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.