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Katherine E. Webster

Publications and source records attributed to Katherine E. Webster.

18 recordsLinked to original sources

Abrupt changes in algal biomass of thousands of US lakes are related to climate and are more likely in low-disturbance watersheds.

Climate change is predicted to intensify lake algal blooms globally and result in regime shifts. However, observed increases in algal biomass do not consistently correlate with air temperature or precipitation, and evidence is lacking for a causal effect of climate or the nonlinear dynamics needed to demonstrate regime shifts. We modeled the causal effects of climate on annual lake chlorophyll (a measure of algal biomass) over 34 y for 24,452 lakes across broad ecoclimatic zones of the United States and evaluated the potential for regime shifts. We found that algal biomass was causally related to climate in 34% of lakes. In these cases, 71% exhibited abrupt but mostly temporary shifts as opposed to persistent changes, 13% had the potential for regime shifts. Climate was causally related to algal biomass in lakes experiencing all levels of human disturbance, but with different likelihood. Climate causality was most likely to be observed in lakes with minimal human disturbance and cooler summer temperatures that have increased over the 34 y studied. Climate causality was variable in lakes with low to moderate human disturbance, and least likely in lakes with high human disturbance, which may mask climate causality. Our results explain some of the previously observed heterogeneous climate responses of lake algal biomass globally and they can be used to predict future climate effects on lakes.

continental United States

A continuous classification of the 476,697 lakes of the conterminous US based on geographic archetypes

A variety of classification approaches are used to facilitate understanding, prediction, monitoring, and the management of lakes. However, broad-scale applicability of current approaches is limited by either the need for in situ lake data, incompatibilities among approaches, or a lack of empirical testing of approaches based on ex situ data. We developed a new geographic classification approach for 476,697 lakes ≥ 1 ha in the conterminous U.S. based on lake archetypes representing end members along gradients of multiple geographic features. We identified seven lake archetypes with distinct combinations of climate, hydrologic, geologic, topographic, and morphometric properties. Individual lakes were assigned weights for each of the seven archetypes such that groups of lakes with similar combinations of archetype weights tended to cluster spatially (although not strictly contiguous) and to have similar limnological properties (e.g., concentrations of nutrients, chlorophyll a (Chl a ), and dissolved organic carbon). Further, archetype lake classification improved commonly measured limnological relationships (e.g., between nutrients and Chl a ) compared to a global model; a discrete archetype classification slightly outperformed an ecoregion classification; and considering lakes as continuous mixtures of archetypes in a more complex model further improved fit. Overall, archetype classification of US lakes as continuous mixtures of geographic features improved understanding and prediction of lake responses to limnological drivers and should help researchers and managers better characterize and forecast lake states and responses to environmental change.

conterminous United States

Ecological prediction at macroscales using big data: Does sampling design matter?

Although ecosystems respond to global change at regional to continental scales (i.e., macroscales), model predictions of ecosystem responses often rely on data from targeted monitoring of a small proportion of sampled ecosystems within a particular geographic area. In this study, we examined how the sampling strategy used to collect data for such models influences predictive performance. We subsampled a large and spatially-extensive dataset to investigate how macroscale sampling strategy affects prediction of ecosystem characteristics in 6,784 lakes across a 1.8 million km2 area. We estimated model predictive performance for different subsets of the dataset to mimic three common sampling strategies for collecting observations of ecosystem characteristics: random sampling design, stratified random sampling design, and targeted sampling. We found that sampling strategy influenced model predictive performance such that (1) stratified random sampling designs did not improve predictive performance compared to simple random sampling designs and (2) although one of the scenarios that mimicked targeted (non-random) sampling had the poorest performing predictive models, the other targeted sampling scenarios resulted in models with similar predictive performance to that of the random sampling scenarios. Our results suggest that although potential biases in datasets from some forms of targeted sampling may limit predictive performance, compiling existing spatially-extensive datasets can result in models with good predictive performance that may inform a wide range of science questions and policy goals related to global change.

Ecological Applications

Prediction of lake depth across a 17-state region in the United States

Lake depth is an important characteristic for understanding many lake processes, yet it is unknown for the vast majority of lakes globally. Our objective was to develop a model that predicts lake depth using map-derived metrics of lake and terrestrial geomorphic features. Building on previous models that use local topography to predict lake depth, we hypothesized that regional differences in topography, lake shape, or sedimentation processes could lead to region-specific relationships between lake depth and the mapped features. We therefore used a mixed modeling approach that included region-specific model parameters. We built models using lake and map data from LAGOS, which includes 8164 lakes with maximum depth (Z max ) observations. The model was used to predict depth for all lakes ≥4 ha ( n = 42 443) in the study extent. Lake surface area and maximum slope in a 100 m buffer were the best predictors of Z max . Interactions between surface area and topography occurred at both the local and regional scale; surface area had a larger effect in steep terrain, so large lakes embedded in steep terrain were much deeper than those in flat terrain. Despite a large sample size and inclusion of regional variability, model performance ( R 2 = 0.29, RMSE = 7.1 m) was similar to other published models. The relative error varied by region, however, highlighting the importance of taking a regional approach to lake depth modeling. Additionally, we provide the largest known collection of observed and predicted lake depth values in the United States.

Inland Waters

Small values in big data: The continuing need for appropriate metadata

Compiling data from disparate sources to address pressing ecological issues is increasingly common. Many ecological datasets contain left-censored data – observations below an analytical detection limit. Studies from single and typically small datasets show that common approaches for handling censored data — e.g., deletion or substituting fixed values — result in systematic biases. However, no studies have explored the degree to which the documentation and presence of censored data influence outcomes from large, multi-sourced datasets. We describe left-censored data in a lake water quality database assembled from 74 sources and illustrate the challenges of dealing with small values in big data, including detection limits that are absent, range widely, and show trends over time. We show that substitutions of censored data can also bias analyses using ‘big data’ datasets, that censored data can be effectively handled with modern quantitative approaches, but that such approaches rely on accurate metadata that describe treatment of censored data from each source.

Ecological Informatics

Evidence for regional nitrogen stress on chlorophyll a in lakes across large landscape and climate gradients

Nitrogen (N) and phosphorus (P) commonly stimulate phytoplankton production in lakes, but recent observations from lakes from an agricultural region suggest that nitrate may have a subsidy‐stress effect on chlorophyll a (Chl a ). It is unclear, however, how generalizable this effect might be. Here, we analyzed a large water quality dataset of 2385 lakes spanning 60 regions across 17 states in the Northeastern and Midwestern U.S. to determine if N subsidy‐stress effects on phytoplankton are common and to identify regional landscape characteristics promoting N stress effects in lakes. We used a Bayesian hierarchical modeling framework to test our hypothesis that Chl a –total N (TN) threshold relationships would be common across the central agricultural region of the U.S. (“the Corn Belt”), where lake N and P concentrations are high. Data aggregated across all regions indicated that high TN concentrations had a negative effect on Chl a in lakes with concurrent high total P. This large‐scale pattern was driven by relationships within only a subset of regions, however. Eight regions were identified as having Chl a –TN threshold relationships, but only two of these regions located within the Corn Belt clearly demonstrated this subsidy‐stress relationship. N stress effects were not consistent across other intense agricultural regions, as we hypothesized. These findings suggest that interactions among regional land use and land cover, climate, and hydrogeology may be important in determining the synergistic conditions leading to N subsidy‐stress effects on lake phytoplankton.

Limnology and Oceanography

Creating multithemed ecological regions for macroscale ecology: Testing a flexible, repeatable, and accessible clustering method

Understanding broad-scale ecological patterns and processes often involves accounting for regional-scale heterogeneity. A common way to do so is to include ecological regions in sampling schemes and empirical models. However, most existing ecological regions were developed for specific purposes, using a limited set of geospatial features and irreproducible methods. Our study purpose was to: (1) describe a method that takes advantage of recent computational advances and increased availability of regional and global data sets to create customizable and reproducible ecological regions, (2) make this algorithm available for use and modification by others studying different ecosystems, variables of interest, study extents, and macroscale ecology research questions, and (3) demonstrate the power of this approach for the research question—How well do these regions capture regional-scale variation in lake water quality? To achieve our purpose we: (1) used a spatially constrained spectral clustering algorithm that balances geospatial homogeneity and region contiguity to create ecological regions using multiple terrestrial, climatic, and freshwater geospatial data for 17 northeastern U.S. states (~1,800,000 km 2 ); (2) identified which of the 52 geospatial features were most influential in creating the resulting 100 regions; and (3) tested the ability of these ecological regions to capture regional variation in water nutrients and clarity for ~6,000 lakes. We found that: (1) a combination of terrestrial, climatic, and freshwater geospatial features influenced region creation, suggesting that the oft-ignored freshwater landscape provides novel information on landscape variability not captured by traditionally used climate and terrestrial metrics; and (2) the delineated regions captured macroscale heterogeneity in ecosystem properties not included in region delineation—approximately 40% of the variation in total phosphorus and water clarity among lakes was at the regional scale. Our results demonstrate the usefulness of this method for creating customizable and reproducible regions for research and management applications.

Ecology and Evolution

Effects of land use on lake nutrients: The importance of scale, hydrologic connectivity, and region

Catchment land uses, particularly agriculture and urban uses, have long been recognized as major drivers of nutrient concentrations in surface waters. However, few simple models have been developed that relate the amount of catchment land use to downstream freshwater nutrients. Nor are existing models applicable to large numbers of freshwaters across broad spatial extents such as regions or continents. This research aims to increase model performance by exploring three factors that affect the relationship between land use and downstream nutrients in freshwater: the spatial extent for measuring land use, hydrologic connectivity, and the regional differences in both the amount of nutrients and effects of land use on them. We quantified the effects of these three factors that relate land use to lake total phosphorus (TP) and total nitrogen (TN) in 346 north temperate lakes in 7 regions in Michigan, USA. We used a linear mixed modeling framework to examine the importance of spatial extent, lake hydrologic class, and region on models with individual lake nutrients as the response variable, and individual land use types as the predictor variables. Our modeling approach was chosen to avoid problems of multi-collinearity among predictor variables and a lack of independence of lakes within regions, both of which are common problems in broad-scale analyses of freshwaters. We found that all three factors influence land use-lake nutrient relationships. The strongest evidence was for the effect of lake hydrologic connectivity, followed by region, and finally, the spatial extent of land use measurements. Incorporating these three factors into relatively simple models of land use effects on lake nutrients should help to improve predictions and understanding of land use-lake nutrient interactions at broad scales.

Michigan

Building a multi-scaled geospatial temporal ecology database from disparate data sources: Fostering open science through data reuse

Although there are considerable site-based data for individual or groups of ecosystems, these datasets are widely scattered, have different data formats and conventions, and often have limited accessibility. At the broader scale, national datasets exist for a large number of geospatial features of land, water, and air that are needed to fully understand variation among these ecosystems. However, such datasets originate from different sources and have different spatial and temporal resolutions. By taking an open-science perspective and by combining site-based ecosystem datasets and national geospatial datasets, science gains the ability to ask important research questions related to grand environmental challenges that operate at broad scales. Documentation of such complicated database integration efforts, through peer-reviewed papers, is recommended to foster reproducibility and future use of the integrated database. Here, we describe the major steps, challenges, and considerations in building an integrated database of lake ecosystems, called LAGOS (LAke multi-scaled GeOSpatial and temporal database), that was developed at the sub-continental study extent of 17 US states (1,800,000 km 2 ). LAGOS includes two modules: LAGOS GEO , with geospatial data on every lake with surface area larger than 4 ha in the study extent (~50,000 lakes), including climate, atmospheric deposition, land use/cover, hydrology, geology, and topography measured across a range of spatial and temporal extents; and LAGOS LIMNO , with lake water quality data compiled from ~100 individual datasets for a subset of lakes in the study extent (~10,000 lakes). Procedures for the integration of datasets included: creating a flexible database design; authoring and integrating metadata; documenting data provenance; quantifying spatial measures of geographic data; quality-controlling integrated and derived data; and extensively documenting the database. Our procedures make a large, complex, and integrated database reproducible and extensible, allowing users to ask new research questions with the existing database or through the addition of new data. The largest challenge of this task was the heterogeneity of the data, formats, and metadata. Many steps of data integration need manual input from experts in diverse fields, requiring close collaboration.

Connecticut, Delaware, Illinois, Indiana, Iowa, Ma

Regional variability among nonlinear chlorophyll-phosphorus relationships in lakes

The relationship between chlorophyll a (Chl a ) and total phosphorus (TP) is a fundamental relationship in lakes that reflects multiple aspects of ecosystem function and is also used in the regulation and management of inland waters. The exact form of this relationship has substantial implications on its meaning and its use. We assembled a spatially extensive data set to examine whether nonlinear models are a better fit for Chl a —TP relationships than traditional log-linear models, whether there were regional differences in the form of the relationships, and, if so, which regional factors were related to these differences. We analyzed a data set from 2105 temperate lakes across 35 ecoregions by fitting and comparing two different nonlinear models and one log-linear model. The two nonlinear models fit the data better than the log-linear model. In addition, the parameters for the best-fitting model varied among regions: the maximum and lower Chl a asymptotes were positively and negatively related to percent regional pasture land use, respectively, and the rate at which chlorophyll increased with TP was negatively related to percent regional wetland cover. Lakes in regions with more pasture fields had higher maximum chlorophyll concentrations at high TP concentrations but lower minimum chlorophyll concentrations at low TP concentrations. Lakes in regions with less wetland cover showed a steeper Chl a —TP relationship than wetland-rich regions. Interpretation of Chl a —TP relationships depends on regional differences, and theory and management based on a monolithic relationship may be inaccurate.

Iowa, Maine, Michigan, New Hampshire, Ohio, Wiscon

Long-term citizen-collected data reveal geographical patterns and temporal trends in lake water clarity

We compiled a lake-water clarity database using publicly available, citizen volunteer observations made between 1938 and 2012 across eight states in the Upper Midwest, USA. Our objectives were to determine (1) whether temporal trends in lake-water clarity existed across this large geographic area and (2) whether trends were related to the lake-specific characteristics of latitude, lake size, or time period the lake was monitored. Our database consisted of >140,000 individual Secchi observations from 3,251 lakes that we summarized per lake-year, resulting in 21,020 summer averages. Using Bayesian hierarchical modeling, we found approximately a 1% per year increase in water clarity (quantified as Secchi depth) for the entire population of lakes. On an individual lake basis, 7% of lakes showed increased water clarity and 4% showed decreased clarity. Trend direction and strength were related to latitude and median sample date. Lakes in the southern part of our study-region had lower average annual summer water clarity, more negative long-term trends, and greater inter-annual variability in water clarity compared to northern lakes. Increasing trends were strongest for lakes with median sample dates earlier in the period of record (1938–2012). Our ability to identify specific mechanisms for these trends is currently hampered by the lack of a large, multi-thematic database of variables that drive water clarity (e.g., climate, land use/cover). Our results demonstrate, however, that citizen science can provide the critical monitoring data needed to address environmental questions at large spatial and long temporal scales. Collaborations among citizens, research scientists, and government agencies may be important for developing the data sources and analytical tools necessary to move toward an understanding of the factors influencing macro-scale patterns such as those shown here for lake water clarity.

Illinois, Indiana, Iowa, Michigan, Minnesota, Miss

Cross-scale interactions: Quantifying multi-scaled cause–effect relationships in macrosystems

Ecologists are increasingly discovering that ecological processes are made up of components that are multi-scaled in space and time. Some of the most complex of these processes are cross-scale interactions (CSIs), which occur when components interact across scales. When undetected, such interactions may cause errors in extrapolation from one region to another. CSIs, particularly those that include a regional scaled component, have not been systematically investigated or even reported because of the challenges of acquiring data at sufficiently broad spatial extents. We present an approach for quantifying CSIs and apply it to a case study investigating one such interaction, between local and regional scaled land-use drivers of lake phosphorus. Ultimately, our approach for investigating CSIs can serve as a basis for efforts to understand a wide variety of multi-scaled problems such as climate change, land-use/land-cover change, and invasive species.

Frontiers in Ecology and the Environment

Shifts in controls on the temporal coherence of throughfall chemical flux in Acadia National Park, Maine, USA

Major ion and mercury (Hg) inputs to terrestrial ecosystems include both wet and dry deposition (total deposition). Estimating total deposition to sensitive receptor sites is hampered by limited information regarding its spatial heterogeneity and seasonality. We used measurements of throughfall flux, which includes atmospheric inputs to forests and the net effects of canopy leaching or uptake, for ten major ions and Hg collected during 35 time periods in 1999–2005 at over 70 sites within Acadia National Park, Maine to (1) quantify coherence in temporal dynamics of seasonal throughfall deposition and (2) examine controls on these patterns at multiple scales. We quantified temporal coherence as the correlation between all possible site pairs for each solute on a seasonal basis. In the summer growing season and autumn, coherence among pairs of sites with similar vegetation was stronger than for site-pairs that differed in vegetation suggesting that interaction with the canopy and leaching of solutes differed in coniferous, deciduous, mixed, and shrub or open canopy sites. The spatial pattern in throughfall hydrologic inputs across Acadia National Park was more variable during the winter snow season, suggesting that snow re-distribution affects net hydrologic input, which consequently affects chemical flux. Sea-salt corrected calcium concentrations identified a shift in air mass sources from maritime in winter to the continental industrial corridor in summer. Our results suggest that the spatial pattern of throughfall hydrologic flux, dominant seasonal air mass source, and relationship with vegetation in winter differ from the spatial pattern of throughfall flux in these solutes in summer and autumn. The coherence approach applied here made clear the strong influence of spatial heterogeneity in throughfall hydrologic inputs and a maritime air mass source on winter patterns of throughfall flux. By contrast, vegetation type was the most important influence on throughfall chemical flux in summer and autumn.

Maine

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.

Freshwater Biology

Long-term hydrologic and biogeochemical responses of a soft water seepage lake in north central Wisconsin

Hydrologic and chemical budgets were determined for the period October 1980 through September 1988 for Vandercook Lake, a 43-ha soft water seepage lake located in predominantly sandy outwash in northern Wisconsin. The 1951–1980 mean annual precipitation for the lake, based on nearby National Weather Service (NWS) stations, was 825 mm; volume-weighted p H of bulk atmospheric deposition during the 8-year study averaged 4.7. From October 1980 through September 1983, annual precipitation was 108% of the 30-year NWS average, annual recharge was 144% of the 8-year study period average, and annual groundwater inflow to the lake ranged from 5 to 9% of the total inflow. From October 1986 through September 1988, annual precipitation was 90% of the NWS average, annual recharge was 30% of the study period average, and the lake received no groundwater inflow. During the study, the lake changed from a system whose buffering mechanism was significantly influenced by mineral weathering in inflowing groundwater to one dominated by in-lake sulfate reduction. The functional differences exhibited by this lake during the 8 years of study demonstrate the tenuousness with which conclusions based on shorter-term studies of similar systems must be considered.

Wisconsin

Transient hydrogeological controls on the chemistry of a seepage lake

A solute mass balance method was used to estimate groundwater inflow and outflow rates for Nevins Lake, Michigan, a seepage lake in the upper peninsula that historically has shown extremely variable water chemistry compared with most other seepage lakes. A 4-year study (1989–1992) of the hydrology and geochemistry of Nevins Lake and its contiguous groundwater system revealed that changes in the mass of dissolved solutes are the result of annual hydraulic gradient reversals. A pronounced acidification of Nevins Lake from 1986 to 1988 was likely caused by drought-induced diminished groundwater inflow rates. In this study, dissolved calcium (the major cation in water of Nevins Lake, groundwater, and precipitation) was used for estimating mass flow rates. During the 1989–1992 period, Nevins Lake showed a reproducible annual cycle in calcium mass. Immediately following spring snowmelt and the resulting hydraulic gradient reversal, the mass of dissolved calcium in the lake increases rapidly, and then it decreases steadily throughout the summer and early fall, at which time the lake becomes hydraulically mounded and receives no groundwater inflow. Groundwater flow rates estimated by the solute mass balance method are sensitive to assumed solute concentrations in discharging groundwater. Pore water samples from the lake bed are shown to be more representative of water discharging to the lake than are samples from piezometers near the lake shore, but spatial and temporal variability in pore water chemistry must be considered. Stable isotope analyses ( 18 O and 2 H) of lake water, groundwater, and pore water samples show that water discharging to Nevins Lake in the spring is entirely recycled lake water, and no groundwater derived from terrestrial recharge reaches the lake. The conceptual model formulated during this study linking lake chemistry and the contiguous groundwater system and general groundwater flow patterns surrounding highly transient lake systems are likely transferable to other similar systems.

Michigan

Using landscape limnology to classify freshwater ecosystems for multi-ecosystem management and conservation

Governmental entities are responsible for managing and conserving large numbers of lake, river, and wetland ecosystems that can be addressed only rarely on a case-by-case basis. We present a system for predictive classification modeling, grounded in the theoretical foundation of landscape limnology, that creates a tractable number of ecosystem classes to which management actions may be tailored. We demonstrate our system by applying two types of predictive classification modeling approaches to develop nutrient criteria for eutrophication management in 1998 north temperate lakes. Our predictive classification system promotes the effective management of multiple ecosystems across broad geographic scales by explicitly connecting management and conservation goals to the classification modeling approach, considering multiple spatial scales as drivers of ecosystem dynamics, and acknowledging the hierarchical structure of freshwater ecosystems. Such a system is critical for adaptive management of complex mosaics of freshwater ecosystems and for balancing competing needs for ecosystem services in a changing world.

Iowa, Maine, Michigan, Ohio, Wisconsin

Spatial variation among lakes within landscapes: Ecological organization along lake chains

Although limnologists have long been interested in regional patterns in lake attributes, only recently have they considered lakes connected and organized across the landscape, rather than as spatially independent entities. Here we explore the spatial organization of lake districts through the concept of landscape position, a concept that considers lakes longitudinally along gradients of geomorphology and hydrology. We analyzed long-term chemical and biological data from nine lake chains (lakes in a series connected through surface or groundwater flow) from seven lake districts of diverse hydrologic and geomorphic settings across North America. Spatial patterns in lake variables driven by landscape position were surprisingly common across lake districts and across a wide range of variables. On the other hand, temporal patterns of lake variables, quantified using synchrony, the degree to which pairs of lakes exhibit similar dynamics through time, related to landscape position only for lake chains with lake water residence times that spanned a wide range and were generally long (close to or greater than 1 year). Highest synchrony of lakes within a lake chain occurred when lakes had short water residence times. Our results from both the spatial and temporal analyses suggest that certain features of the landscape position concept are robust enough to span a wide range of seemingly disparate lake types. The strong spatial patterns observed in this analysis, and some unexplained patterns, suggest the need to further study these scales and to continue to view lake ecosystems spatially, longitudinally, and broadly across the landscape.

Ecosystems