Connectivity and complexity of floodplain habitats govern zooplankton dynamics in a large temperate river system

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Feeding Ecology and Energetic Relationships with Habitat of Blue Catfish, Ictalurus furcatus, and Flathead Catfish, Pylodictis olivaris, in the Lower Mississippi River, U.S.A.

We examined feeding of blue catfish, Ictalurus furcatus, and flathead catfish, Pylodictis olivaris, collected from floodplain lake, secondary (side) river channel, and main river channel habitats in the lower Mississippi River (LMR), U.S.A. We described the feeding ecology of two large river catfish species within the context of whether off-channel habitats in the LMR (i.e., floodplain lakes and secondary channels) potentially provided energetic benefits to these fishes as purported in contemporary theory on the ecology of large rivers. We used diet composition and associated caloric densities of prey consumed as indicators of energetic benefit to catfishes. Differences in diet among habitats were strong for blue catfish, but weak for flathead catfish; consumed foods generally differed among habitats in caloric (energy) content. Caloric densities of consumed foods were generally greatest in floodplain lakes, least in the main river channel, and intermediate in secondary river channels. Strong between-year variation in diet was observed, but only for blue catfish. Blue catfish fed disproportionately on lower-energy zebra mussels in the main river channel during 1997, and higher-energy chironomids and oligochaetes in floodplain lakes during 1998. Results suggested that although off-channel habitats potentially provided greater energetic return to catfishes in terms of foods consumed, patterns of feeding and subsequent energy intake may vary annually. Energetic benefits associated with off-channel habitats as purported under contemporary theory (e.g., the ‘flood-pulse concept’) may not be accrued by catfishes every year in the LMR.     

Phytoplankton diversity (alpha,beta, and gamma) from the Araguaia River tropical floodplain lakes (central Brazil)

Alpha, beta and gamma are three components of species diversity. Knowing these attributes in floodplain lake phytoplankton communities is vital when selecting conservation areas. Species diversity is commonly used with other taxonomic groups, but rarely with phytoplankton. We compared the number of phytoplankton species (alpha diversity) from 21 Middle Araguaia River floodplain lakes in the 2000 and 2001 rainy and dry seasons. From these samples we estimated complete survey species richness (gamma diversity), quantified differences in species composition between lakes (beta diversity) and assessed the influence of abiotic variables on beta diversity. We recorded a total of 577 taxa. The Sjack1 estimator indicated that 62.31% of taxa were sampled in the 2000 rainy and 67.65% dry seasons, and 68.36% in the 2001 rainy and 73.5% dry seasons. In almost all seasons, alpha diversity negatively correlated with latitude. Beta diversity (β-1) was higher in high water periods, especially in 2000. This may have been caused by isolated heavy rainfall, which would have increased environmental heterogeneity and raised beta diversity. DCA showed differences in phytoplankton composition between rainy and dry seasons in 2000 and 2001, reflecting the influence of flood pulses on phytoplankton composition. The Mantel test indicated spatial distribution patterns where geographically more distant lakes had less-similar phytoplankton communities. Handling editor: J. Padisak     

The influence of floodplain habitat on the quantity and quality of riverine phytoplankton carbon produced during the flood season in San Francisco Estuary

Primary productivity, community respiration, chlorophyll a concentration, phytoplankton species composition, and environmental factors were compared in the Yolo Bypass floodplain and adjacent Sacramento River in order to determine if passage of Sacramento River through floodplain habitat enhanced the quantity and quality of phytoplankton carbon available to the aquatic food web and how primary productivity and phytoplankton species composition in these habitats were affected by environmental conditions during the flood season. Greater net primary productivity of Sacramento River water in the floodplain than the main river channel was associated with more frequent autotrophy and a higher P:R ratio, chlorophyll a concentration, and phytoplankton growth efficiency (α^B). Total irradiance and water temperature in the euphotic zone were positively correlated with net primary productivity in winter and early spring but negatively correlated with net primary productivity in the late spring and early summer in the floodplain. In contrast, net primary productivity was correlated with chlorophyll a concentration and streamflow in the Sacramento River. The flood pulse cycle was important for floodplain production because it facilitated the accumulation of chlorophyll a and wide diameter diatom and green algal cells during the drain phase. High chlorophyll a concentration and diatom and green algal biomass enabled the floodplain to export 14–37% of the combined floodplain plus river load of total, diatom and green algal biomass and wide diameter cells to the estuary downstream, even though it had only 3% of the river streamflow. The study suggested the quantity and quality of riverine phytoplankton biomass available to the aquatic food web could be enhanced by passing river water through a floodplain during the flood season.     

Relative importance of periphyton and phytoplankton in turbid and clear vegetated shallow lakes from the Pampa Plain (Argentina): a comparative experimental study

We analyzed experimentally the relative contribution of phytoplankton and periphyton in two shallow lakes from the Pampa Plain (Argentina) that represent opposite scenarios according to the alternative states hypothesis for shallow lakes: a clear lake with submerged macrophytes, and a turbid lake with high phytoplankton biomass. To study the temporal changes of both microalgal communities under such contrasting conditions, we placed enclosures in the littoral zone of each lake, including natural phytoplankton and artificial substrata, half previously colonized by periphyton until a mature stage and half clean to analyze periphyton colonization. In the clear vegetated shallow lake, periphyton chlorophyll a concentrations were 3–6 times higher than those of the phytoplankton community. In contrast, phytoplankton chlorophyll a concentrations were 76–1,325 times higher than those of periphyton in the turbid lake. Here, under light limitation conditions, the colonization of the periphyton was significantly lower than in the clear lake. Our results indicate that in turbid shallow lakes, the light limitation caused by phytoplankton determines a low periphyton biomass dominated by heterotrophic components. In clear vegetated shallow lakes, where nitrogen limitation probably occurs, periphyton may develop higher biomass, most likely due to their higher efficiency in nutrient recycling.     

Variation in phytoplankton composition and limnological features in a water–water ecotone of the Lower Paraná Basin (Argentina)

1. Structure and dynamics of phytoplankton were studied at six sites in a reach of the Lower Paraná River floodplain, from the main course to a marginal shallow lake, connected through a stream. This system is defined as a water–water ecotone and thus, the aim of this work is to analyse the spatial changes of phytoplankton along the transitional system.
2. Temporal fluctuations in number of species and phytoplankton density were more pronounced in the river than in the shallow lake. Even though most species were ubiquitous, there was a clear segregation of the most abundant populations across the ecotone. Aulacoseira granulata var. granulata and other centric diatoms decreased towards the lentic system.
3. Transparency varied according to river discharge, being higher in the lake during high water phases, while the opposite behaviour was observed during dry periods. These fluctuations were related to precipitation and resuspension processes.
4. Nitrate concentration diminished towards the lake while soluble reactive phosphorus remained fairly homogeneous across the ecotone.     

Seasonal succession of phytoplankton in a lake of the Paraná river floodplain,Argentina

Phytoplankton biomass, morphological and taxonomic composition, species diversity and productivity were analyzed in a shallow lake of the Middle Paraná River floodplain (El Tigre, 31 ° 41 S and 60° 42 W), between November 1986 and July 1988. Lake inundation (filling and through-flow phases) constituted an intense long-term perturbation in the physical and chemical environment. As the lake filled with river water, K-selected species (netplanktonic filamentous bluegreens, > 37 µm, with low surface area/volume (SA/V) ratios) that had existed prior to filling (late spring 1986) were replaced in summer-fall by r-selected species (nannoplanktonic chlorophytes and cryptophytes, < 37 µm, mainly stout forms with high SA/V ratios). During the through-flow phase, lentic phytoplankton was replaced by lotic flagellate populations due to the direct flushing by river water. During the period of falling water (drainage and isolation phases), nanoplanktonic algae with similar characteristics to those of the filling phase dominated in late winter-spring. Later in the isolation phase, these were succeeded by K-selected species (netplanktonic algae, mainly motile spherical dinoflagellates and filamentous bluegreens with low SA/V ratios). Simultaneously, primary production per unit biomass decreased and total biomass and specific diversity increased. Seasonal changes of phytoplankton in floodplain lakes can be interpreted as the interaction between true successional development (as observed in the drainage and isolation phases) and intermediate disturbance. Using Reynolds' terminology, short-term disturbance (slight inflow of nutrient-rich river water) caused reversion to an earlier stage in the former succession, and long-term disturbance (lake inundation) truncated the successional progression and a new (or shifted) succession was initiated.     

Water quality dynamics of oxbow lakes in young glacial landscape of NE Poland in relation to their hydrological connectivity

In this paper, the functioning of floodplain lakes was documented on the basis of water quality and hydrological data in a postglacial river floodplain representative of the temperate climatic zone. Nine oxbow lakes in the middle section of the River ?yna floodplain located in north-eastern Poland were investigated. Three groups of oxbow lakes distinguished on the basis of hydrological connectivity and retention time (lotic, semi-lotic and lentic) showed significant spatial and temporal differences in chemical parameters of water and matter cycling in relation to water table fluctuations, sediment character and aquatic plant diversity. Lotic habitats had parameters similar to river water, in particular to EC, main cation and anion concentrations. When the connection to a river is maintained, oxbow lakes prolong their existence due to multiple exchanges of matter. Very short-term water retention in lotic habitats is not favorable for sedimentation of such particles as Ca or SO₄, Na, K, Cl which are easily moved by river water. This is contrary to lentic (disconnected) water bodies, where retention time is incomparably longer and internal cycling limits ecosystem productivity. Disconnected oxbow lakes undergo the highest seasonal variations in nutrient availability related in summer to intensive development of free-floating phytocenoses that limit light and aeration conditions, whereas and in winter to long-term ice cover. The presented data indicate an important function of oxbow lakes that relies on the regulation of nutrient transfer towards rivers. The effectiveness of TIN withdrawal from lentic oxbow lakes is estimated as 75%, whereas and in other macroelements from 50% to 82%, when compared to the lotic type. The retention ability of natural riverine ecosystems should be considered in proper designs of created wetlands.     

Dynamics of phytoplankton pigment characteristics in the Biya head water

Concentrations of phytoplankton photosynthetic pigments were measured at the Biya head water (in the immediate vicinity of Lake Teletskoe) every 10 days from July 1998 through December 2001. Comparison of phytoplankton pigment characteristics at the river’s head waters and at the surface of the lake’s pelagic zone for the ice-free period shows that the monitoring data give a clear picture of the formation and functioning of the lake phytoplankton. Analysis of seasonal fluctuations of pigment characteristics and their proportions revealed characteristic stages in the development of lake algal cenoses. We have demonstrated similarities and differences in the seasonal dynamics of the phytoplankton between Lake Teletskoe, a deep oligotrophic lake in the south of West Siberia, as compared with the general pattern observed in temperate lakes. According to the concentration of chlorophyll a, the trophic status of the lake’s pelagic zone and the Biya head water is ultraoligotrophic-oligotrophic. Yellow:green pigment ratio ranks Lake Teletskoe among “carotenoid lakes“. Low coefficients of linear correlation between chlorophyll a, water level and biogenic matter concentration shows that the relationships analyzed are far more complex than has been inferred from the combined effect of abiotic and biotic factors.     

Composition changes of phototrophic microbial communities along the salinity gradient in the solar saltern evaporation ponds of Eilat,Israel

There are several conflicting hypothesis that deal with the influence of flooding in the natural river–floodplain systems. According to the Flood Pulse Concept, the flood pulses are not considered to be a disturbance, while some recent studies have proven that floods can be a disturbance factor of phytoplankton development. In order to test whether flooding acts as a disturbance factor in the shallow Danubian floodplain lake (Lake Sakadaš), phytoplankton dynamics was investigated during two different hydrological years—extremely dry (2003) without flooding and usually flooded (2004). A total of 18 phytoplankton functional groups were established. The sequence of phytoplankton seasonality can be summarized P/D → E (W1, W2) → C/P (only in potamophase) → S2/H1/S_N/S1 → W1/W2 → P/D. The canonical correspondence analysis (CCA) demonstrated that the water level was a significant environmental variable in 2004. Due to the higher total biomass of Bacillariophyceae established under potamophase conditions, floodings in the early spring seem to be a stimulating factor for phytoplankton development. On the other hand, the flood pulses in May and June had dilution effects on nutrients, so that a significantly lower phytoplankton biomass was established indicating that flooding pulses can be regarded as a disturbance event. Such conditions supported diatom development (D, P, C species) and prolonged its dominance in the total phytoplankton biomass. A long-lasting Cyanoprokaryota bloom (various filamentous species—S1, S2, S_N and H1 representatives) with very high biomass characterized the limnophase (dry conditions) in summer and autumn of both years. In-lake variables (lake morphology, internal loadings of nutrients from sediments, light conditions) seem to be important for the appearance of Cyanoprokaryota bloom. The equilibrium phase was found during the Cyanoprokaryota bloom only in the extremely dry year. This study showed that depending on the time scale occurrence, flood pulses can be a stimulating or a disturbance factor for phytoplankton development in Lake Sakadaš. Handling editor: J. Padisak     

Composition,size, and biomass of zooplankton in large productive Florida lakes

Crustacean zooplankton data were compiled from long-term observational studies at seven large shallow Florida lakes, to determine whether there are general characteristics in regard to species composition, body size, and biomass. In particular, we examined whether patterns in body size and species richness fit empirical models developed by Stanley Dodson. The lakes included range in size from 125 to 1730 km² and encompass mesotrophic to hyper-eutrophic conditions. We found that zooplankton biomass was strongly dominated by one species of calanoid copepod—Arctodiaptomus dorsalis. Large daphnids were absent, and Cladocera assemblages were dominated by small taxa such as Ceriodaphnia, Chydorus, and Eubosmina. The total number of species of pelagic cladocerans (8–12) was consistent with Dodson’s predictions based on lake area. The average size of crustacean zooplankton in Florida lakes is small in comparison with temperate communities. A. dorsalis is the smallest calanoid copepod in North America, and the mean length of Cladocera (0.6 mm) is consistent with Dodson’s results that size decreases from temperate to tropical zones. Total biomass of crustacean zooplankton was very low, ratios of zooplankton to phytoplankton biomass (0.01–0.1) are among the lowest reported in the literature, and the zooplankton displayed short-lasting early spring peaks in biomass. Cladocera were almost entirely absent in spring and summer. Factors known to occur in Florida lakes, which appear to explain these characteristics of biomass, include intense fish predation and high summer water temperature.     

Spatial and Temporal Trade-Offs by Bluegills in Floodplain River Ecosystems

Ecological trade-offs by organisms to minimize mortality and maximize growth is a foundational theme in ecology. Yet, these trade-offs are rarely examined within spatially complex, temporally variable ecosystems, such as floodplain rivers. Here, we evaluate ecological trade-offs across space and time for the bluegill (Lepomis macrochirus) in two unregulated river ecosystems in southeastern USA. Life-history differences among spatially segregated main channel and floodplain lake populations were used to assess effects of habitat type on bluegill fitness. Growth, condition, and gonadal somatic index were all significantly enhanced in floodplain lakes relative to the main channel. Furthermore, stomach fullness was significantly higher, and predator densities significantly lower in floodplain lakes thereby providing an ecological explanation for the life-history plasticity observed across the riverscape. However, historical observations suggested that although floodplain lakes are highly productive for bluegills, they are also prone to complete desiccation by drought approximately every 5 years, revealing the ultimate value of channel habitat, which does not dry, as desiccation refugia. Bluegills are faced with a balancing act associated with variation in foraging opportunities, and risks to predation and desiccation, that change in both the temporal and the spatial dimensions of floodplain rivers. The differential responses to these opportunities and risks help to explain why both habitats remain actively populated by bluegills, as well as many other organisms, in these and many other natural rivers.     

The distribution of macroinvertebrate assemblages in a reach of the River Allier (France), in relation to riverbed characteristics

Investigations on large canalised rivers, for example the Danube, have shown that transported particulate matter, which is typically inorganic, is predominantly deposited in waters near the river’s main channel. This investigation deals with the lower section of the River Havel (NE Germany), a canalised lowland river with a very flat floodplain. This river is highly polluted by nutrients from urban areas (Berlin) and a long chain of river lakes produces high concentrations of phytoplankton. Due to the high proportion of planktogenic detritus, it was hypothesised that greater quantities of nutrient-rich fine particulate organic matter (FPOM) would be deposited in floodplain waters located further from the main channel than has been reported for large rivers. The total nutrient, P-binding metal (Fe, Al, and Mn), organic and inorganic carbon (TOC, TIC) contents of the upper organic sediment layer (0–4 cm) were analysed in samples collected from 48 floodplain water and river sites. The sediment bulk density, calculated on the basis of dry mass content and loss on ignition, was used to characterise the waters according to the impact of the river current. The results showed that the variability of total phosphorus (TP) was best explained by the variability of total iron (TFe, R² = 0.52). The floodplain water sediments could clearly be separated into two groups on the basis of the sediment particle size composition, and of the element ratios TOC:TP, TN:TP, primarily TFe:TP. The sediments from impounded river sections and from mouth sections of backwaters (approx. 100–200 m) were characterised by a high proportion particles from the 0.1 to 0.5 mm size fraction and by homogeneous, low TFe:TP, TOC:TP and TN:TP ratios. Sediments from distal sections of backwaters and of oxbow lakes tended to exhibit high element ratios with much higher variability. These results were interpreted as a spatially limited impact of the river on the floodplain water sediments. Contrary to expectation, the phosphorus bound in river seston was predominantly and very homogeneously deposited in the impounded river and mouth sections of backwaters. This implies that the inundation of the floodplain waters during spring floods seems to have no important material impact on the sediments in waters of low hydrological connectivity with the River Havel.     

Cyanobacterial blooms in a temperate river-floodplain ecosystem: the importance of hydrological extremes

In the past decade, extreme hydrological events were expressed with extreme droughts and floods in temperate regions. The aim of this paper is to explain how such changes in hydrology can influence cyanobacterial populations in floodplain ecosystems. We therefore analyzed a 6-year (2003–2008) study of the phytoplankton in the Kopački Rit floodplain, one of the largest natural floodplains in the middle section of the Danube River (Europe). During the studied period, the shallow floodplain lake shifted between a state of turbid water, characterized by high phytoplankton biomass and regular appearance of cyanobacteria blooms, to a state of clear water with very low phytoplankton biomass and absence of cyanobacteria, and back to the turbid state. Apparently, the major forces driving the cyclic shift were closely related to extremely high and long-lasting flood events. Significant increase in water level, low hydraulic residence time of water, decrease in transparency and low-light climate, together with mass developed aquatic macrophyte vegetation in the whole inundated floodplain were unfavorable conditions for growth and proliferation of cyanobacteria. With the establishment of the flood regime characterized by long-lasting periods without flooding, in-lake processes prevailed leading to cyanobacterial bloom. The most successful were filamentous non-N-fixing cyanobacteria tolerant to mixed and low-light conditions (Planktothrix and Limnothrix) and invasive species Cylindrospermopsis raciborskii. Their massive development led to the establishment of a phytoplankton steady state. All our results demonstrate that the altered intensity and frequency of flood events will have pronounced effects on the appearance of cyanobacterial blooms and generally on alternative stable states in the floodplain. Relating to this, management objectives should be focused on qualifications of changes in hydrology and projecting those effects for potential floodplain restoration.     

Influence of hydrological seasonality on bacterioplankton in two neotropical floodplain lakes

Seasonal fluctuation in river stage strongly affects the ecological functioning of tropical floodplain lakes. This study was conducted to assess the influence of hydrological seasonality on bacterial production and abundance in two floodplain lakes of the Autana River, a blackwater river in the Middle Orinoco basin, Venezuela. Water samples for nutrient chemistry, chlorophyll a, and microbiological determinations were collected in two floodplain lakes and in the mainstem of the river during 1997–98. DOC and chlorophyll a concentrations were similar between mainstem and lake sites during high water when river and lakes were well connected but became different during the period of low water when the interaction was minimal. Higher values of bacterial production were observed in the floodplain lakes (0.62–1.03 g C l^–1 h^–1) compared to the mainstem sites (0.17–0.19 g C l^–1 h^–1) during the period of low water, while during the period of high water river and lake sites showed similar levels (0.04 g C l^–1 h^–1). Bacterial numbers followed bacterial production in the floodplain lakes, reaching higher numbers during the period of low water (1.41–2.40 × 10⁶ cells ml^–1). Availability of substrate and inorganic nutrients, pH, and inputs and losses of bacterial cells could be determining the observed seasonal patterns in bacterial production and abundance. The Autana lakes exhibited a strong seasonal pattern in the chemical and biological conditions, showing higher productivity during the lentic phase that lasted between 5 and 6 months.     

Performance trade-off across a natural resource gradient

An important environmental factor determining both phytoplankton and zooplankton community composition is lake depth and thermal stratification. However, there is little information on how the interaction between zooplankton grazers and their phytoplankton food changes along an environmental gradient of lake depth. We contrasted resource availability for daphniid zooplankton populations living in two shallow, unstratified lakes and in two deep, stratified lakes using a novel growth bioassay. Stratified lakes had consistently lower resource richness than shallow unstratified lakes. To test whether resources were important in explaining differences in daphniid composition of shallow and deep lakes, we performed reciprocal transplant experiments. We raised daphniids typical of shallow (Ceriodaphnia reticulata) and deep (Daphnia dentifera) lakes in the resources from replicate shallow and deep lakes and monitored survival and reproduction. The two species exhibited a performance trade-off, measured by life table r and R ₀, across a gradient in natural resource richness. D. dentifera had higher relative fitness than C. reticulata when raised in the poorest resource environment from a deep lake. However, under richer resource conditions typical of shallow lakes, C. reticulata outperformed D. dentifera. We further created a gradient in natural resource quantity (by dilution) to test whether this trade-off in species relative fitness involved aspects of resource quality. No trade-off in species performance was evident across the dilution gradient, indicating that resource quality was important to the trade-off. We conclude that shifts in daphniid species composition along a gradient of lake depth involve an adaptive trade-off in ability to exploit rich versus poor resource quality. Received: 11 May 1998 / Accepted: 15 January 1999     

Comparative analysis of nutrients,chlorophyll and transparency in two large shallow lakes (Lake Taihu,P.R. China and Lake Okeechobee,USA)

This article compares limnological attributes of two of the world’s largest shallow lakes—Lake Okeechobee in Florida, USA and Lake Taihu in P.R. China. Both the systems support an array of ecological and societal values including fish and wildlife habitat, public water supply, flood protection, and recreation. Both have extensive research programs, largely because of concern regarding the lakes’ frequent cyanobacterial blooms. By evaluating these systems together, we compare and contrast properties that can generally advance the understanding and management of large shallow lowland lakes. Because of shallow depth, long fetch, and unconsolidated mud sediments, water chemistry, and transparency in both the lakes are strongly influenced by resuspended sediments that affect light and nutrient conditions. In the central region of both the lakes, where depth is the greatest, evaluation of limiting factors by a trophic state index approach indicates that light most often limits phytoplankton biomass. In contrast, the more sheltered shoreline areas of both the lakes display evidence of nitrogen (N) limitation, which also has been confirmed in nutrient assays conducted in earlier studies. This N limitation most likely is a result of excessive levels of phosphorus (P) that have developed in the lakes due to high external loads over recent decades and the currently high internal P recycling. Comparisons of these lakes show that Lake Taihu has higher N than, similar total phosphorus (TP) and similar light conditions to that of Lake Okeechobee, but less chlorophyll a (CHL). The latter may be as a result of lower winter temperatures in Lake Taihu (around 5°C) compared to Lake Okeechobee (around 15°C), which could reduce phytoplankton growth and abundance through the other seasons of the year. In these systems, the important role of light, temperature, and nutrients in algal bloom dynamics must be considered, especially due to possible adverse and unintended effects that might occur with projects such as sediment removal, and in the long term, in regard to buffering lake responses to external load reduction. Handling editor: D. Hamilton     

Scaling of Pelagic Metabolism to Size,Trophy and Forest Cover in Small Danish Lakes

ABSTRACT We tested whether pelagic light and nutrient availability, metabolism, organic pools and CO₂-supersaturation were related to lake size and surrounding forest cover in late summer–autumn measurements among 64 small (0.02–20 ha), shallow seepage lakes located in nutrient-rich, calcareous moraine soils in North Zealand, Denmark. We found a strong implicit scaling to lake size as light availability increased significantly with lake size while nutrient availability, phytoplankton biomass and dissolved organic matter declined. Forest lakes had significantly stronger net heterotrophic traits than open lakes as higher values were observed for light attenuation above and in the water, dissolved organic matter, pelagic community respiration (R) relative to maximum gross primary production (R/GPP) and CO₂-supersaturation. Total-phosphorus was the main predictor of phytoplankton biomass (Chl) despite a much weaker relationship than observed in previous studies of larger lakes. Maximum gross primary production increased with algal biomass and decreased with dissolved organic matter, whereas community respiration increased with dissolved organic matter and particularly with gross primary production. These results suggest that exogenous organic matter supplements primary production as an energy source to heterotrophs in these small lakes, and particularly so in forest lakes experiencing substantial shading from the forest and dissolved humic material. This suggestion is supported by 20–30-fold CO₂ supersaturation in the surface water of the smallest forest lakes and more than sixfold supersaturation in 75% of all measurements making these lakes among the most supersaturated temperate lakes examined so far.