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Arch. Environ. Contam. Toxicol. 39, 523–530 (2000) DOI: 10.1007/s002440010136

A R C H I V E S O F

Environmental Contamination a n d Toxicology © 2000 Springer-Verlag New York Inc.

Development of an Index of Biotic Integrity for the Little Arkansas River Basin, Kansas M. J. Lydy,1 A. J. Strong,1 T. P. Simon2 1 2

Department of Biological Sciences, Wichita State University, Wichita, Kansas 67260-0026, USA U.S. Fish and Wildlife Service, 620 S. Walker Street, Bloomington, Indiana 47403-2121, USA

Received: 26 January 2000/Accepted: 3 July 2000

Abstract. An index of biotic integrity (IBI) was developed for the Little Arkansas River Basin (LARB) in south-central Kansas by establishing a reference condition for the watershed. Twelve metrics, in six categories, were chosen for use in the IBI. Fish assemblages from 30 sites were selected to represent the highest quality sites (reference sites) remaining in the LARB. In addition, 20 historical sites were used to show changes in the watershed over the last century. The modified IBI was then tested at 10 sites within the basin to assess the affects of urban and agricultural disturbances on fish community structure in the Wichita area. IBI scores were statistically lower for the urban versus the agricultural sites. Overall, IBI scores rated from poor to fair, supporting the contention that the fish communities within the LARB are impaired.

The best type of approach for determining the health and integrity of an aquatic system is to use biological communities. Fish communities make good biological indicators for a number of reasons. First, extensive life history information is available for most sport and commercial species of fish, and at least some information is available for all species in North America. Fish species cover a range of trophic levels, and their position at the top of the aquatic food web allows for a comprehensive look at aquatic condition. Fish species are relatively easy to identify, and the public can relate to information relating to fish populations. Fish are usually present in all but the most polluted waters. Finally, fish populations tend to be fairly stable throughout the summer months when most collecting is done, and many species are relatively long-lived, which provides a temporal look at stream conditions (Karr 1981). An index of biotic integrity (IBI) is an ecological approach to biomonitoring incorporating multiparameters of a fish community in a composite index that reflects anthropogenic disturbances in aquatic systems (Karr et al. 1986; Fausch et al. 1990; Simon and Lyons 1995; Simon 1998a). The IBI allows stream health, or the inherent potential of the stream, to be assessed

Correspondence to: M. J. Lydy

based on fish community structure and function. Structural IBI components include species richness, habitat guilds, and numbers of individuals (Simon and Lyons 1995). Functional components consist of feeding and trophic categories, reproductive indices, environmental tolerance, and individual stress and condition groupings (Karr et al. 1986; Fausch et al. 1990; Simon and Lyons 1995). Each attribute of the fish assemblage or “metric” is calibrated based on a regional reference or “least impacted” condition (Hughes 1995). A reference condition is defined as a stream in an area with minimal anthropogenic disturbances. A numerical IBI score is then determined based on a composite value summed from each of the individual metrics and is based on whether the test site substantially or minimally differs from the reference condition. The IBI was first used to assess the biotic integrity of surface water in Midwestern streams and rivers (Karr 1981; Karr et al. 1986). Since those early studies, the IBI has been modified for use regionally and is now considered a family of indices (Simon and Lyons 1995), which have been modified for use internationally (Oberdorff and Hughes 1992). Besides the IBIs original development for warm-water streams, the IBI has been modified for use in coldwater streams (Mundahl and Simon 1999), the littoral zones of the Great Lakes (Minns et al. 1994), great rivers (Goldstein et al. 1994; Simon and Emery 1995), inland lakes (Whittier 1999), and palustrine wetlands (Simon 1998b; Simon and Stewart 1998). The objectives of this study were to develop potential metrics for the Little Arkansas River Basin (LARB) quantify fish assemblage differences among urban versus agricultural landuse sites and provide a baseline for future water quality assessments.

Materials and Methods Study Area The LARB is located in south-central Kansas in the Central Great Plains Ecoregion (Omernik 1987). The river is a moderate-sized stream 165 km in length and drains approximately 3,626 km2. The stream headwaters are located in the Smokey Hills physiographic

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province, and the remainder of the basin lies in the Wellington McPherson lowlands and Arkansas River lowlands physiographic regions (Figure 1). The headwaters of the river originate in Sedgwick County and flow southeast until it joins the Arkansas River in downtown Wichita (KWRB 1975). The LARB differs from streams used in defining the original IBI (Karr 1981; Karr et al. 1986) based on lower species richness and a lack of optimal in-stream habitat.

Reference Site Selection Thirty reference or “least-impacted” sites were chosen from the tributaries and main channel of the LARB so that basin-wide conditions could be characterized (Figure 1). Sites were not spatially distributed equally throughout the watershed because “least impacted” conditions were clumped along the main channel and associated tributaries. The criteria for defining reference sites in the current study included the following characteristics: avoiding point-source discharge; evaluating areas with complete riparian vegetation throughout the sampling reach; locating least-impacted microhabitats within run, riffle, and pool areas; sampling in areas with no impoundments within three meander cycles; avoiding sampling near bridges allowing at least a 100-m buffer zone from the sampling sites (Meador et al. 1993). We also graphically included the results of historical sampling efforts at 20 historical sites. The historical sites scored highest on many of the metrics; therefore, it appears valid to include these sites as “least impacted” sites.

Land-Use Site Selection Ten sites were selected to test the newly developed IBI (Figure 1). These sites were spatially clumped toward the lower end of the basin because we wanted to compare urban Wichita sites with surrounding agricultural sites and because the LARB empties into the Arkansas River in downtown Wichita. Based on a thorough visual assessment both at the sample site and upstream, sites were placed into one of two categories: agricultural or urban. Sites were classified as agricultural (nag ⫽ 5) if more than 50% of the predominant surrounding land use was farming, while urban sites (nu ⫽ 5) had more than 50% of the predominant surrounding land use as commercial, residential, or industrial.

Fish Collection Fish were collected from May to October 1996 for the development of IBI work and from April to October 1998 for land-use assessment. Collection techniques followed the methods of Meador et al. (1993). Briefly, sampling was conducted during the summer and early fall to take advantage of low and stable flow conditions. Reach sampling distance was approximately 15 times the stream width, with a maximum reach length of 300 m. In small headwater streams (less than 10 m wide), a Smith-Root Model 15-C backpack electroshocker was used. Fish sampling on the larger streams (⬎ 10 m), was conducted using one or a combination of the following methods, backpack electroshocker, Smith-Root tote-barge electroshocker, or Smith-Root boat electroshocker. Fish were collected using pulsed direct current with 2– 4 amp output. In addition, each reach also was sampled with a 4.5-m long, 6.35-mm mesh common minnow seine. Block nets were not used during fish sampling due to the relatively large width at many of the sampling sites. Sampling was continued until no new species were captured by repeat sampling of the reach. We combined our catch data from seining and electrofishing to represent relative abundance at each site.

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Upon collection, fish were temporarily held in aerated coolers to keep specimen mortality at a minimum. All fish were identified to species and voucher specimens are housed in the Environmental Toxicology Core Facility at Wichita State University, Wichita, KS. Thirty individuals of each species were weighed, measured for both standard and total length, checked for external anomalies, and returned to the river or stream. Any remaining individuals not processed were identified, counted, checked for anomalies, and then released. Anomalies included eroded fins, lesions, tumors, and ulcers (collectively called DELT, which stands for deformities, eroded fins, lesions, and tumors) (Sanders et al. 1999).

Statistics Relationships between individual IBI metrics and drainage basin area (DBA) were determined using bivariate linear regression (SAS 1996). Average IBI scores were compared among different land-use categories using one-way analysis of variance (ANOVA).

Results and Discussion Development of an IBI for the LARB Metrics. Twelve metrics are presented for the modified IBI that represented the structure and function of fish assemblages in the LARB (Table 1). Guild assignments for the fish species that were collected in the basin were classified for calculation of the IBI metrics using Simon (1998a) (Table 2). Metrics were grouped into six broad categories, including species richness and composition, sensitivity and tolerance, trophic guilds, abundance, reproductive guild, and individual health and condition. We designated each species found in the LARB into appropriate reproductive (Simon 1998c), trophic (Goldstein and Simon 1998), and tolerance guilds (Simon 1991). Species unique to the Little Arkansas River were assigned to the proper guilds based on Cross and Collins (1995) and Pflieger (1997). Reference conditions (Hughes 1995) and the calibrated IBI are based on structural and functional attributes of “leastimpacted” sites in the watershed. These sites were calibrated for each of the candidate metrics by developing maximum species richness (MSR) curves based on collection information from the 30 sampling sites. We also graphically included the results of historical sampling efforts at 20 sites to show changes in the watershed over the last century (Figure 1) (Strong 1997; Strong et al. 1998). We acknowledge that some limitations exist in using historical data collected with different sampling objectives and gear; however, we considered it necessary to include this type of information because of the current lack of high-quality sites in the basin. The historical sites scored highest on many of the metrics, therefore, it appears valid to include these sites as “least impacted” sites. Recently, several authors have used historical collections to describe “least-impacted conditions” in watersheds that have been heavily modified. For example, Hughes et al. (1995) used historical records to examine the reference condition for streams of the Coastal Mountain range of Oregon, and Simon (1998b) used historical collections from palustrine wetlands in the Central Corn Belt Plain to describe the reference condition for dunal, palustrine wetlands in the Lake Michigan watershed

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Fig. 1. Location of present (1996, F) and historical (E) sampling sites used to establish reference conditions for the Little Arkansas River Basin, Kansas. Numbers represent the 50 sampling sites used for establishing reference conditions. In addition, location of the 10 land-use sites are provided (■ ⫽ urban sites and 䊐 ⫽ agricultural sites). Land-use sites are designated with different letters

of northwest Indiana. Both studies used historical data to evaluate species richness and assumed that the early collectors used best available methods to collect “representative” samples in their studies. Our investigation of the LARB also relied on “representative” collection methods that sampled the fish assemblage at 30 sites based on the resident fauna without attempting to get every species or individual. Thus, it is not imperative that the exact same methods be employed but that the historical data represent a single effort on a single date. We did not see large differences in either species diversity or catch-per-unit-effort between the two gear types. Thus, we assume that it is acceptable to follow Simon (1998b) and Hughes et al. (1995) in the use of historical data for defining reference conditions. We see the use of this data as a strength in our manuscript because it shows past conditions before significant changes in land use affected the stream quality of the Little Arkansas River. MSR curves were modified and calibrated specifically for the LARB to assess drainage area relationships and metric expec-

tations (Figure 2). Drainage area was estimated from topographical maps by using a dot grid area estimate method (Barrett and Philbrook 1970). Scoring criteria for each metric was determined by establishing the maximum species richness line so that 95% of the data was beneath the line (Fausch et al. 1984). The remaining area under the MSR curve was then trisected and scores of “1,” “3,” or “5” were assigned to represent whether the area approximated, deviated somewhat, or deviated strongly from the “least impacted” reference condition (Fausch et al. 1984). Many of the individual metric expectations were not dependent on drainage area; however, the total number of species and the number of individuals collected differed according to drainage area. Scoring criteria were then developed to represent modified metrics for all size streams in the LARB watershed (Table 1). Fish collection data for each of the 30 sites sampled for this project were applied accordingly to the 12 metrics modified for the LARB. A score was then assigned dependent on the number of fish applicable to the metric. The metrics were summed

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Table 1. Metrics developed for the index of biotic integrity for use in the Little Arkansas River Basin, Kansas (modified from Karr et al. 1986) Scoring Criteria 5 Species richness and composition 1. Total number of species 2. Number of minnow species 3. Number of centrarchid species 4. Number of benthic invertivore species Sensitivity and tolerance 5. Number of sensitive species 6. Percent individuals as green sunfish Trophic guilds 7. Percent individuals as detritivores 8. Percent individuals as invertivores 9. Percent individuals as carnivores Abundance 10. Relative number of individuals Reproductive guild 11. Percent individuals as simple lithophils Individual health and condition 12. Percent of individuals with DELT

3 Varies with drainage area 3–4 2 2

ⱖ5 ⱖ3 ⱖ3

1

ⱕ2 ⱕ1 ⱕ1

ⱖ4 ⬍ 15%

2–3 15–30%

ⱕ1 ⬎ 30%

⬍ 15% ⬎ 40% ⬎ 10%

15–30% 20–40% 5–10%

⬎ 30% ⬍ 20% ⬍ 5%

Varies with drainage area ⬎ 15%

8–15%

⬍ 8%

⬍ 0.1%

0.1–1.3%

⬎ 1.3%

and a total IBI score and an integrity class assigned for each of the 30 sites. Six different scoring categories (from excellent to no fish) were chosen to describe the biotic integrity and the characteristics of the fish community dependent on the IBI score (Table 3).

chemical environmental disturbances (Page 1983). Darters require high dissolved oxygen concentrations, are intolerant of toxicants and siltation, and thrive over clean substrates. We substituted the number of benthic invertivore species (metric 4) as a functional substitute metric by classifying species that were riffle and run habitat specialists and are benthic invertebrate feeders.

Species Richness and Composition. The total number of species metric (metric 1) was not modified for the LARB. This metric is considered to be one of the most powerful in resolving water resource issues, since a positive correlation exists between high quality resources and the numbers of species for warm-water assemblages (Fausch et al. 1984; Ohio EPA 1988; Simon 1998a; Barbour et al. 1999). Few sites in the LARB, either historically or in recent sampling, possessed adequate number of suckers in order to develop a metric. Therefore, a total count of all species collected that belong to the family Cyprinidae (metric 2) was substituted for this metric, since minnows represent more than half of all of the species of fish found throughout the state (Cross and Collins 1995). Cyprinids are effectively sampled with electrofishing gear and are a dominant family component of Kansas riverine fish fauna. Some cyprinids have very specific habitat and food requirements and are susceptible to habitat and chemical degradation (Gorman 1987). The number of centrarchid species (metric 3) included the number of sunfish species (family Centrarchidae) and included the black basses (Micropterus sp). Simon and Emery (1995) also included black bass in an IBI developed for the Ohio River, and Niemela et al. (1999) used the modified metric for the Red River of the North. Few darter species naturally occur in the LARB. Darters are important environmental indicators because they are insectivorous, benthic habitat specialists, and sensitive to physical and

Sensitivity and Tolerance. The number of sensitive species (metric 5) is used by Ohio EPA (1988) as a modification of the original IBI metric, number of intolerant species. Intolerant species represent the upper 5% of the species that disappear as a response to disturbance. Based on criteria in Karr et al. (1986), few of these species occur in the LARB. The Ohio EPA (1988) suggested that the sensitive species metric includes some moderately intolerant species that decline with decreasing environmental quality and disappear, as viable populations, when the aquatic environment degrades to the “fair” category. Ohio EPA (1988) and Simon (1991) used the sensitive species metric for evaluating sites ranging from headwater to large rivers. The percent of individuals as green sunfish (metric 6) was retained from Karr et al. (1986) as a negative tolerance metric (Table 1). The percentage of green sunfish increases with increasing degradation. It was not necessary to modify this metric by including other tolerant species as in other studies, since green sunfish are commonly collected in most Kansas streams and rivers (Simon 1998a). However, if it becomes necessary, species such as mosquitofish Gambusia affinis, carp Cyprinus carpio, bluntnose minnow Pimephales notatus, and fathead minnow P. promelas should also be included as tolerant. Trophic Guilds. A lack of a suitable definition of omnivores, which narrows eligible species, required the percentage of

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Table 2. Fish from the Little Arkansas River Basin, Kansas, that were used in the development of the index of biotic integrity metrics Minnow species Centrarchid species Benthic invertivores Sensitive species Detritivores Invertivores Carnivores Simple lithophils

Campostoma anomalum, Cyprinella lutrensis, Cyprinus carpio, Notemigonus crysoleucas, Notropis topeka, Notropis ludibundis, Phenacobius mirabilis, Pimephales notatus, Pimephales promelas, Pimephales vigilax Lepomis cyanellus, Lepomis humilis, Lepomis macrochirus, Lepomis megalotis, Micropterus salmoides, Pomoxis annularis Notropis ludibundis, Phenacobius mirabilis, Pimephales vigilax, Moxostoma macrolepidotum, Percina phoxocephala Notropis topeka, Moxostoma macrolepidotum, Fundulus zebrinus, Lepomis humilis, Lepomis megalotis, Percina phoxocephala Dorosoma cepedianum, Cyprinus carpio, Pimephales notatus, Pimephales promelas, Carpiodes carpio, Ictiobus bubalus, Ictiobus niger Notropis ludibundis, Notropis topeka, Phenacobius mirabilis, Pimephales vigilax, Moxostoma macrolepidotum, Fundulus zebrinus, Gambusia affinis, Lepomis humilis, Lepomis megalotis, Percina phoxocephala Pylodictis olivaris, Micropterus salmoides, Pomoxis annularis Dorosoma cepedianum, Phenacobius mirabilis, Carpiodes carpio, Ictiobus bubalus, Ictiobus niger, Moxostoma macrolepidotum, Percina phoxocephala

omnivores metric to be changed to the percentage of detritivores (metric 7). Goldstein and Simon (1998) defined detritivores based on morphological features as having a long-coiled gut and a dark peritoneum. Karr (1981) and Karr et al. (1986) defined omnivores as those species that include a diet of at least 25% plant and 25% animal foods (including detritus) and have the ability (usually indicated by the presence of a long-gut and dark peritoneum) to utilize both. The proportion of invertivores (metric 8) is a suggested alternative to that of the original metric of Karr et al. (1986) the proportion of insectivorous cyprinidae. This modified metric is inclusive of all macroinvertebrates, not just insects, and is intended to respond to a depletion of the benthic macroinvertebrate community that composes the primary food-base for most invertivorous fishes. Invertivorous species are an important link in transferring energy between lower trophic levels to keystone predator species. As disturbance increases, the diversity of macroinvertebrate larvae decreases, triggering an increase in the detritivorous trophic level. Thus, this metric varies inversely with metric 7 with increased environmental degradation. Karr (1981) developed the carnivore metric (metric 9) to measure fish community integrity in the upper trophic levels. It is only in high-quality environments that upper trophic levels are able to flourish. Carnivores are generally not abundant in headwater streams; however, black bass were found in all size streams in the LARB. Karr (1981) suggested that the proportion of carnivores should be a reflection of drainage basin area. Such a correlation in streams greater than 20 mi2 was not found in Midwestern studies (Ohio EPA 1989; Simon 1991) or in the present study. Abundance, Reproductive Guild, and Individual Health and Condition. The relative number of individuals metric (metric 10) was retained from the original IBI (Karr et al. 1986). This metric is used to evaluate fish populations within a stream or river site. Poor water quality will expectedly result in fewer individuals than stream sites of good quality when similar areas and sampling techniques are used. The Ohio EPA (1988) replaced the original index metric, proportion of hybrids (Karr et al. 1986), with the number of lithophilic spawners metric (11), which was used for the LARB IBI as well. The hybrid metric was abandoned because the original intent of the metric was to assess the extent to which

degradation altered reproductive isolation among species. Difficulties of identification, lack of occurrence in headwater and impacted streams, and presence in high quality streams among certain taxa, e.g., cyprinids and centrarchids, caused a lack of sensitivity for the hybrid metric. The percentage of individuals with deformities, eroded fins, lesions, or tumors (DELT) (metric 12) increases in frequency of occurrence as an indication of physical stress due to environmental degradation, chemical pollutants, overcrowding, improper diet, excessive siltation, and other perturbations (Sanders et al. 1999). Leonard and Orth (1986) found that this metric corresponded to increased stream degradation in West Virginia. The presence of black spot is not included in the above analyses, since infestation varies in degree and is a function of the presence of snails, thus it is not solely related to environmental degradation (Berra and Au 1981). Whittier et al. (1987) showed no relationship between Ohio stream quality and black spot. Other parasites were also excluded due to the lack of a consistent relationship with environmental degradation.

Scoring Modifications Rankin and Yoder (1999) suggested that the effect of samples with extremely low numbers in the catch may present a scoring problem in some of the proportional metrics. Adjustments must be made to reduce the possibility of bias toward higher scoring of degraded sites. Aquatic habitats impacted by anthropogenic disturbances may exhibit a disruption in the food base, and the sample may possess very few individuals. At such low population sizes, the normal structure of the community is unpredictable (Rankin and Yoder 1999). Based on Ohio EPA experiences, the proportion of detritivores, insectivorous fishes, and percent individuals affected by anomalies do not always match expected trends at these sample sizes. Although scores are expected to deviate strongly from those of high-quality areas, this is not always observed because of low numbers of individuals or absence of certain taxa. Scoring extremely degraded sites without modifying scoring criteria for the proportional metrics can overestimate the total index score for these sites. The following scoring modifications proposed by Rankin and Yoder (1999) were used when scoring sites in the LARB. The percentage of individuals as detritivores

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Fig. 2. Maximum species richness curve determining scoring criteria for metrics used in a modified index of biotic integrity with increasing drainage area for the LARB, Kansas (n ⫽ 30 F for present and n ⫽ 20 E for historical data)

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Table 3. Total IBI scores, integrity classes, and attributes of the IBI modified from Karr et al. (1986) Total IBI Score

Integrity Class

Attributes

58–60

Excellent

48–52

Good

40–44

Fair

28–34

Poor

12–22

Very poor

Comparable to the best situation without human disturbance; all regionally expected species for the habitat and size, including the most sensitive forms, are present with a full array of age (size) classes; balanced trophic structure Species richness somewhat below expectation, especially due to the loss of the most sensitive species; some species are present with less than optimal abundance or size distributions; trophic structure shows signs of stress Signs of additional deterioration, including loss of sensitive forms, fewer species, and highly skewed tropic structure (e.g., increasing frequency of detritivores); older age classes of top predators may be rare Dominated by detritivores, tolerant forms and habitat generalists; few top carnivores; growth rates and condition factors commonly depressed; hybrids and diseased fish commonly present Few fish present, mostly introduced or tolerant forms; hybrids common; disease, parasites, fin damage and other anomalies regular Repeated sampling finds no fish

No fish

was scored a “1” when fewer than 50 individuals were collected, or were dominated (⬎ 50%) by such generalist feeders as red shiner (Cyprinella lutrensis) and golden shiner (Notemigonus crysoleucas). The percentage of individuals as invertivores, simple lithophils, and individuals possessing DELT anomalies were scored a “1” when fewer than 50 total individuals were collected at a site. In addition, when a high proportion of short-lived fishes were collected in an area that is suspected of chemical contamination, biologists may choose to score the DELT metric a “1,” since short-lived fish will not have had sufficient time to develop anomalies from exposure to chemical contaminants.

Assessment of Biotic Integrity for Different Land-Use Activities The IBI scores differed for the two land-use activities studied. IBI scores at the agricultural sites were statistically higher than scores at the urban sites (F ⫽ 5.3, df ⫽ 1,8, p ⫽ 0.02). The mean IBI score for the agricultural sites was 37.6 ⫾ 1.9 (1 SE), and the mean score for the urban sites was 30.8 ⫾ 1.2 (1 SE). The fish community attributes typical of these integrity scores include deterioration in community structure, including decreases in the number of sensitive species, increases in the number of detritivores and tolerant species, and high numbers of diseased fish. Mean IBI scores have integrity class ratings of “poor” to “fair” for the urban and agricultural sites, respectively. Therefore, the IBI developed for the LARB appears to have successfully reflected the “more degraded” urban sites in comparison to the “less impacted” agricultural sites within the basin. In a companion paper, Eaton and Lydy (2000) did find significantly more organochlorine insecticide residues in fish tissue and sediments in the urban area of Wichita in comparison with surrounding agricultural areas that may help explain the noted difference. Even though there were significant differences among agricultural and urban sites, the IBI scores found overall reflect impacted water quality for the streams within the LARB. Few carnivores were found at any of the 10 sites, and only a single site scored above a 1 for this metric. The percentage as invertivores had less

than a third of the sites scoring above 1, while the percentage of individuals as detritivores for many of the sites was relatively high suggesting that tropic guilds were unstable. Both agricultural and urban areas have contributed to this impairment by impacting water quality and optimal aquatic habitat. For example, agricultural practices in the watershed have resulted in runoff not only of harmful chemicals but also of organic material and sediments that lead to streambed degradation. The increase of bedload sediments eliminated many species of fish that reside or spawn in the interstitial spaces of the streambed. In addition, streambed degradation often eliminates aquatic insect larvae and the species that feed on them. On the other hand, urban areas contribute a wide array of industrial and municipal wastes, and often large amounts of organochlorine pesticides (KDHE 1996). The newly developed IBI calibrated for the LARB has shown that severe degradation and land-use practices have impaired the fish community. This basin is not unique for the state of Kansas and compounding influences have impacted many of the streams (KDHE 1996). This new index provides resource agencies an initial monitoring tool to assess stream conditions in the LARB and provides targets for attaining baseline restoration efforts. However, additional validation of the metrics is needed before the index can be considered fully functional.

Acknowledgments. We appreciate all of the landowners in the Little Arkansas River Basin that allowed us to work on their land and all those that provided field support for this project. This research was supported by a grant from the Kansas Department of Wildlife and Parks.

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