The ecological organisation of lake districts: general introduction

1. Here we introduce a special issue of Freshwater Biology that focuses on the landscape‐scale spatial and temporal patterns exhibited by multiple lakes within lake districts. We call this patterning, and the processes that lead to it, the ‘ecological organisation of lake districts.’ 2. Papers in this special issue share the common goal of examining landscape‐level processes that lead to spatial and temporal patterns of lake characteristics in individual or multiple lake districts. Several papers focus on the degree to which multiple lakes have synchronous among year variability in various physical, chemical, and biological variables. Others focus on the landscape‐level processes that lead to spatial patterning of lake characteristics. Finally, a few papers examine the relationship between spatial patterning and temporal dynamics. Papers in this special issue present results from 10 lake districts from North America, Europe, and Antarctica.     

Structuring features of lake districts: landscape controls on lake chemical responses to drought

1. Within a lake district of relatively homogeneous geomorphology, the responses of lakes to climate are influenced by the complexity of the hydrogeologic setting, position in the landscape, and lake‐specific biological and physical features. We examined lake chemical responses to drought in surface water‐ and groundwater‐dominated districts to address two general questions. (1) Are spatial patterns in chemical dynamics among lakes uniform and synchronous within a lake district, suggesting broad geomorphic controls; variable in a spatially explicit pattern, with synchrony related to landscape position, suggesting hydrologic flowpath controls; or spatially unstructured and asynchronous, suggesting overriding control by lake‐specific factors? (2) Are lake responses to drought a simple function of precipitation quantity or are they dictated by more complex interactions among climate, unique lake features, and hydrologic setting? 2. Annual open‐water means for epilimnetic concentrations of chloride, calcium, sulfate, ANC, DOC, total nitrogen, silica, total phosphorus, and chlorophyll a measured between 1982 and 1995 were assembled for lakes in the Red Lake and ELA districts of north‐western Ontario, the Muskoka – Dorset district in south‐central Ontario, and the Northern Highland district of Wisconsin. Within each district, we compared responses of lakes classified by landscape position into highland or lowland, depending on relative location within the local to regional hydrologic flow system. Synchrony, defined as a measure of the similarity in inter‐annual dynamics among lakes within a district, was quantified as the Pearson product‐moment correlation (r) between two lakes with observations paired by year. To determine if solute concentrations were directly related to interannual variations in precipitation quantity, we used regression analysis to fit district‐wide slopes describing the relationship between each chemical variable and annual (June to May) and October to May (Oct–May) precipitation. 3. Among lakes in each of the three Ontario districts, the pattern of chemical response to interannual shifts in precipitation was spatially uniform. In these surface water‐ dominated districts, solute concentrations were generally a simple function of precipitation. Conservative solutes, like calcium and chloride, tended to be more synchronous and were negatively related to precipitation. Solutes such as silica, total phosphorus, and chlorophyll a, which are influenced by in‐lake processes, were less synchronous and relationships with precipitation tended to be positive or absent. 4. In the groundwater‐dominated Northern Highland lakes of Wisconsin, we observed spatial structure in drought response, with lowland lakes more synchronous than highland lakes. However, there was no evidence for a direct relationship between any solute and precipitation. Instead, increases in the concentration of the conservative ion calcium during drought were not followed by a symmetrical return to pre‐drought conditions when precipitation returned to normal or above‐average values. 5. For calcium, time lags in recovery from drought appeared related to hydrologic features in a complex way. In the highland Crystal Lake, calcium concentrations tracked lake stage inversely, with a return to pre‐drought concentrations and lake stage five years after the drought. This pattern suggests strong evaporative controls. In contrast, after five years of normal precipitation, calcium in the lowland Sparkling Lake had not returned to pre‐drought conditions despite a rebound in lake stage. This result suggests that calcium concentrations in lowland lakes were controlled more by regional groundwater flowpaths, which track climatic signals more slowly. 6. Temporal dynamics driven by climate were most similar among lakes in districts that have a relatively simple hydrology, such as ELA. Where hydrologic setting was more complex, as in the groundwater‐dominated Northern Highland of Wisconsin, the expression of climate signals in lakes showed lags and spatial patterns related to landscape position. In general, we expect that landscape and lake‐specific factors become increasingly important in lake districts with more heterogeneous hydrogeology, topography or land use. These strong chemical responses to climate need to be considered when interpreting the responses of lakes to other regional disturbances.     

Regional coherence of climatic and lake thermal variables of four lake districts in the Upper Great Lakes Region of North America

1. Within a region with common climatic conditions, lake thermal variables should exhibit coherent variability patterns to the extent to which they are not influenced by lake specific features such as morphometry and water clarity. We tested the degree of temporal coherence in interannual variability for climatic variables (air temperature and solar radiation) among four lake districts in the Upper Great Lakes Region. We also tested the degree of coherence of lake thermal variables (near‐surface temperature, eplimnetic temperature, hypolimnetic temperature and thermocline depth) for lakes within these districts. 2. Our four lake districts included the Experimental Lakes Area in north‐western Ontario, the Dorset Research Centre area north of Toronto, Ontario, the Northern Highland Lake District in northern Wisconsin, and the Yahara Lakes near Madison in southern Wisconsin. Seventeen lakes were analyzed for lake thermal variables dependent on stratification. Another five lakes were added for the analysis of near‐surface temperature. 3. The analysis tested whether for monthly and summer means, the climate (air temperature and solar radiation) across the four lake districts was coherent interannually and whether variables which measure the thermal structure of the lakes were coherent interannually among lakes within each lake district and across the four lake districts. 4. Temporal coherence was estimated by the correlation between lake districts for meteorological variables and between lake pairs for lake thermal variables. Mean coherence and the percentage of correlations exceeding the 5% significance level were derived both within and between lake districts for lake thermal variables. 5. Across the four lake districts, summer mean air temperature was highly coherent while summer solar radiation was less coherent. Approximately 60–80% of the interannual variation in mean summer air temperature at a site occurred across the entire region. Less than 45% of the variation in solar radiation occurred across sites. 6. Epilimnetic temperature and the near‐surface temperature were highly coherent both within and between lake districts. The coherence of thermocline depth within and between lake districts was weaker. Hypolimnetic temperature was not coherent between lake districts for most lake pairs. It was coherent among lakes within some lake districts. 7. The influences of local weather and differences among lakes in water clarity are discussed in the context of differences in levels of coherence among lake thermal variables and among lake pairs for a given variable.     

Integration of lakes and streams in a landscape perspective: the importance of material processing on spatial patterns and temporal coherence

1. We studied the spatial and temporal patterns of change in a suite of twenty‐one chemical and biological variables in a lake district in arctic Alaska, U.S.A. The study included fourteen stream sites and ten lake sites, nine of which were in a direct series of surface drainage. All twenty‐four sites were sampled between one and five times a year from 1991 to 1997. 2. Stream sites tended to have higher values of major anions and cations than the lake sites, while the lake sites had higher values of particulate carbon, nitrogen, phosphorous and chlorophyll a. There were consistent and statistically significant differences in concentrations of variables measured at the inlet versus the outlet of lakes, and in variables measured at upstream versus downstream sites in the stream reaches which connect the lakes. In‐lake processing tended to consume alkalinity, conductivity, H⁺, DIC, Ca²⁺, Mg²⁺, CO₂, CH₄, and NO₃^–, and produce K⁺ and dissolved organic carbon (DOC). In‐stream processing resulted in the opposite trends (e.g. consumption of K⁺ and DOC), and the magnitudes of change were often similar to those measured in the lakes but with the opposite sign. 3. Observed spatial patterns in the study lakes included mean concentrations of variables which increased, decreased or were constant along the lake chain from high to low altitude in the catchment (stream sites showed no spatial patterns with any variables). The strongest spatial patterns were of increasing conductivity, Ca²⁺, Mg²⁺, alkalinity, dissolved inorganic carbon and pH with lake chain number (high to low altitude in the basin). These patterns were partly determined by the effect of increasing catchment area feeding into lakes further downslope, and partly by the systematic processing of materials in lakes and in the stream segments between lakes. 4. Synchrony (the temporal coherence or correlation of response) of variables across all lakes ranged from 0.18 for particulate phosphorus to 0.90 for Mg²⁺ the average synchrony for all twenty‐one variables was 0.50. The synchronous behaviour of lake pairs was primarily related to the spatial location or proximity of the lakes for all variables taken together and for many individual variables, and secondarily, to the catchment to lake area ratio and the water residence time. 5. These results illustrate that, over small geographic areas, and somewhat independent of lake or stream morphometry, the consistent and directional (downslope) processing of materials helps produce spatial patterns which are coherent over time for many limnological variables. We combine concepts from stream, lake and landscape ecology, and develop a conceptual view of landscape mass balance. This view highlights that the integration of material processing in both lakes and rivers is critical for understanding the structure and function of surface waters, especially from a landscape perspective.     

A landscape approach to examining spatial patterns of limnological variables and long-term environmental change in a southern Canadian lake district

1. We explored patterns of limnological variables (physical, chemical and biological) with relation to landscape position (expressed as lake order) in 86 study lakes located on shield bedrock in south‐central Ontario, Canada. 2. Using anova s with lake order as the categorical variable, landscape position explained significant amounts of variation in major ion chemistry, physical and catchment characteristics, hypolimnetic oxygen, and community composition in algal (diatom, chrysophyte) and invertebrate (chironomid) assemblages preserved in surficial sediments. Several nutrient variables (TP, total phosphorus and TN, total nitrogen) and dissolved organic carbon did not have significant relationships with lake order. 3. The strongest relationships with lake order (as a fraction of variation explained in anova s) included silica concentrations (r² = 0.40) and SO₄ (r² = 0.29) concentrations, surface area (r² = 0.50) and hypolimnetic oxygen (r² = 0.29). 4. Bedrock geology (carbonate metasedimentary versus non‐carbonate bedrock) had strong influences on spatial gradients of pH and major ion chemistry. It was difficult to separate geological influences from spatial influences on limnological variables in this study, as drainage patterns in the region are highly influenced by surface features of underlying geological formations because of the very thin glacial till or exposed bedrock that exists in most catchments. 5. Patterns of limnological variables indicated that low‐order, headwater lakes had the lowest concentrations of major ions, and, from algal inferences of pH change, had been most susceptible to acidic deposition. High‐order, downstream lakes were larger and deeper, and had higher concentrations of hypolimnetic oxygen, indicating that optimal lake trout habitat was primarily located in high‐order lakes. 6. Variance partitioning analyses indicated that lake order as a metric of landscape position explained comparable portions of community variation in algal and invertebrate assemblages compared with geographic position (latitude, longitude) and Cartesian coordinate position (e.g. x, y, x², y², etc.) metrics. Lake order explained more community variation in chironomid assemblages compared with other landscape metrics, possibly because of the strong relationships between lake order and lake morphometry variables.     

Distribution of recreational boating across lakes: do landscape variables affect recreational use?

1. Because people impact lake ecosystems, it is important to consider factors influencing the human use of freshwater resources. We investigated the influence of the landscape position, as well as lake area, recreational facilities, and distance to highways and urban centres, on lake use by boaters in the Northern Highland Lake District of Wisconsin, U.S.A.
2. In aerial surveys of ninety-nine randomly selected lakes, we did not see boats on over half of the lakes. Of the lakes with boats, we found a strong correlation between the number of boats and lake area. Recreational boats tended to be found on large, accessible lakes with good boating facilities. Boats were not seen on small, stained lakes with few recreational facilities.
3. Regression models showed that lake size and landscape position explained 63% of the variability in the average number of boats per lake and landscape position explained 24% of the variability in boat density on all ninety-nine lakes. Social variables representing the quality of boating facilities and the perception of good fishing explained 70% of the variability in number of boats per lake and 54% of the variation in boat density on all lakes. A combined model using both physical and social variables increased the explanatory power for both number and density. Lake use by boaters was correlated with landscape position, the quality of fishing and the availability of recreational facilities. When the analysis was restricted to the forty-six lakes where boats were found, only the availability of recreational facilities proved a significant predictor of boat density.
4. Our results suggest that lake choice by recreational boaters may be best predicted by a combination of the location of a lake in a regional hydrologic landscape, and considerations of available facilities and perceptions regarding fishing.     

Climate and landscape influence on indicators of lake carbon cycling through spatial patterns in dissolved organic carbon

Freshwater ecosystems are strongly influenced by both climate and the surrounding landscape, yet the specific pathways connecting climatic and landscape drivers to the functioning of lake ecosystems are poorly understood. Here, we hypothesize that the links that exist between spatial patterns in climate and landscape properties and the spatial variation in lake carbon (C) cycling at regional scales are at least partly mediated by the movement of terrestrial dissolved organic carbon (DOC) in the aquatic component of the landscape. We assembled a set of indicators of lake C cycling (bacterial respiration and production, chlorophyll a, production to respiration ratio, and partial pressure of CO₂), DOC concentration and composition, and landscape and climate characteristics for 239 temperate and boreal lakes spanning large environmental and geographic gradients across seven regions. There were various degrees of spatial structure in climate and landscape features that were coherent with the regionally structured patterns observed in lake DOC and indicators of C cycling. These different regions aligned well, albeit nonlinearly along a mean annual temperature gradient; whereas there was a considerable statistical effect of climate and landscape properties on lake C cycling, the direct effect was small and the overall effect was almost entirely overlapping with that of DOC concentration and composition. Our results suggest that key climatic and landscape signals are conveyed to lakes in part via the movement of terrestrial DOC to lakes and that DOC acts both as a driver of lake C cycling and as a proxy for other external signals.     

Synchronization in ecological systems by weak dispersal coupling with time delay

One of the most salient spatiotemporal patterns in population ecology is the synchronization of fluctuating local populations across vast spatial extent. Synchronization of abundance has been widely observed across a range of spatial scales in relation to the rate of dispersal among discrete populations. However, the dependence of synchrony on patterns of among-patch movement across heterogeneous landscapes has been largely ignored. Here, we consider the duration of movement between two predator–prey communities connected by weak dispersal and its effect on population synchrony. More specifically, we introduce time-delayed dispersal to incorporate the finite transmission time between discrete populations across a continuous landscape. Reducing the system to a phase model using weakly connected network theory, it is found that the time delay is an important factor determining the nature and stability of phase-locked states. Our analysis predicts enhanced convergence to stable synchronous fluctuations in general and a decreased ability of systems to produce in-phase synchronization dynamics in the presence of delayed dispersal. These results introduce delayed dispersal as a tool for understanding the importance of dispersal time across a landscape matrix in affecting metacommunity dynamics. They further highlight the importance of landscape and dispersal patterns for predicting the onset of synchrony between weakly coupled populations.     

Landscape drivers of regional variation in the relationship between total phosphorus and chlorophyll in lakes

1. For north temperate lakes, the well‐studied empirical relationship between phosphorus (as measured by total phosphorus, TP), the most commonly limiting nutrient and algal biomass (as measured by chlorophyll a, CHL) has been found to vary across a wide range of landscape settings. Variation in the parameters of these TP–CHL regressions has been attributed to such lake variables as nitrogen/phosphorus ratios, organic carbon and alkalinity, all of which are strongly related to catchment characteristics (e.g. natural land cover and human land use). Although this suggests that landscape setting can help to explain much of the variation in ecoregional TP–CHL regression parameters, few studies have attempted to quantify relationships at an ecoregional spatial scale. 2. We tested the hypothesis that lake algal biomass and its predicted response to changes in phosphorus are related to both local‐scale features (e.g. lake and catchment) and ecoregional‐scale features, all of which affect the availability and transport of covarying solutes such as nitrogen, organic carbon and alkalinity. Specifically, we expected that land use and cover, acting at both local and ecoregional scales, would partially explain the spatial pattern in parameters of the TP–CHL regression. 3. We used a multilevel modelling framework and data from 2105 inland lakes spanning 35 ecoregions in six US states to test our hypothesis and identify specific local and ecoregional features that explain spatial heterogeneity in TP–CHL relationships. We include variables such as lake depth, natural land cover (for instance, wetland cover in the catchment of lakes and in the ecoregions) and human land use (for instance, agricultural land use in the catchment of lakes and in the ecoregions). 4. There was substantial heterogeneity in TP–CHL relationships across the 35 ecoregions. At the local scale, CHL was negatively and positively related to lake mean depth and percentage of wooded wetlands in the catchment, respectively. At the ecoregional scale, the slope parameter was positively related to the percentage of pasture in an ecoregion, indicating that CHL tends to respond more rapidly to changes in TP where there are high levels of agricultural pasture than where there is little. The intercept (i.e. the ecoregion‐average CHL) was negatively related to the percentage of wooded wetlands in the ecoregion. 5. By explicitly accounting for the hierarchical nature of lake–landscape interactions, we quantified the effects of landscape characteristics on the response of CHL to TP at two spatial scales. We provide new insight into ecoregional drivers of the rate at which algal biomass responds to changes in nutrient concentrations. Our results also indicate that the direction and magnitude of the effects of certain land use and cover characteristics on lake nutrient dynamics may be scale dependent and thus likely to represent different underlying mechanisms regulating lake productivity.     

Remote European mountain lake ecosystems: regionalisation and ecological status

1. A survey of c. 350 remote high altitude and high latitude lakes from 12 different mountain regions across Europe was undertaken to explore ecosystem variability, climate forcing, environmental conditions and pollution threats at a scale not previously attempted.
2. Lakes were sampled for a range of contemporary and sub-fossil organisms including planktonic crustaceans, rotifers, littoral invertebrates, chironomids, diatoms and cladocerans. Survey and cartographic data were used to determine environmental characteristics at each site. Organic pollutants and trace metal concentrations were measured in the lake sediment.
3. A number of separate studies were undertaken which examined the environmental characteristics of the surveyed lakes (climate forcing and chemical composition), distribution of biota relative to local, regional and biogeographical factors and pollution threats (acidification, heavy metals and persistent organic pollutants) to these sensitive ecosystems.
4. There is a strong regional element to the way that environmental factors combine (including climate and pollution threats) and the biota responds in mountain lakes across Europe. From a management perspective it is clear that lake classification and the development of useful typologies and assessments of reference conditions should be undertaken at regional rather than pan-European scales.
5. There are some common features across lake districts related to the timing of industrialisation, but the studies carried out on metals, organic pollutants and nitrate deposition indicate that each lake district has distinct pollution threats. Climate warming already affects most of the lake districts and there are considerable uncertainties as to how this will modify conditions in remote European mountain systems.
6. The lake district concept goes beyond a geographical construct and merits further theoretical and experimental development as an ecological concept.     

Testing for temporal coherence across spatial extents: the roles of climate and local factors in regulating stream macroinvertebrate community dynamics

Temporal coherence or spatial synchrony refers to the tendency of population, community or ecosystem dynamics to behave similarly among locations through time as a result of spatially‐correlated environmental stochasticity (Moran effect), dispersal or trophic interactions. While terrestrial studies have treated synchrony mainly as a population‐level concept, the majority of freshwater studies have focused on community‐level patterns, particularly in lake planktonic communities. We used spatially and temporally hierarchical data on benthic stream invertebrates across six years, with three seasonal samples a year, in 11 boreal streams to assess temporal coherence at three spatial extents: 1) among regions (watersheds), 2) among streams within a region, and 3) among riffles within a stream, using the average of correlation coefficients for stream/riffle pairs across years. Our results revealed the primacy of strongly synchronized climatic factors (precipitation, air temperature) in inducing temporal coherence of macroinvertebrate assemblages across geographically distinct sites (i.e. Moran effect). Coherence tended to decrease with increasing spatial extent, but positive coherence was detected for most biological variables even at the largest extent (about 350 km). The generally high level of coherence reflected the strong seasonality of boreal freshwater communities. A hydrologically exceptional year enhanced the synchrony of biological variables, particularly total macroinvertebrate abundance. Regionally low precipitation in that year led to a substantial decrease in benthic densities across a broad spatial extent, followed by a rapid post‐drought recovery. Coherence at the among‐riffle (within‐stream) extent was lower than expected, implying that local‐scale habitat filters determine community dynamics at smaller spatial extents. Thus, temporal coherence of stream benthic communities appears to be controlled by partly different processes at different spatial scales.     

Temporal coherence of two alpine lake basins of the Colorado Front Range, U.S.A.

1. Knowledge of synchrony in trends is important to determining regional responses of lakes to disturbances such as atmospheric deposition and climate change. We explored the temporal coherence of physical and chemical characteristics of two series of mostly alpine lakes in nearby basins of the Colorado Rocky Mountains. Using year‐to‐year variation over a 10‐year period, we asked whether lakes more similar in exposure to the atmosphere be‐haved more similarly than those with greater influence of catchment or in‐lake processes. 2. The Green Lakes Valley and Loch Vale Watershed are steeply incised basins with strong altitudinal gradients. There are glaciers at the heads of each catchment. The eight lakes studied are small, shallow and typically ice‐covered for more than half the year. Snowmelt is the dominant hydrological event each year, flushing about 70% of the annual discharge from each lake between April and mid‐July. The lakes do not thermally stratify during the period of open water. Data from these lakes included surface water temper‐ature, sulphate, nitrate, calcium, silica, bicarbonate alkalinity and conductivity. 3. Coherence was estimated by Pearson's correlation coefficient between lake pairs for each of the different variables. Despite close geographical proximity, there was not a strong direct signal from climatic or atmospheric conditions across all lakes in the study. Individual lake characteristics overwhelmed regional responses. Temporal coherence was higher for lakes within each basin than between basins and was highest for nearest neighbours. 4. Among the Green Lakes, conductivity, alkalinity and temperature were temporally coherent, suggesting that these lakes were sensitive to climate fluctuations. Water tem‐perature is indicative of air temperature, and conductivity and alkalinity concentrations are indicative of dilution from the amount of precipitation flushed through by snowmelt. 5. In Loch Vale, calcium, conductivity, nitrate, sulphate and alkalinity were temporally coherent, while silica and temperature were not. This suggests that external influences are attenuated by internal catchment and lake processes in Loch Vale lakes. Calcium and sulphate are primarily weathering products, but sulphate derives both from deposition and from mineral weathering. Different proportions of snowmelt versus groundwater in different years could influence summer lake concentrations. Nitrate is elevated in lake waters from atmospheric deposition, but the internal dynamics of nitrate and silica may be controlled by lake food webs. Temperature is attenuated by inconsistently different climates across altitude and glacial meltwaters. 6. It appears that, while the lakes in the two basins are topographically close, geologically and morphologically similar, and often connected by streams, only some attributes are temporally coherent. Catchment and in‐lake processes influenced temporal patterns, especially for temperature, alkalinity and silica. Montane lakes with high altitudinal gradients may be particularly prone to local controls compared to systems where coherence is more obvious.     

A geomorphic template for the analysis of lake districts applied to the Northern Highland Lake District, Wisconsin, U.S.A.

1. We tested the degree to which a lake's landscape position constrains the expression of limnological features and imposes a characteristic spatial pattern in a glacial lake district, the Northern Highland Lake District in north‐central Wisconsin. 2. We defined lake order as a metric to analyze the effect of landscape position on limnological features. Lake order, analogous to stream order, is based solely on geographical information and is simple to measure. 3. We examined the strength of the relationship between lake order and a set of 25 variables, which included measures of lake morphometry, water optical properties, major ions, nutrients, biology, and human settlement patterns. 4. Lake order explained a significant fraction of the variance of 21 of the 25 variables tested with ANOVA. The fraction of variance explained varied from 12% (maximum depth) to 56% (calcium concentration). The variables most strongly related to lake order were: measures of lake size and shape, concentrations of major ions (except sulfate) and silica, biological variables (chlorophyll concentration, crayfish abundance, and fish species richness), and human‐use variables (density of cottages and resorts). Lake depth, water optical properties, and nutrient concentrations (other than silica) were poorly associated with lake order. 5. Potential explanations for a relationship with lake order differed among variables. In some cases, we could hypothesize a direct link. For example, major ion concentration is a function of groundwater input, which is directly related to lake order. We see these as a direct influence of the geomorphic template left by the retreat of the glacier that led to the formation of this lake district. 6. In other cases, a set of indirect links was hypothesized. For example, the effect of lake order on lake size, water chemistry, and lake connectivity may ultimately explain the relation between lake order and fish species richness. We interpret these relationships as the result of constraints imposed by the geomorphic template on lake development over the last 12 000 years. 7. By identifying relationships between lake characteristics and a measure of landscape position, and by identifying geomorphologic constraints on lake features and lake evolution, our analysis explains an important aspect of the spatial organization of a lake district.     

Spatial synchrony of wader populations in inland lakes of the Iberian Peninsula

Spatial synchronization refers to similarity in temporal variations between spatially separated populations. Three mechanisms have been associated with the spatial synchrony of populations: Moran effect, dispersal and trophic interactions. In this study, we explored the degree of spatial synchrony of three wader species populations (Pied Avocet, Black-winged Stilt and Kentish Plover) using monthly estimates of their abundance in inland lakes of the Iberian Peninsula. The effect of several types of wetland variables (structural, hydroperiod and landscape) on spatial synchronization was explored. Groups of lakes with significant synchronization were identified for all three species. The lakes with wastewater input presented longer hydroperiods than those that did not receive these effluents, and this factor was positively related to the spatial synchrony of the Pied Avocet and Kentish Plover populations. The distance between lakes (used as an indicator of the dispersal effect on synchronization) was significant only in Pied Avocet. No structural or landscape variables were related to spatial synchronization in any species. It was impossible to identify any variable related to the spatial synchronization of Black-winged Stilt abundance as a possible result of the high ecological plasticity of this species. Our data provides the first evidence for mechanisms that act on the spatial synchronizing of wader populations in temporary continental lakes in central Spain, and show that the hydroperiod of lakes acts as an important factor in the spatial synchronization of aquatic species and that its effect is mediated by the reception of urban wastewater.     

Landscape influences on climate‐related lake shrinkage at high latitudes

Climate‐related declines in lake area have been identified across circumpolar regions and have been characterized by substantial spatial heterogeneity. An improved understanding of the mechanisms underlying lake area trends is necessary to predict where change is most likely to occur and to identify implications for high latitude reservoirs of carbon. Here, using a population of ca. 2300 lakes with statistically significant increasing and decreasing lake area trends spanning longitudinal and latitudinal gradients of ca. 1000 km in Alaska, we present evidence for a mechanism of lake area decline that involves the loss of surface water to groundwater systems. We show that lakes with significant declines in lake area were more likely to be located: (1) in burned areas; (2) on coarser, well‐drained soils; and (3) farther from rivers compared to lakes that were increasing. These results indicate that postfire processes such as permafrost degradation, which also results from a warming climate, may promote lake drainage, particularly in coarse‐textured soils and farther from rivers where overland flooding is less likely and downslope flow paths and negative hydraulic gradients between surface water and groundwater systems are more common. Movement of surface water to groundwater systems may lead to a deepening of subsurface flow paths and longer hydraulic residence time which has been linked to increased soil respiration and CO₂ release to the atmosphere. By quantifying relationships between statewide coarse resolution maps of landscape characteristics and spatially heterogeneous responses of lakes to environmental change, we provide a means to identify at‐risk lakes and landscapes and plan for a changing climate.     

Spatial scale and ecological relationships between the macroinvertebrate communities of stony habitats of streams and lakes

1. Comparative studies of distinct, but not ecologically isolated, systems such as lakes and streams may improve our understanding of the importance of ecological linkages in aquatic ecosystems. 2. In this study we compared the macroinvertebrate benthos of stony habitats in Swedish lakes and streams. Community composition was used to evaluate zoogeographic patterns and functional feeding guilds were used to identify mechanisms potentially affecting such patterns. 3. Stream communities were generally more diverse and species‐rich and had a higher proportion of grazers, shredders and passive‐filter feeders than lakes. Lake communities had a higher proportion of predators and collector‐gatherers. Of the 10 most common taxa, only Leptophlebia mayflies, clams (Sphaeriidae) and the isopod Asellus aquaticus were recorded in both lakes and streams. 4. Among‐site variance in macroinvertebrate communities accounted for by regional‐scale variables was low (6.4% for lakes and 10.1% for streams), compared with that by local‐scale variables (21% for lakes and 37.6% for streams). For lakes, the among‐site variance in macroinvertebrate communities was best explained by habitat‐scale characteristics followed by ecosystem, riparian, catchment, geographic position and ecoregion. For streams, the variance in macroinvertebrate communities was best explained by ecosystem characteristics followed by habitat, catchment, riparian, ecoregion and geographic position. 5. Conspicuous differences in spatial pattern were revealed between lakes and streams. For lakes, the most unequivocal differences in community composition and function occurred at the transition zone between the mixed forests in the south and the boreal coniferous forests in the north. Surprisingly, streams did not respond as strongly to profound landscape‐level differences in climate and vegetation cover. 6. The spatial differences noted between macroinvertebrate communities of lakes and streams may be because of differences in retention of detrital matter. Our findings imply that detrital inputs are qualitatively similar, but that the retention and processing of coarse particulate organic matter was presumably higher in lake littoral regions than in stream riffle habitats. 7. Although our findings support the conjecture that species distribution is determined fundamentally by conditions prevailing at the local‐scale, regional factors such as land use/type and the role of history were important and seemingly act as strong determinants of large‐scale patterns in biodiversity.     

Unraveling the effects of water–sediment conditions and spatial patterns on Unionida assemblages in seasonally connected floodplain lakes

Floodplain lakes are good metacommunity systems to study the environmental and spatial processes structuring local assemblages. They are more connected during high-water periods and are more isolated during low-water periods. We evaluated the effects of lake spatial patterning and water and sediment conditions on Unionida species assemblages. Moran Eigenvector Maps were used to generate spatial variables representing spatial patterns at different scales. We sampled 35 lakes from the Pantanal floodplain, Brazil. To understand the effects of environmental and spatial variables, we performed Redundancy Analyses and variation partitioning to separate environmental and spatial pattern effects. Environmental variables explained almost twice the variation in the Pantanal mussel assemblages than did spatial variables. Unionida species presence was driven mainly by variations in sediment coarse sand and silt contents. The weak spatial patterns observed may be related to increased connectivity between lakes during floods, which facilitates mussel host fish dispersal. Mussel abundances were driven mainly by organic matter availability, but varied between species. Changes in lake connectivity can affect the regional sediment dynamics and affect mussel assemblages.

     

长江中下游湖群大型底栖动物群落结构及影响因素

长江中下游地区是我国淡水湖泊分布最为密集的区域,其中面积大于10 km²的湖泊总面积占相同级别中国淡水湖泊总面积的51.3%。目前对本地区湖泊大型底栖动物研究主要是关于单个湖泊或几个湖泊之间的比较,将区域内湖泊作为一个整体来分析的研究较少。为揭示现阶段长江中下游浅水湖泊底栖动物群落现状及其主要影响因素,于2008年和2009年夏季对本地区5个湖群69个湖泊大型底栖动物和水化学进行了调查,并分析区域过程和局域环境条件在决定该地区底栖群落结构中的相对重要性。结果表明水体矿化度、电导率及氮磷指标在不同湖群间具有显著差异,而高锰酸盐指数、叶绿素a及营养状态指数无显著差异。密度方面,以寡毛类和摇蚊幼虫为优势类群的湖泊共46个,占总数量的66.7%,以螺类为优势类群之一的湖泊16个,占总数量的23.2%;生物量方面,以螺类为优势的湖泊数量最多(33个),占总数量的46.4%,但以寡毛类和摇蚊幼虫占优势的湖泊亦有27个,占总数量的39.1%,双壳类仅在9个湖泊占据优势。典范对应分析结果表明该地区底栖动物群落结构是局域环境条件和区域过程共同作用的结果,两类因子共解释了33.9%的底栖动物群落变异,其中局域环境因子占被解释量的48.1%,空间变量占35.4%。空间变量较高的解释量表明对整个长江中下游地区湖泊而言,区域过程对底栖动物的分布也起着非常重要的作用。     

Temporal coherence in the limnology of a suite of lakes in Wisconsin, U.S.A.

SUMMARY. 1. Temporal coherence between pairs of lakes over 7 years was measured for thirty-seven limnological variables in seven lakes at the North Temperate Lakes Long Term Ecological Research site in Northern Wisconsin. This analysis tested. First, whether lakes more similar in exposure to the atmosphere were more temporally coherent than lakes which differed more in exposure and, second, whether temporal coherence in lakes progressively decreased from variables more directly influenced by climatic factors such as thermal and hydrologic properties, to those chemical and biological properties which may be less directly influenced by climatic factors. 2. The lakes were a heterogeneous set in respect to exposure to climatic factors as estimated by the ratio of‘lake area/mean depth’and by other morphometric features, and they also differed in their position in the landscape, fertility and fish assemblages. Limnological variables formed a progression from those expected to respond directly to climatic factors to those which would not. They ranged from water level and temperatures to chemical variables such as pH, calcium concentrations and total dissolved phosphorus to biological variables such as chlorophyll concentrations, invertebrate and fish abundances. 3. Coherence was estimated by the correlation between lake pairs for each of the different variables. The mean correlation and the percentage of strong correlations were calculated for each lake pair across all variables, and for each variable across all lake pairs, and both measures of coherence gave similar answers to the questions posed above. 4. Temporal coherence between takes was higher for lakes similar in their exposure to climatic factors; mean correlation (r?) being +0.3 to +0.7 for these lakes and <+0.3 for lakes not similar in exposure. None of the lake pairs had high coherence across all variables. 5. Temporal coherence between lakes was greater for limnological variables directly influenced by climatic factors than for variables either indirectly affected by climate or complexly influenced by other types of factors. Water-level variables had a coherence near 1, r?=0.9. All biological variables had low coherence, some near r?=0.0. Chemical variables more likely to be influenced directly by climatic factors appeared to be more coherent than those more influenced by hydrology or biology. Most silica and phosphorus variables had coherences less than r?=0.15. 6. Coherence was not as strongly related to similarity in landscape position as it was to similarity in exposure to climatic factors, and was not jelated to the proximity of the lake pairs or to their similarity in fertility. 7. A conceptual model was presented to explain how climatic signals are filtered by the lake's exposure to climatic factors and by terrestrially mediated and in-lakc processes to reduce the coherence of lake pairs owing to time lags, frequency shifts and complex interference pattems. 8. Coherence is an important property to evaluate because it influences how broadly we can extrapolate results from a lake or set of lakes (for example, to a lake district) and because manipulative whole lake experiments often use adjacent lakes as reference lakes to evaluate treatment effects. The low coherence in this set of lakes in general and of individual variables such as chlorophyll concentrations and yellow perch year-class strength are discussed. 9. We found no studies that explicitly addressed interlake coherence; one long-term study of forest production made it clear that coherence among sites in a landscape will be a function of scale.     

Landscape spatial patterns in freshwater snail assemblages across Northern Highland catchments

1. Limnologists and landscape ecologists have illustrated how the spatial position of a lake in a landscape influences many of its properties, from the physical to the social. Taking a community ecology perspective, we investigated whether freshwater gastropod assemblages respond to lake landscape position.
2. We determined: (a) whether there is any spatial pattern among lakes in either the species richness or composition of gastropod assemblages; (b) the form of any spatial pattern; and (c) if any explanatory variables (e.g. dispersal corridors and limiting local conditions) show a similar pattern.
3. In three different hydrological catchments, snail species richness increased from isolated highland lakes to stream-connected lowland lakes, probably reflecting increased colonization potential and less limiting local factors for lowland drainage lakes. Catchments appear to differ from one another with regard to relative species abundance, both in terms of macrophyte-associated snail fauna and snails from all habitats aggregated. One or more historical events, such as chance dispersal, may have produced this pattern. Taken together, these results suggest that within-catchment constraints produce repeated gradients in species richness, regardless of what species composition persists in the catchment.