomass, and spring total phosphorus in lakes that contained landlocked, anadromous, or no alewives. Both the duration of residence and the intraspecific variation in foraging morphology strongly influenced zooplankton community structure. Lakes with landlocked alewives had small-bodied zooplankton year-round, and lakes with no alewives had large-bodied zooplankton year-round. In contrast, zooplankton communities in lakes with anadromous alewives cycled between large-bodied zooplankton in the winter and spring and small-bodied zooplankton in the summer. In summer, differences in feeding morphology of alewives caused zooplankton biomass to be lower and body size to be smaller in lakes with anadromous alewives than in lakes with landlocked alewives. Furthermore, intraspecific variation altered the strength of the trophic cascade caused by alewives. Our results demonstrate that intraspecific phenotypic variation of predators can regulate community structure and ecosystem processes by modifying the form and strength of complex trophic interactions.     

Extinction, recolonization, and dispersal through time in a planktonic crustacean

Dormant propagule banks are important reservoirs of biological and genetic diversity of local communities and populations and provide buffering mechanisms against extinction. Although dormant stages of various plant and animal species are known to remain viable for decades and even centuries, little is known about the effective influence of recolonization from such old sources on the genetic continuity of intermittent populations under natural conditions. Using recent and old dormant eggs recovered from a dated lake sediment core in Kenya, we traced the genetic composition of a local population of the planktonic crustacean Daphnia barbata through a sequence of extinction and recolonization events. This was combined with a phylogeographic and population-genetic survey of regional populations. Four successive populations, fully separated in time, inhabited Lake Naivasha from ca. 1330 to 1570 AD, from ca. 1610 to 1720 AD, from ca. 1840 to 1940 AD, and from 1995 to the present (2001 AD). Our results strongly indicate genetic continuity between the 1840-1940 and 1995-2001 populations, which are separated in time by at least 50 years, and close genetic relatedness of them both to the 1330-1580 population. A software tool (Colonize) was developed to find the most likely source population of the refounded 1995-2001 population and to test the number of colonists involved in the recolonization event. The results confirmed that the 1995-2001 population most probably developed out of a limited number of surviving local dormant eggs from the previous population, rather than out of individuals from regional (central and southern Kenya) or more distant (Ethiopia, Zimbabwe) populations that may have immigrated to Lake Naivasha through passive dispersal. These results emphasize the importance of prolonged dormancy for the natural long-term dynamics of crustacean zooplankton in fluctuating environments and suggest an important role of old local dormant egg banks in aquatic habitat restoration.     

Lake of flies, or lake of fish? A trophic model of Lake Malawi

Ecosystem-focused models have, for the first time, become available for the combined demersal and pelagic components of a large tropical lake ecosystem, Lake Malawi. These provide the opportunity to explore continuing controversies over the production efficiencies and ecological functioning of large tropical lakes. In Lake Malawi these models can provide important insight to the effect of fishing on fish composition, and the potential competition that the lakefly Chaoborus edulis may have with fisheries production. A mass-balanced trophic model developed for the demersal fish community of the southern and western areas of Lake Malawi was integrated with an existing trophic model developed for the open-water pelagic. Input parameters for the demersal model were obtained from a survey of fish distributions, fish food consumption studies, and from additional published quantitative and qualitative information on the various biotic components of the community. The model was constructed using the Ecopath approach and software. The graphically presented demersal food web spanned four trophic levels and was based primarily on consumption of detritus, zooplankton and sedimented diatoms. Zooplankton was imported into the system at trophic levels three and four through fish predation on carnivorous and herbivorous copepods and Chaoborus larvae. It is proposed that the primary consumption of copepods was by fish migrating into the pelagic zone. Chaoborus larvae in the demersal were probably consumed near the lakebed as they conducted a daily migration from the pelagic to seek refuge in the sediments. This evidence for strong benthic-pelagic coupling provided the opportunity for linking the demersal model to the existing model for the pelagic community so producing the first model for the complete ecosystem. Energy fluxes through the resulting combined model demonstrated that the primary import of biomass to the demersal system was detritus of pelagic origin (72.1%) and pelagic zooplankton (10.6%). Only 15.8% of the biomass consumed within the demersal system was of demersal origin. Lakefly production is efficiently utilised by the lake fish community, and any attempt to improve fishery production through introduction of a non-native plantivorous fish species would have a negative impact on the stability and productivity of the lake ecosystem.     

Is catchment productivity a useful predictor of taxa richness in lake plankton communities?

The influence of catchment variables on lake organisms is understudied. The terrestrial zone in the vicinity of lakes is, however, probably highly important for biota due to the effects on water chemistry and to various processes operating across ecosystem boundaries. We examined the relative importance of lake and catchment variables, as well as large-scale geographical factors, on the taxa richness of phyto- and zooplankton in 100 small lakes in Finland. In variation partitioning, the variability of phytoplankton richness was most strongly related to the effects of lake variables, the joint effects of lake and catchment variables, and the joint effects of all three groups of variables. Zooplankton richness, in turn, was most strongly related to the effects of lake and catchment variables and the joint effect of lake and catchment variables. The exact results of the variation partitioning depended on the catchment sizes considered in the regression models. Among lake variables, planktonic richness was strongly related to variables indicating productivity. Among catchment variables, the normalized difference vegetation index (NDVI), indicating catchment productivity, showed a relatively strong association with planktonic richness. These results provide evidence that catchment variables such as the NDVI may be efficient predictors of planktonic richness in small lakes. It is possible that individual lakes embedded in a highly productive landscape have higher taxa richness than solitary, potentially productive lakes because of the high influx of dispersing propagules from the regional pool. We also suggest that catchment variables may respond to environmental changes at different scales than the lake variables, and explicit consideration of catchment productivity would therefore be useful when planning research and monitoring programs for freshwater organisms.     

Optimization of exergy and implications of body sizes of phytoplankton and zooplankton in an aquatic ecosystem model

Size appears to be an important parameter in ecological processes. All physiological processes vary with body size ranging from small microorganisms to higher mammals. In this model, five state variables — phosphorus, detritus, phytoplankton, zooplankton and fish are considered. We study the implications of body sizes of phytoplankton and zooplankton for total system dynamics by optimizing exergy as a goal function for system performance indicator. The rates of different sub-processes of phytoplankton and zooplankton are calculated, by means of allometric relationships of their body sizes. We run the model with different combinations of body sizes of phytoplankton and zooplankton and observe the overall biomass of phytoplankton, zooplankton and fish. The highest exergy values in different combinations of phytoplankton and zooplankton size indicate the maximum biomass of fish with relative proportions of phytoplankton and zooplankton. We also test the effect of phosphorus input conditions corresponding to oligotrophic, mesotrophic, eutrophic system on its dynamics. The average exergy to be maximized over phytoplankton and zooplankton size was computed when the system reached a steady state. Since this state is often a limit cycle, and the exergy copies this behaviour, we averaged the exergy computed for 365 days (duration of 1 year) in the stable period of the run. In mesotrophic condition, maximum fish biomass with relative proportional ratio of phytoplankton, zooplankton is recorded for phytoplankton size class 3.12 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume). In oligotrophic condition the highest average exergy is obtained in between phytoplankton size 1.48 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume), whereas in eutrophic condition the result shows the highest exergy in the combination of phytoplankton size 5.25 (log V μm³ volume) and zooplankton size 4 (log V μm³ volume).     

Benthic-planktonic coupling, regime shifts, and whole-lake primary production in shallow lakes

Alternative stable states in shallow lakes are typically characterized by submerged macrophyte (clear-water state) or phytoplankton (turbid state) dominance. However, a clear-water state may occur in eutrophic lakes even when macrophytes are absent. To test whether sediment algae could cause a regime shift in the absence of macrophytes, we developed a model of benthic (periphyton) and planktonic (phytoplankton) primary production using parameters derived from a shallow macrophyte-free lake that shifted from a turbid to a clear-water state following fish removal (biomanipulation). The model includes a negative feedback effect of periphyton on phosphorus (P) release from sediments. This in turn induces a positive feedback between phytoplankton production and P release. Scenarios incorporating a gradient of external P loading rates revealed that (1) periphyton and phytoplankton both contributed substantially to whole-lake production over a broad range of external P loading in a clear-water state; (2) during the clear-water state, the loss of benthic production was gradually replaced by phytoplankton production, leaving whole-lake production largely unchanged; (3) the responses of lakes to biomanipulation and increased external P loading were both dependent on lake morphometry; and (4) the capacity of periphyton to buffer the effects of increased external P loading and maintain a clear-water state was highly sensitive to relationships between light availability at the sediment surface and the of P release. Our model suggests a mechanism for the persistence of alternative states in shallow macrophyte-free lakes and demonstrates that regime shifts may trigger profound changes in ecosystem structure and function.     

The variations of exergies and structural exergies along eutrophication gradients in Chinese and Italian lakes

Taking 29 Chinese lakes and 29 Italian lakes as two separate case studies, the paper presented the variations of exergies and structural exergies along eutrophication gradients in Chinese and Italian lakes. The exergies (Ex) and structural exergies (Exst) were calculated based on phytoplankton biomass (BA) and zooplankton biomass (BZ). A trophic state index (TSI) scaling from 0 to 100 was developed to classify trophic status for Chinese and Italian lakes based on three indicators, chlorophyll-a concentration (Chl-a, in mg/m³), total phosphorus (TP in mg/m³) and transparency in Secchi disk depth (SD in m). The relationships between TSI and Ex, Exst, total biomass, BZ/BA ratios were analyzed. The following results were obtained: (1) with the increase of TSI in Chinese and Italian lakes, there is an increasing trend for Ex, and a decreasing trend for Exst, generally. The obvious negative correlations exist between TSI and Exst, at the significant level of 0.01 for Italian lakes, and 0.05 for Chinese lakes. The obvious positive correlations exist between TSI and Ex, at the significant level of 0.01 for Chinese lakes, and for Italian lakes in Spring, Autumn and the all-year. (2) The structural exergy is more dependent on the ratio of phytoplankton biomass to zooplankton biomass (BZ/BA) than the exergy, and the exergy is more dependent on total biomass than the structural exergy. (3) The phytoplankton biomass (BA) and zooplankton biomass (BZ) are increased with the increasing TSI in Chinese and Italian lakes, and phytoplankton biomass (BA) increases more rapidly then zooplankton biomass (BZ) does. This results in the definite decrease of BZ/BA ratio with the increasing trophic status index. Such changes of BA, BZ and BZ/BA ratio could explain successfully the variations of exergies and structural exergies along eutrophication gradients in Chinese and Italian lakes. From the two separate case studies of Chinese lakes and Italian lakes, it could be concluded that exergy and structural exergy are feasible to serve as the system-level ecological indicators to give appropriate information on the trophic status of different lakes.