Potential of multivariate quantitative methods for delineation and visualization of ecoregions

Multivariate clustering based on fine spatial resolution maps of elevation, temperature, precipitation, soil characteristics, and solar inputs has been used at several specified levels of division to produce a spectrum of quantitative ecoregion maps for the conterminous United States. The coarse ecoregion divisions accurately capture intuitively-understood regional environmental differences, whereas the finer divisions highlight local condition gradients, ecotones, and clines. Such statistically generated ecoregions can be produced based on user-selected continuous variables, allowing customized regions to be delineated for any specific problem. By creating an objective ecoregion classification, the ecoregion concept is removed from the limitations of human subjectivity, making possible a new array of ecologically useful derivative products. A red-green-blue visualization based on principal components analysis of ecoregion centroids indicates with color the relative combination of environmental conditions found within each ecoregion. Multiple geographic areas can be classified into a single common set of quantitative ecoregions to provide a basis for comparison, or maps of a single area through time can be classified to portray climatic or environmental changes geographically in terms of current conditions. Quantified representativeness can characterize borders between ecoregions as gradual, sharp, or of changing character along their length. Similarity of any ecoregion to all other ecoregions can be quantified and displayed as a "representativeness" map. The representativeness of an existing spatial array of sample locations or study sites can be mapped relative to a set of quantitative ecoregions, suggesting locations for additional samples or sites. In addition, the shape of Hutchinsonian niches in environment space can be defined if a multivariate range map of species occurrence is available.     

Topographic, bioclimatic, and vegetation characteristics of three ecoregion classification systems in North America: comparisons along continent-wide transects

Ecoregion classification systems are increasingly used for policy and management decisions, particularly among conservation and natural resource managers. A number of ecoregion classification systems are currently available, with each system defining ecoregions using different classification methods and different types of data. As a result, each classification system describes a unique set of ecoregions. To help potential users choose the most appropriate ecoregion system for their particular application, we used three latitudinal transects across North America to compare the boundaries and environmental characteristics of three ecoregion classification systems [Küchler, World Wildlife Fund (WWF), and Bailey]. A variety of variables were used to evaluate the three systems, including woody plant species richness, normalized difference in vegetation index (NDVI), and bioclimatic variables (e.g., mean temperature of the coldest month) along each transect. Our results are dominated by geographic patterns in temperature, which are generally aligned north-south, and in moisture, which are generally aligned east-west. In the west, the dramatic changes in physiography, climate, and vegetation impose stronger controls on ecoregion boundaries than in the east. The Küchler system has the greatest number of ecoregions on all three transects, but does not necessarily have the highest degree of internal consistency within its ecoregions with regard to the bioclimatic and species richness data. In general, the WWF system appears to track climatic and floristic variables the best of the three systems, but not in all regions on all transects.     

Landscape-level ecological regions: Linking state-level ecoregion frameworks with stream habitat classifications

Regionalization is a form of spatial classification, where boundaries are drawn around areas that are relatively homogeneous in landscape characteristics. The process of delineating ecological regions, or ecoregions, includes the analysis of ecosystem structure. To date, ecoregions have been developed at national and state scales for research and resource management. Stream classification is another method to order the variability of aquatic habitats that spans spatial scales from microhabitat to valley segment. In this study, landscape-level ecoregions are developed for the upper Grande Ronde River basin in northeastern Oregon, 3000 sq km in area. The ecoregion framework presented here is proposed to bridge the gap between stream habitat and state-level ecoregion classifications. Classification at this scale is meant to address issues of management at local scales: to aid in sampling design, in extrapolation of the results of site-specific studies, and in the development of best management practices that are more predictive of ecosystem response than current methods.     

Grouping Lakes for Water Quality Assessment and Monitoring: The Roles of Regionalization and Spatial Scale

Regionalization frameworks cluster geographic data to create contiguous regions of similar climate, geology and hydrology by delineating land into discrete regions, such as ecoregions or watersheds, often at several spatial scales. Although most regionalization schemes were not originally designed for aquatic ecosystem classification or management, they are often used for such purposes, with surprisingly few explicit tests of the relative ability of different regionalization frameworks to group lakes for water quality monitoring and assessment. We examined which of 11 different lake grouping schemes at two spatial scales best captures the maximum amount of variation in water quality among regions for total nutrients, water clarity, chlorophyll, overall trophic state, and alkalinity in 479 lakes in Michigan (USA). We conducted analyses on two data sets: one that included all lakes and one that included only minimally disturbed lakes. Using hierarchical linear models that partitioned total variance into within-region and among-region components, we found that ecological drainage units and 8-digit hydrologic units most consistently captured among-region heterogeneity at their respective spatial scales using all lakes (variation among lake groups = 3% to 50% and 12% to 52%, respectively). However, regionalization schemes capture less among-region variance for minimally disturbed lakes. Diagnostics of spatial autocorrelation provided insight into the relative performance of regionalization frameworks but also demonstrated that region size is only partly responsible for capturing variation among lakes. These results suggest that regionalization schemes can provide useful frameworks for lake water quality assessment and monitoring but that we must identify the appropriate spatial scale for the questions being asked, the type of management applied, and the metrics being assessed.     

Variability of Lotic Macroinvertebrate Assemblages and Stream Habitat Characteristics Across Hierarchical Landscape Classifications

Streams are naturally hierarchical systems, and their biota are affected by factors effective at regional to local scales. However, there have been only a few attempts to quantify variation in ecological attributes across multiple spatial scales. We examined the variation in several macroinvertebrate metrics and environmental variables at three hierarchical scales (ecoregions, drainage systems, streams) in boreal headwater streams. In nested analyses of variance, significant spatial variability was observed for most of the macroinvertebrate metrics and environmental variables examined. For most metrics, ecoregions explained more variation than did drainage systems. There was, however, much variation attributable to residuals, suggesting high among-stream variation in macroinvertebrate assemblage characteristics. Nonmetric multidimensional scaling (NMDS) and multiresponse permutation procedure (MRPP) showed that assemblage composition differed significantly among both drainage systems and ecoregions. The associated R-statistics were, however, very low, indicating wide variation among sites within the defined landscape classifications. Regional delineations explained most of the variation in stream water chemistry, ecoregions being clearly more influential than drainage systems. For physical habitat characteristics, by contrast, the among-stream component was the major source of variation. Distinct differences attributable to stream size were observed for several metrics, especially total number of taxa and abundance of algae-scraping invertebrates. Although ecoregions clearly account for a considerable amount of variation in macroinvertebrate assemblage characteristics, we suggest that a three-tiered classification system (stratification through ecoregion and habitat type, followed by assemblage prediction within these ecologically meaningful units) will be needed for effective bioassessment of boreal running waters.     

The use of terrestrial ecoregions as a regional-scale screen for selecting representative reference sites for water quality monitoring

A process has been developed to select watersheds that are representative of terrestrial ecoregions and that are relatively undisturbed by human activity. Using an existing land classification system at two hierarchical levels of resolution, representative subsets (ecodistricts) of large-scale ecoregions were selected, on the basis of their physiographic, biological, and climatological attributes, to represent the ecoregions. This was achieved using a frequency distribution analysis of existing attribute data and choosing the ecodistrict most closely resembling the most common set of conditions for that ecoregion. Within each ecodistrict, river basins were selected through a best-judgement evaluation of land use, coupled with an assessment of the size and location of each river basin, in order to meet the condition of minimal human impact. Preliminary assessment of water quality data collected from six watersheds selected in this way suggests that the process is effective in targeting regional scale river basins exhibiting the desired characteristics.     

Ecoregions in the Southern Balkans: Should Their Boundaries Be Revised?

Ecoregion delineations have gained increased attention in Europe, especially following the Water Framework Directive 2000/60/EC (WFD), which provides the European Union’s first policy-relevant ecoregion map. However, the WFD’s ecoregions were created through a minor adaptation of Illies’ (Limnofauna Europaea. Gustav Fisher Verlag, Stuttgart, 1967/1978) freshwater zoogeographic regions, and the map’s specific boundaries have not been widely evaluated with respect to the WFD’s uses or their biogeographic accuracy. We examined the WFD ecoregion boundaries in Greece and its neighboring Balkan states by comparing them with the most prominent freshwater biogeographic boundaries as shown by riverine freshwater fish assemblages. Classification and ordination analyses of 23 river basin fish assemblages helped delineate natural faunal break boundaries in freshwater species assemblage distributions depicting major biogeographic barriers to aquatic biota dispersal. However, these biogeographic boundaries differ from those delineated in the WFD map, suggesting boundary errors and inconsistencies in the delineation method of the WFD ecoregions. We reviewed specific boundary disagreements and produced a map showing the region’s most prominent freshwater biogeographic boundaries by charting them on watershed borders among the four biotically dissimilar river basin groups in the southern Balkans. This regional evaluation reveals both a need to reconcile disparate approaches to ecoregion mapping and to promote the development of a new policy-relevant inland waters ecoregion framework that would support broad-scale water management and aquatic conservation.     

Comparing Hydrogeomorphic Approaches to Lake Classification

A classification system is often used to reduce the number of different ecosystem types that governmental agencies are charged with monitoring and managing. We compare the ability of several different hydrogeomorphic (HGM)—based classifications to group lakes for water chemistry/clarity. We ask: (1) Which approach to lake classification is most successful at classifying lakes for similar water chemistry/clarity? (2) Which HGM features are most strongly related to the lake classes? and, (3) Can a single classification successfully classify lakes for all of the water chemistry/clarity variables examined? We use univariate and multivariate classification and regression tree (CART and MvCART) analysis of HGM features to classify alkalinity, water color, Secchi, total nitrogen, total phosphorus, and chlorophyll a from 151 minimally disturbed lakes in Michigan USA. We developed two MvCART models overall and two CART models for each water chemistry/clarity variable, in each case comparing: local HGM characteristics alone and local HGM characteristics combined with regionalizations and landscape position. The combined CART models had the highest strength of evidence (ω_i range 0.92–1.00) and maximized within class homogeneity (ICC range 36–66%) for all water chemistry/clarity variables except water color and chlorophyll a. Because the most successful single classification was on average 20% less successful in classifying other water chemistry/clarity variables, we found that no single classification captures variability for all lake responses tested. Therefore, we suggest that the most successful classification (1) is specific to individual response variables, and (2) incorporates information from multiple spatial scales (regionalization and local HGM variables).     

Ecoregions and ecodistricts: Ecological regionalizations for the Netherlands' environmental policy

For communicating data on the state of the environment to policy makers, various integrative frameworks are used, including regional integration. For this kind of integration we have developed two related ecological regionalizations, ecoregions and ecodistricts, which are two levels in a series of classifications for hierarchically nested ecosystems at different spatial scale levels. We explain the compilation of the maps from existing geographical data, demonstrating the relatively holistic, a priori integrated approach. The resulting maps are submitted to discriminant analysis to test the consistancy of the use of mapping characteristics, using data on individual abiotic ecosystem components from a national database on a 1-km² grid. This reveals that the spatial patterns of soil, groundwater, and geomorphology correspond with the ecoregion and ecodistrict maps. Differences between the original maps and maps formed by automatically reclassifying 1-km² cells with these discriminant components are found to be few. These differences are discussed against the background of the principal dilemma between deductive, a priori integrated, and inductive, a posteriori, classification.     

Development and Validation of an Aquatic Fine Sediment Biotic Index

The Fine Sediment Biotic Index (FSBI) is a regional, stressor-specific biomonitoring index to assess fine sediment (<2 mm) impacts on macroinvertebrate communities in northwestern US streams. We examined previously collected data of benthic macroinvertebrate assemblages and substrate particle sizes for 1,139 streams spanning 16 western US Level III Ecoregions to determine macroinvertebrate sensitivity (mostly at species level) to fine sediment. We developed FSBI for four ecoregion groupings that include nine of the ecoregions. The grouping were: the Coast (Coast Range ecoregion) (136 streams), Northern Mountains (Cascades, N. Rockies, ID Batholith ecoregions) (428 streams), Rockies (Middle Rockies, Southern Rockies ecoregions) (199 streams), and Basin and Plains (Columbia Plateau, Snake River Basin, Northern Basin and Range ecoregions) (262 streams). We excluded rare taxa and taxa identified at coarse taxonomic levels, including Chironomidae. This reduced the 685 taxa from all data sets to 206. Of these 93 exhibited some sensitivity to fine sediment which we classified into four categories: extremely, very, moderately, and slightly sensitive; containing 11, 22, 30, and 30 taxa, respectively. Categories were weighted and a FSBI score calculated by summing the sensitive taxa found in a stream. There were no orders or families that were solely sensitive or resistant to fine sediment. Although, among the three orders commonly regarded as indicators of high water quality, the Plecoptera (5), Trichoptera (3), and Ephemeroptera (2) contained all but one of the species or species groups classified as extremely sensitive. Index validation with an independent data set of 255 streams found FSBI scores to accurately predict both high and low levels of measured fine sediment.     

Regional Representativeness of Swedish Reference Lakes

/ Recent focus has been placed on ecoregion delineations for providing an appropriate framework for monitoring and assessment of region-specific attainable water/habitat quality. Using an ecoregion approach to stratify variance, this study was conducted to determine whether earlier (subjectively) selected Swedish reference lakes may be considered as regionally representative reference sites when compared with a randomly selected lake population. Predictive modeling by discriminant function analysis with lakes classified by ecoregion and lake surface area and six physicochemical variables showed that the greater majority of reference lakes may be considered as regionally representative. The highest proportion of lake "misclassifications" occurred in the boreonemoral ecoregion, a relatively diverse ecoregion of southern Sweden. This apparent bias may be in part be due to the criteria used in selecting regional reference lakes. In the earlier selection of reference lakes emphasis was placed on lakes not being adversely affected by land usage or pollutant discharges, consequently forest lakes were often overrepresented and sites in agricultural areas underrepresented in the selected reference sites. As a complement to predictive modeling, PCA ordination showed the placement of reference lakes within the random lake population and indicated where reference sites might be missing along potentially important ecological gradients. KEY WORDS: Regionalization; Ecoregion; Representativeness; Reference; Ordination; Modeling; Temperate lakes     

Using an ecoregion framework to analyze land-cover and land-use dynamics

The United States has a highly varied landscape because of wide-ranging differences in combinations of climatic, geologic, edaphic, hydrologic, vegetative, and human management (land use) factors. Land uses are dynamic, with the types and rates of change dependent on a host of variables, including land accessibility, economic considerations, and the internal increase and movement of the human population. There is a convergence of evidence that ecoregions are very useful for organizing, interpreting, and reporting information about land-use dynamics. Ecoregion boundaries correspond well with patterns of land cover, urban settlement, agricultural variables, and resource-based industries. We implemented an ecoregion framework to document trends in contemporary land-cover and land-use dynamics over the conterminous United States from 1973 to 2000. Examples of results from six eastern ecoregions show that the relative abundance, grain of pattern, and human alteration of land-cover types organize well by ecoregion and that these characteristics of change, themselves, change through time.     

Magnitude,Frequency, and Duration Relations for Suspended Sediment in Stable (“Reference”) Southeastern Streams1

Abstract: Sediment is listed as one of the leading causes of water‐quality impairments in surface waters of the United States (U.S.). A water body becomes listed by a State, Territory or Tribe if its designated use is not being attained (i.e., impaired). In many cases, the prescribed designated use is aquatic health or habitat, indicating that total maximum daily loads (TMDL) targets for sediment should be functionally related to this use. TMDL targets for sediment transport have been developed for many ecoregions over the past several years using suspended‐sediment yield as a metric. Target values were based on data from “reference” streams or reaches, defined as those exhibiting geomorphic characteristics of equilibrium. This approach has proved useful to some states developing TMDLs for suspended sediment, although one cannot conclude that if a stream exceeds the target range, the aquatic ecosystem will be adversely impacted. To address this problem, historical flow‐transport and sediment‐transport data from hundreds of sites in the Southeastern U.S. were re‐examined to develop parameters (metrics) such as frequency and duration of sediment concentrations. Sites determined as geomorphically stable from field evaluations and from analysis of gauging‐station records were sorted by ecoregion. Mean‐daily flow data obtained from the U.S. Geological Survey were applied to sediment‐transport rating relations to determine suspended‐sediment load for each day of record. The frequency and duration that a given concentration was equaled or exceeded were then calculated to produce a frequency distribution for each site. “Reference” distributions were created using the stable sites in each ecoregion by averaging all of the distributions at specified exceedance intervals. As with the “reference” suspended‐sediment yields, there is a broad range of frequency and duration distributions that reflects the hydrologic and sediment‐transport regimes of the ecoregions. Ecoregions such as the Mississippi Valley Loess Plains (#74) maintain high suspended sediment concentrations for extended periods, whereas coastal plain ecoregions (#63 and 75) show much lower concentrations.     

Evaluating Microbial Indicators of Environmental Condition in Oregon Rivers

Traditional bacterial indicators used in public health to assess water quality and the Biolog system were evaluated to compare their response to biological, chemical, and physical habitat indicators of stream condition both within the state of Oregon and among ecoregion aggregates (Coast Range, Willamette Valley, Cascades, and eastern Oregon). Forty-three randomly selected Oregon river sites were sampled during the summer in 1997 and 1998. The public health indicators included heterotrophic plate counts (HPC), total coliforms (TC), fecal coliforms (FC) and Escherichia coli (EC). Statewide, HPC correlated strongly with physical habitat (elevation, riparian complexity, % canopy presence, and indices of agriculture, pavement, road, pasture, and total disturbance) and chemistry (pH, dissolved O2, specific conductance, acid-neutralizing capacity, dissolved organic carbon, total N, total P, SiO2, and SO4). FC and EC were significantly correlated generally with the river chemistry indicators. TC bacteria significantly correlated with riparian complexity, road disturbance, dissolved O2, and SiO2 and FC. Analyzing the sites by ecoregion, eastern Oregon was characterized by high HPC, FC, EC, nutrient loads, and indices of human disturbance, whereas the Cascades ecoregion had correspondingly low counts of these indicators. The Coast Range and Willamette Valley presented inconsistent indicator patterns that are more difficult to characterize. Attempts to distinguish between ecoregions with the Biolog system were not successful, nor did a statistical pattern emerge between the first five principle components and the other environmental indicators. Our research suggests that some traditional public health microbial indicators may be useful in measuring the environmental condition of lotic systems.     

Ecoregions of the Conterminous United States: Evolution of a Hierarchical Spatial Framework

A map of ecological regions of the conterminous United States, first published in 1987, has been greatly refined and expanded into a hierarchical spatial framework in response to user needs, particularly by state resource management agencies. In collaboration with scientists and resource managers from numerous agencies and institutions in the United States, Mexico, and Canada, the framework has been expanded to cover North America, and the original ecoregions (now termed Level III) have been refined, subdivided, and aggregated to identify coarser as well as more detailed spatial units. The most generalized units (Level I) define 10 ecoregions in the conterminous U.S., while the finest-scale units (Level IV) identify 967 ecoregions. In this paper, we explain the logic underpinning the approach, discuss the evolution of the regional mapping process, and provide examples of how the ecoregions were distinguished at each hierarchical level. The variety of applications of the ecoregion framework illustrates its utility in resource assessment and management.     

A regional framework for establishing recovery criteria

Effective assessments of aquatic ecosystem recovery require ecologically sound endpoints against which progress can be measured. Site-by-site assessments of end points and potential recovery trajectories are impractical for water resource agencies. Because of the natural variation among ecosystems, applying a single set of criteria nationwide is not appropriate either. This article demonstrates the use of a regional framework for stratifying natural variation and for determining realistic biological criteria. A map of ecoregions, drawn from landscape characteristics, formed the framework for three statewide case studies and three separate studies at the river basin scale. Statewide studies of Arkansas, Ohio, and Oregon, USA, streams demonstrated patterns in fish assemblages corresponding to ecoregions. The river basin study in Oregon revealed a distinct change at the ecoregion boundary; those in Ohio and Montana demonstrated the value of regional reference sites for assessing recovery. Ecoregions can be used to facilitate the application of ecological theory and to set recovery criteria for various regions of states or of the country. Such a framework provides an important alternative between site-specific and national approaches for assessing recovery rates and conditions.     

Correspondence between spatial patterns in fish assemblages in ohio streams and aquatic ecoregions

Land classification systems can be useful for assessing aquatic ecosystems if relationships among them exist. Because the character of an aquatic ecosystem depends to a large extent upon the character of the landscape it drains, spatial patterns in aquatic ecosystems should correspond to patterns in the landscape. To test this hypothesis, the US state of Ohio was divided into four aquatic ecoregions based on an analysis of spatial patterns in the combination of land-surface form, land use, potential natural vegetation, and soil parent material. During the period July–October 1983, fish assemblages were sampled in 46 streams that were representative of the ecoregions, and that had watersheds relatively undisturbed by human activities. Spatial patterns of the fish assemblages were examined relative to the ecoregions; distinct regional differences were identified. The assemblages differed most between the Huron/Erie Lake Plain region and the Western Allegheny Plateau region; assemblages in the Eastern Corn Belt Plains and the Erie/Ontario Lake Plain-Interior Plateau regions were intermediate. This pattern also reflects the gradient in landscape character as one moves from the northwest to the southeast of Ohio.     

Perspectives on the nature and definition of ecological regions

Among environmental managers, recognition of the importance of integrating management activities across agencies and programs that have different responsibilities for the same geographic areas has created an awareness of the need for a common hierarchical framework of ecological regions (ecoregions) to implement the strategy. Responding to this need in the United States, nine federal agencies have signed a memorandum of understanding on the subject of developing a common framework of ecoregions. However, considerable disagreement over how to define ecoregions and confusion over the strengths and limitations of existing frameworks stand in the way of achieving this goal. This paper presents some perspectives on the nature and definition of ecoregions related to this confusion and provides a brief overview of the weight of evidence approach to mapping ecoregions, using an example initiated by the US Environmental Protection Agency. To effectively implement ecosystem assessment, management, and research at local, regional, and national levels, research is needed to increase our understanding of ecoregions. We must find ways to illustrate the nature of ecoregion boundaries and the variability of characteristics within ecoregions as they relate to management issues. Research must also be conducted on comparing existing frameworks and developing indices of ecological integrity to effectively evaluate their usefulness.     

Geospatial and Temporal Analysis of a 20‐Year Record of Landsat‐Based Water Clarity in Minnesota's 10,000 Lakes

A large 20‐year database on water clarity for all Minnesota lakes ≥8 ha was analyzed statistically for spatial distributions, temporal trends, and relationships with in‐lake and watershed factors that potentially affect lake clarity. The database includes Landsat‐based water clarity estimates expressed in terms of Secchi depth (SD_Landsat), an integrative measure of water quality, for more than 10,500 lakes for time periods centered around 1985, 1990, 1995, 2000, and 2005. Minnesota lake clarity is lower (more turbid) in the south and southwest and clearer in the north and northeast; this pattern is evident at the levels of individual lakes and ecoregions. Temporal trends in clarity were detected in ~11% of the lakes: 4.6% had improving clarity and 6.2% had decreasing clarity. Ecoregions in southern and western Minnesota, where agriculture is the predominant land use, had higher percentages of lakes with decreasing clarity than the rest of the state, and small and shallow lakes had higher percentages of decreasing clarity trends than large and deep lakes. The mean SD_Landsat statewide remained stable from 1985 to 2005 but decreased in ecoregions dominated by agricultural land use. Deep lakes had higher clarity than shallow lakes statewide and for lakes grouped by land cover. SD_Landsat decreased as the percentage of agriculture and/or urban area increased at county and catchment levels and it increased with increasing forested land.