Biological indicators or indices of biotic integrity (IBI) have been developed for land management and regulatory agencies to categorize the condition of a given ecosystem. IBIs are more widely used in lotic systems and those that can be used over wide geographic regions or multiple systems are deemed most valuable. Lacustrine wetlands have intrinsic complexity and multidimensionality making them very difficult to classify. This, in turn, greatly affects the transferability of indices created for explicit regions and wetland types. Similarly, due to scarcity, relatively pristine reference conditions are seldom included in IBI calibration and represent a critical end of the disturbance continuum.

The robustness and transferability of macroinvertebrate and fish IBIs created for fringing lacustrine and drowned river mouth wetlands of the Great Lakes for use in wetlands occurring in inland-lakes on islands within the Great Lakes were tested. Islands within the Laurentian Great Lakes contain unique and critical habitats that have received little attention, but also require specialized tools for monitoring and management. Inland-lake wetlands of Beaver Island, Lake Michigan, were ranked a priori along a disturbance gradient based on adjacent land use/cover. Transferability of the pre-existing Great Lakes IBIs was determined by correlating the site-specific IBI rank with the site-specific disturbance ranks. Results indicated that the IBIs were not directly transferable and may require substantial modification.

Introduction

Traditional approaches to ecosystem health assessment have relied heavily on the analysis of water chemistry and physical parameters (Minns et al., 1994; Drake and Pereira, 2002). These methods, however, often fail to account for effects of anthropogenic habitat alterations, episodic events, and exotic species (Burton et al., 1999; Simon and Lyons, 1995). Fish and macroinvertebrates are natural monitors of the system they inhabit, as they respond predictably to many abiotic and biotic factors (Simon and Lyons, 1995) while integrating time; and thus, can reveal cumulative and episodic disturbances when used as indicators in an index of biotic integrity (IBI). Multimetric indices typically describe the structure and/or function of biotic assemblages, display predictable responses to anthropogenic disturbance, and indicate overall biological condition (Simon and Lyons, 1995; Barbour et al., 1995; Drake and Pereira, 2002; Seilheimer et al., 2009). IBIs are valuable management tools for the identification, establishment, and tracking of wetland conditions through time (Seilheimer et al., 2009). This is important, as wetlands are sensitive and often one of the first habitats to show signs of disturbance from adjacent land use (Uzarski et al., 2005). Degraded wetlands can be monitored throughout the restoration process utilizing IBIs, and relatively pristine wetlands can be identified and protected. The use of biotic indicators also reduces or eliminates costly chemical analyses and has the advantage of being easily implemented and applied to historical datasets (Seilheimer et al., 2009). By integrating the effects of cumulative and episodic impacts, biota serve as bioindicators and can be used to assess overall ecosystem condition.

IBIs are often region and/or habitat specific since the taxonomic groups which IBIs are based are often system-specific. This can greatly limit their applicability to other systems.

The IBIs evaluated in this paper, originally developed to assess Great Lake coastal wetland condition (Burton et al., 1999; Uzarski et al., 2004; Uzarski et al., 2005; Ruetz et al., 2007), were not based entirely upon species-specific metrics, but included some metrics reflecting both trophic structure and patterns of diversity among functional guilds, and therefore, should have wider applicability than those based solely upon individual species (Table 1). Similarly, the fish IBIs (Uzarski et al., 2005; Ruetz et al., 2007) shared many of the same biodiversity, functional group, and species-specific metrics (Table 2), many of which describe trophic roles and reflect food web structure. Finally, the geographic range included in the development and use of the Great Lake coastal wetland IBIs completely surrounds and encompasses Beaver Island. We hypothesized that because of the geographic overlap, similar habitat type (coastal lacustrine wetlands), and the many metrics that focus on ecological function over specific species that the Great Lake coastal wetland IBIs should be readily transferable to Great Lake island inland-lake wetlands.

Table 1.

Invertebrate IBI metric presence across the Burton et al. (1999) and the Uzarski et al. (2004) invertebrate IBIs and the five inland-lakes of Beaver Island. For metric presence in each lake, 1 = present, 0 = absent. An “X” indicates metric presence in the IBI.

MetricsTypha (1999)Inner Schoenoplectus (1999)Outer Schoenoplectus (1999)Inner Schoenoplectus (2004)Outer Schoenoplectus (2004)GeneserathFontGreene'sFoxBarney's
Odonata taxa richness 
(Genera)           
%. Odonata 
Crustacea plus Mollusca taxa 
richness (Genera)           
Total Genera richness 
% Gastropoda 
% Sphaeriidae 
% Amphipoda    
Ephem. plus Trichopt. taxa    
richness (Genera)           
% Crustacea plus Mollusca  
Total number of families    
% Isopoda     
Evenness    
Shannon Diversity    
Simpson index    
MetricsTypha (1999)Inner Schoenoplectus (1999)Outer Schoenoplectus (1999)Inner Schoenoplectus (2004)Outer Schoenoplectus (2004)GeneserathFontGreene'sFoxBarney's
Odonata taxa richness 
(Genera)           
%. Odonata 
Crustacea plus Mollusca taxa 
richness (Genera)           
Total Genera richness 
% Gastropoda 
% Sphaeriidae 
% Amphipoda    
Ephem. plus Trichopt. taxa    
richness (Genera)           
% Crustacea plus Mollusca  
Total number of families    
% Isopoda     
Evenness    
Shannon Diversity    
Simpson index    
Table 2.

Fish IBI metric presence across Uzarski et al.'s (2005) and Ruetz et al.'s (2007) fish IBIs and the five inland-lakes of Beaver Island. For metric presence in each lake, 1 = present, 0 = absent.

MetricDRM SAVTyphaSchoenoplectusGeneserathFontGreene'sFoxBarney's
% Omnivore Abund  
% Piscivore Richness  
% Carnivore (insectivore+piscivore+  
zooplanktivore) Richness         
Smallmouth Bass CPUE  
Insectivorous Cyprinidae Richness  
% Centrarchidae Abund  
Centrarchidae Richness  
Mean evennesss  
Rock Bass CPUE 
Bluegill CPUE  
Lepomis CPUE  
Mean Shannon Diversity Index   
Longnose Gar CPUE   
Largemouth Bass CPUE   
Percent insectivore abundance  
Mean catch per net-night   
Total Richness   
% Non-native richness   
% Insectivorous Cyprinidae abundance   
White Sucker mean CPUE   
Black bullhead mean CPUE   
Alewife mean CPUE   
Pugnose Shiner CPUE   
MetricDRM SAVTyphaSchoenoplectusGeneserathFontGreene'sFoxBarney's
% Omnivore Abund  
% Piscivore Richness  
% Carnivore (insectivore+piscivore+  
zooplanktivore) Richness         
Smallmouth Bass CPUE  
Insectivorous Cyprinidae Richness  
% Centrarchidae Abund  
Centrarchidae Richness  
Mean evennesss  
Rock Bass CPUE 
Bluegill CPUE  
Lepomis CPUE  
Mean Shannon Diversity Index   
Longnose Gar CPUE   
Largemouth Bass CPUE   
Percent insectivore abundance  
Mean catch per net-night   
Total Richness   
% Non-native richness   
% Insectivorous Cyprinidae abundance   
White Sucker mean CPUE   
Black bullhead mean CPUE   
Alewife mean CPUE   
Pugnose Shiner CPUE   

There are over 30,000 islands throughout the Great Lakes basin (US EPA Great Lakes National Program Office; estimates >35,000 from the Great Lakes Information Network), many of which have inland lakes, and few that have received any real scientific attention. Many of these island lakes are at risk from future development, thus developing tools or modifying existing tools to monitor and evaluate ecosystem condition are needed. Toward this end, we evaluated the transferability of the invertebrate and fish IBIs originally developed for specific plant zones (Typha and the inner and outer Schoenoplectus zones), of Great Lakes coastal wetlands (Burton et al., 1999; Uzarski et al., 2004, 2005) and modified versions for drowned river mouth habitats (Ruetz et al., 2007) to the small inland-lakes of Beaver Island. The inland-lakes of Beaver Island (in northern Lake Michigan) provide a case study of IBI transferability that will guide future steps in developing monitoring tools for the wider Great Lake island geography.

Methods

Study sites

Beaver Island, located in northern Lake Michigan, is approximately 51 km northwest of the city of Charlevoix, MI. We sampled Lake Geneserath, Font Lake, Fox Lake, Barney's Lake, and Greene's Lake. Three replicate wetlands (predominately Schoenoplectus stands) from each lake were sampled during the 2009 growing season (June–August) following Great Lakes Coastal Wetland Consortium (GLCWC) fish and invertebrate protocols (Burton et al., 1999; Uzarski et al., 2004, 2005; Ruetz et al., 2007). Fish and invertebrates were sampled and analyzed by wetland (replicate) and subsequently combined in the IBIs.

Disturbance/environmental gradient

A gradient was constructed based upon land use/ cover data in a 250 m buffer zone around each lake to determine relationships with IBIs. Land use/cover data were obtained from the most recent maps from the Michigan Geographic Data Library. ArcGISTM Spatial Analyst was used to create a 250 m buffer zone around each lake (ESRI 2011). The Area Calculator function in XToolsTM was used for the final calculations of area. Seven land cover categories were used in the development of the disturbance gradient: urbanization, transitional lands, agriculture, road length, forest, woody wetland, and herbaceous wetland. Within each of these seven categories are one or more classes identified in the National Land Cover Data (NLCD) Land Cover Classification System Key (NLCD2001). The urbanization category consists of three different types of development: low intensity residential, high intensity residential, and commercial/ industrial/ transportation. Transitional lands include areas of sparse (<25%) cover that are undergoing changes due to land use activities (e.g. forest clearcuts). Agriculture includes pasture/hay, row crops, small grains, fallow, and urban/recreational grasses. The forest, woody wetland, and herbaceous wetland variables were combined into a single large “natural” category in analysis to isolate the disturbance effects. Each of these categories was evaluated as a percentage of the 250 m buffer.

Principal components analysis (PCA) was used to condense the land cover categories into multivariate environmental gradients (Principal components, PCs) which were subsequently related to the IBI scores using Pearson correlations (SAS 9.2). An inspection of the Pearson correlation coefficients, as well as plots of the PCs against the individual IBI metrics, was used to evaluate the transferability of the IBIs. Alpha values were relaxed to α = 0.1 due to the low sample size.

Fish sampling

Fish were sampled in accordance with the Great Lakes Coastal Wetland Consortium (GLCWC) guidelines (Uzarski et al., 2004, 2005). Three randomly selected wetland fragments were selected from each lake and one fyke net was set for one net-night in each. The number of nets fished per lake was approximately proportional to lake perimeter, as we felt this was the best indicator of habitat in these littoral zone dominated lakes (Table 3). The catch was standardized across the nets at each lake to calculate IBI scores. Sites were randomly selected, and based on the minimum inundation required to set nets (approximately 25 cm). Two sizes of fyke nets, both with 4.8 mm mesh, 7.3 m leads, and 1.8 m wings were employed to correspond with varying water depths (GLCWC accepted gears and protocols). Small nets with 0.5 m x 1 m mouths were set in depths from approximately 0.25 m to 0.50 m. Larger nets with 1 m x 1 m mouths were used in water depths greater than 0.50 m. The nets were haphazardly placed perpendicular to the shore, and the wetland of interest, with the mouth at the edge of the vegetation and the entire lead extending through the vegetation. Wings were set at 45 degree angles to the lead. This type of placement ensured that nets would only collect fish from within the vegetated area. Nets were collected after 24 h and fish were identified to species, counted, and immediately released.

Table 3.

Fish and macroinvertebrate sampling effort per lake.

LakeNet-nights fishedDip net sweepsLake perimeter (m)
Geneserath 8300.43 
Fox 2902.43 
Barneys 2512.90 
Font 6572.78 
Greene's 2356.49 
LakeNet-nights fishedDip net sweepsLake perimeter (m)
Geneserath 8300.43 
Fox 2902.43 
Barneys 2512.90 
Font 6572.78 
Greene's 2356.49 

Benthic macroinvertebrate sampling

Three replicate macroinvertebrate samples were collected from each wetland following protocols developed by Burton et al. (1999) and Uzarski et al. (2004). Each replicate was taken from a haphazardly selected wetland point at least 20 m away from any other sampling point. Sampling points were selected to be as visually representative as possible. At each point a standard D-frame dip net containing 0.5mm mesh was swept for two minutes within an area of approximately 3 m2. The net was swept throughout the entire water column from the sediment to the surface and replicate points within each wetland were sampled at various depths to ensure most micro-habitats were sampled and to obtain the most representative sample of each wetland as possible. Contents of the net were placed in white enamel pans for systematic macroinvertebrate collection. Ideally, 150 macroinvertebrates were collected from each sample during a 0.5 person-hour collection period. If 150 invertebrates were not collected during this time, collection continued until the closest increment of 50 was attained. Macroinvertebrates were taken to the laboratory for identification and sorted to genus. The taxonomic keys by Merritt and Cummins (1996) and Thorp and Covich (1991) were used for identification.

Fish and macroinvertebrate IBIs

The invertebrate IBIs that were evaluated were developed by Burton et al. (1999) and Uzarski et al. (2004). The three invertebrate IBIs (Burton et al., 1999) Typha, Uzarski et al. (2004) inner Schoenoplectus, Uzarski et al. (2004) outer Schoenoplectus) were similar in metric composition each IBI having only one completely unique metric that was not shared with one or both of the other IBIs. All of the metrics in the Typha IBI except the metric ‘Relative abundance Amphipoda (%)’ were shared with the inner Schoenoplectus and outer Schoenoplectus IBIs. The inner Schoenoplectus IBI metric ‘Ephemeroptera plus Trichoptera taxa richness (Genera)’ was the single unique metric for this zone. The ‘Total number of families’ metric was the unique metric for the outer Schonoplectus zone. The metric ‘Relative abundance Crustacea plus Mollusca (%)’ was shared between the inner and outer Schoenoplectus IBIs (Table 1).

Table 4.

Land use/land cover percentages in the 250 m buffer zones surrounding each of the five lakes.

Lake Road Length (m) Urban Trans. Agricult. Forest Woody Wetland Herb. Wetland Total% Accounted for 
Barney's 1026 0.35% 2.07% 10.96% 53.21% 15.85% 2.18% 84.61% 
Font 4115 0.16% 0.00% 3.09% 54.73% 35.51% 2.15% 95.64% 
Fox  709 0.00% 0.00% 0.00% 75.86% 12.17% 7.09% 95.12% 
Greene's  692 0.00% 0.00% 0.00% 100.00% 0.00% 0.00% 100.00% 
Geneserath 5120 0.09% 0.00% 1.52% 61.62% 36.05% 0.00% 99.29% 
Lake Road Length (m) Urban Trans. Agricult. Forest Woody Wetland Herb. Wetland Total% Accounted for 
Barney's 1026 0.35% 2.07% 10.96% 53.21% 15.85% 2.18% 84.61% 
Font 4115 0.16% 0.00% 3.09% 54.73% 35.51% 2.15% 95.64% 
Fox  709 0.00% 0.00% 0.00% 75.86% 12.17% 7.09% 95.12% 
Greene's  692 0.00% 0.00% 0.00% 100.00% 0.00% 0.00% 100.00% 
Geneserath 5120 0.09% 0.00% 1.52% 61.62% 36.05% 0.00% 99.29% 

Uzarski et al. (2005) and Ruetz et al. (2007) developed the three fish IBIs evaluated. In the Uzarski et al. (2005) Schoenoplectus fish IBI, seven of the fourteen metrics were unique and unshared with the other two fish IBIs. Five metrics were shared only with the Ruetz et al. (2007) drowned river mouth SAV IBI. One metric, ‘Percent insectivore abundance’, was shared only with the Typha zone IBI. The only metric all three fish IBIs had in common was ‘Rock bass (Ambloplites rupestris) mean catch per net-night’. The Typha fish IBI contained only 3 unique metrics, and shared six of the eleven metrics exclusively with the drowned river mouth SAV IBI. The metric ‘Percent insectivore catch’ was the single metric shared only with the Schoenoplectus zone IBI. Six of the eleven metrics comprising the drowned river mouth SAV IBI were shared with the Typha IBI (Table 2). It is important to note that while the metrics are shared, scoring is different for each unique invertebrate and fish IBI for all zones.

Results

Environmental gradient

Anthropogenic disturbance was relatively minimal around all of the lakes. Forest dominated the land use/land cover, and agriculture was the largest disturbance category around all the lakes (Table 4). The greatest amount of disturbance around these lakes was negligible in comparison to the systems that Uzarski et al. (2004, 2005) studied.

Barney's Lake experienced the greatest amount of disturbance with 11% of land in agricultural activity. The remaining four lakes had substantially less disturbance, with their combined agricultural, urbanization, and transitional land totaling less than 5.5%. Fox Lake and Greene's Lake experienced very little disturbance with only 709 m and 692 m of roads within the buffer respectively. Greene's Lake had least amount of roads and 100% of the land surrounding Greene's was forested. Lake Geneserath contained the largest amount of roads with 5120 m. The roads are predominately unpaved on Beaver Island.

IBI application and transfer

Out of the 23 metrics across all three fish IBIs, eight metrics could not be applied (Table 2). Three of the eight metrics used species that were not present or caught in any of the lakes, including Pugnose Shiner (Notropis anogenus), Alewife (Alosa pseudoharengus), and Smallmouth Bass (Micropterus salmoides). Black Bullhead (Ameiurus melas) was also not prevalent in any of the lakes; however, we used Brown Bullhead (Ameriurus nebulosus) Catch per unit effort (CPUE) to replace this metric as they are of the same genus and functional feeding group (Table 2 in Uzarski et al., 2005). White Sucker (Catostomus commersonii) was only caught in Font Lake which was problematic when applying this metric across all lakes. Insectivorous cyprinids accounted for two of the eight problematic metrics (insectivorous cyprinid richness, and% insectivorous cyprinid abundance). Lake Geneserath was the only lake which contained an insectivorous cyprinid (western Blacknose Dace [Rhinichthys obtusus]) which rendered these metrics unusable. The percent non-native richness metric affected all of the lakes equally, as no invasive species were present in any of the lakes. There were ten metrics comprising the three Burton et al. (1999) macroinvertebrate IBIs, all of which could be evaluated. However, in the Uzarski et al. (2004) macroinvertebrate IBI the relative abundance of isopoda (%) metric could only be evaluated at three of the five lakes due to non-presence in the others (Tables 1 and 2).

Table 5.

Land use disturbance variables for the 250 m buffer zone.

Lake Ranking Lake PC 
l = Relatively Pristine Greene's −1.261 
Fox −1.049 
Genes erath −0.925 
Font −0.221 
5 = Impacted Barney's 3.456 
Lake Ranking Lake PC 
l = Relatively Pristine Greene's −1.261 
Fox −1.049 
Genes erath −0.925 
Font −0.221 
5 = Impacted Barney's 3.456 

Based on land-use patterns, PCA identified Greene's Lake and Fox Lake as the least impacted respectively. Font and Barney's Lake were the most impacted (Table 5). Principal component 1 explained 77.7% of the variation. Three disturbance categories (urbanization, agriculture, and transitional area) were positive loadings on PC1, while undisturbed land area and cumulative road length were negative loadings and were positively correlated with the drowned river mouth SAV IBI (r = 0.84, p = 0.077) (Figure 1). All of the tested macroinvertebrate and fish IBIs showed a positive (though non-significant, p > 0.1) relationship to PC1 (i.e. sites with greater disturbance also had higher IBI scores). The positive relationship between IBI scores and PC1 was opposite of what we hypothesized, though the drowned river mouth SAV fish IBI was the only significant correlation.

Figure 1.

Correlation between principal component 1 for land use/land cover within a 250 m buffer around each lake and the Ruetz et al. (2007) drowned river mouth submerged aquatic vegetation (SAV) fish index of biotic integrity (IBI) applied to the Beaver Island lakes.

Figure 1.

Correlation between principal component 1 for land use/land cover within a 250 m buffer around each lake and the Ruetz et al. (2007) drowned river mouth submerged aquatic vegetation (SAV) fish index of biotic integrity (IBI) applied to the Beaver Island lakes.

Out of all the fish and macroinvertebrate metrics only four macroinvertebrate metrics were correlated to the disturbance gradient as hypothesized and could be used in future recalibration and/or creation of an IBI for the Beaver Island lakes. These metrics included Crustacea plus Mollusca taxa richness (Genera), relative abundance Crustacea plus Mollusca, Simpson Evenness index for macroinvertebrates, and Shannon diversity for macroinvertebrates. There were two primary complications with the remaining fish and macroinvertebrate metrics. First, in many of the correlations between PC1 and the individual metrics the lakes were grouped extremely close, with Barney's Lake often being an outlier (Figure 1). This presented difficulties in the reliable interpretation of metric scores. The second difficulty was that the relationship between the individual metric and PC1 was often bell shaped, making it difficult to recalibrate or create a new IBI.

Appendix A contains all metric and total IBI scores for each IBI examined, as well as the category scores developed by Burton et al. (1999), Uzarski et al. (2004, 2005), and Ruetz et al. (2007) (Appendix A*). The Burton et al. (1999) invertebrate Typha IBI scored all of the lakes in the 21 to 31 range, giving them “moderately degraded” status according to Burton et al. (1999). The 1999 invertebrate inner and outer Schoenoplectus IBIs also gave the lakes “moderately degraded” status with the total IBI scores in the range of 26–34 and 28–36 for the inner and outer Schoenoplectus IBIs, respectively. The Uzarski et al. (2004) invertebrate inner Schoenoplectus IBI scored the lakes in the 37–46 range giving Lake Geneserath and Fox Lake “moderately degraded” status and Barney's Lake, Greene's Lake, and Font Lake “moderately impacted status.” The Uzarski et al. (2004) invertebrate outer Schoenoplectus IBI yielded scores in the 39–45 range giving all of the lakes “moderately impacted” status. The Ruetz et al. (2007) fish IBI scores ranged from 27–50. With the break between relatively “healthy” and “degraded” ecosystems occurring at 33, all of the lakes, aside from Greene's, fell in the “healthy” category.

Discussion

Small lakes and ponds have been historically under-sampled (Downing et al., 2006; Downing, 2010). Only recently has the true impacts and importance of small water bodies to global processes been investigated (Downing et al., 2006; Downing, 2010). With emerging studies showing that small water bodies play a more significant role than previously thought, (Downing et al., 2006; Downing, 2010), it is important to monitor and protect them. This is particularly true of the countless small lakes existing on islands within the Laurentian Great Lakes basin that support evolutionarily unique communities (Hecnar et al., 2002; Harris et al., 2011). The fish and macroinvertebrate IBIs developed by Burton et al. (1999), Uzarski et al. (2004, 2005), and Ruetz et al. (2007) are tools used to evaluate the integrity and ecosystem condition of Great Lakes coastal wetlands and drowned river mouth submerged aquatic vegetation habitats, respectively. They did not, however, effectively transfer to the small inland-lake wetlands of Beaver Island. Thus, new tools need to be developed to support effective resource monitoring and management of island lakes within the Great Lakes basin. The differences in diversity between Great Lakes coastal wetlands and the wetlands of the relatively pristine inland-lakes of Beaver Island rendered several of the metrics evaluated unusable. Metrics are typically based upon the expected characteristics of a specific assemblage in a specific system and ecoregion (Plafkin et al., 1989 in Drake and Valley, 2005). Of the 18 metrics that compose the three fish IBIs we used, ten require specific species’ CPUE data. Four of these species were not present in any of the wetlands sampled. These metrics weighed heavily on the transferability of these IBIs. As a whole, the application of the fish invertebrate IBIs did not produce IBI scores as high as expected, considering the very low amount of anthropogenic disturbance surrounding all of the lakes. Differences in diversity, abundance, ecosystem size and ecosystem type between Great Lakes coastal wetlands and the wetlands of the island inland-lakes may account for this unexpected result; and the development of metrics specific for these inland-lake wetlands on islands within the Great Lakes basin appear to be necessary.

Conclusions

In the development of IBIs, it is important that a broad environmental gradient, including relatively pristine, or the best available reference conditions, be used (Seilheimer, 2009; Brousseau et al., 2011). The lack of transferability of these IBIs may have been related to the lack of truly pristine sites during original IBI development, the narrow disturbance gradient of the inland-lakes, differences between the Great Lakes coastal wetlands and the small, inland-lake wetlands, small sample sizes used to generate and analyze data and/or other unidentified factors (e.g. watershed size) (Figure 2). This study emphasizes the need for a large disturbance gradient including relatively pristine reference conditions, and unique IBIs developed specifically for the Great Lake island inland-lake coastal wetlands identified in this study. The lack of undisturbed reference sites was stated by Wilcox et al. (2002) as a potential problem in the development of IBIs for wetlands. Cooper et al. (2007) sampled macroinvertebrate communities from one heavily impacted system, and three moderately impacted systems, but were unable to detect a relationship between community composition and anthropogenic disturbance. Unfortunately moderately and highly degraded sites are common, whereas more pristine sites are growing less common over time, but are nonetheless vitally important to create a broad enough gradient to detect real changes. An IBI can be an effective tool for evaluation of aquatic ecosystems (Miller et al., 1988; Oberdorff and Hughes, 1992; Kerans and Karr, 1994) when the assumptions used to create the IBI, such as a broad range of environmental conditions and shared habitat conditions, are scientifically based and validated.

Figure 2.

Hypothesized relationship between a complete disturbance gradient (pristine to highly degraded) and IBI scores. Included is the perceived scope of field conditions of the present and earlier IBI studies.

Figure 2.

Hypothesized relationship between a complete disturbance gradient (pristine to highly degraded) and IBI scores. Included is the perceived scope of field conditions of the present and earlier IBI studies.

Acknowledgements

This project was funded by the Faculty Research and Creative Endeavors Committee of Central Michigan University. We thank Central Michigan University Biological Station staff for their help and accommodation. Special thanks are extended to Tom Clement for assistance in the field, as well as Jessica Sherman and Nathan Barton for their review of the manuscript. We would also like to thank the anonymous reviewers whose comments greatly improved earlier versions of this manuscript.

Note

*An appendix for this article is available online at:http://www.aehms.org/Journal_16(3)_Calabro_Appendix.html

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