1 2 Biological Integrity in Urban Streams: Toward Resolving 3 Multiple Dimensions of Urbanization 4 5 B. Michael Walton*a, 6 Mark Sallingb, 7 James Wylesb, 8 Julie Wolina, 9 10 a 11 Cleveland State University, Cleveland OH 44115; 12 b 13 State University, Cleveland, OH 44115; Department of Biological, Geological, and Environmental Sciences, Northeast Ohio Data and Information Center, Levin College of Urban Affairs, Cleveland 14 15 November 19, 2004 16 17 *Corresponding author 18 19 20 21 22 Running head: Biological Integrity in Urban Streams Biological Integrity in Urban Streams Walton et al. -2 23 Abstract 24 Most studies of urban streams have relied on single variables to characterize the degree of 25 urbanization, which may not reflect interactions among features of urban landscapes 26 adequately. We report on an approach to the characterization of urbanization effects on 27 streams that used principal components analysis and multiple regression to explore the 28 combined, interactive effects of land use/land cover, human population demography, and 29 stream habitat quality on an index of biological integrity (IBI) of fish communities. 30 Applied to a substantially urbanized region in northeast Ohio, USA, the analysis 31 demonstrated the interactive nature of urbanization effects. Urban land use and stream 32 habitat quality were significant predictors of IBI, but were no better than and, in some 33 cases, poorer predictors than other gradients and interactions among gradients. High 34 integrity sites were characterized by low forest cover and high grassland cover at sub- 35 catchment scale, but high forest cover within a 500 m radius local zone of the sample 36 point, conditions often found in protected parklands in the region. The analysis also 37 indicated that variability in stream habitat quality was unrelated to landscape or 38 demographic features, a result we attribute to the interaction between the geological and 39 urbanization histories of the region. 40 Keywords: biological integrity, fish, land use, urban streams, water quality 41 42 43 44 45 Biological Integrity in Urban Streams Walton et al. -3 46 1. Introduction 47 Urbanization poses vexing challenges to the ecological sustainability and restoration of 48 stream ecosystems. Stream habitat and biota in urban settings are often profoundly 49 degraded in comparison to natural or less-impacted rural conditions, e.g., (Klein, 1979; 50 Steedman, 1988; Schuler, 1994; May, et al., 1997; Boward, et al., 1999; Morley and Karr, 51 2002; Morse, et al., 2003; Miltner, et al., 2004), even at modest amounts urban 52 development (Weaver and Garman, 1994; Booth and Jackson, 1997). Given these 53 impacts and the accelerating pace of urbanization (Cohen, 2003), there has been great 54 interest in describing quantitative relationships among the intensity of urbanization, 55 constraints on stream recovery, and potential thresholds of degradation imposed by urban 56 development, e.g, reviews by Paul and Meyer (2001) and Allan (2004). 57 However, considerable variability surrounds these general biological integrity- 58 urbanization relationships, so that equivalent levels of urbanization can be associated 59 with a wide range of biological indicator scores (Wang, et al., 1997; Klauda, et al., 1998). 60 For example, streams sites in urban regions of Ohio can vary by more than four-fold in 61 biological integrity within the same level of upstream urban land use (Yoder, et al., 2000; 62 Miltner, et al., 2004). Further, slopes and thresholds of urbanization effects differ among 63 urban regions (Yoder, et al., 1999, 2000; Coles, et al., 2004). 64 Some of this variability is surely attributable to identifiable, allied stressors 65 affecting stream sites, such as point-source pollutants and combined or sanitary sewer 66 outfalls that may exert effects in addition to the generalized impacts of urbanization 67 (Yoder, et al., 2000; Miltner, et al., 2004). However, significant fractions may also be 68 associated with complex interactions among features of urbanized landscapes and the Biological Integrity in Urban Streams Walton et al. -4 69 effects of urbanization within regionally-specific contexts of geology, climate, or history 70 of development and anthropogenic disturbance (Allan, 2004). Such complexity is 71 unlikely to be captured by any single measure of urbanization (Booth and Jackson, 1997; 72 Yoder, et al., 2000; Allan, 2004; Coles, et al., 2004). 73 Hence, a method for quantifying urban effects on streams that integrates multiple 74 dimensions of urbanization and interactions among these factors is desirable. In this 75 report, we describe such an approach and illustrate its application with an analysis of 76 small stream sites in a highly urbanized region of northeastern Ohio, USA. The region 77 has figured prominently in past analyses using single indicators of urbanization, e.g., 78 Yoder et al. (2000), thereby facilitating comparisons with those earlier studies. The 79 analysis integrates the influences of three sets of variables characterizing the urban 80 environment: major land use / land cover features; human population and housing 81 density; and stream habitat quality. We compared the relative impacts of these variables 82 on measures of biological integrity based on fish communities to address the following 83 questions: (1) Are multivariate descriptors of urbanization better predictors of biological 84 integrity than a single variable measure of urban effects, e.g., % urban land, population or 85 housing density?; (2) Does a multivariate approach provide useful, additional insight into 86 effects of urbanization not revealed by single measures of urbanization, e.g., spatial 87 interactions among variables that mitigate or exacerbate the general effects of 88 urbanization, or spatial interactions that represent regionally-specific patterns of urban 89 development?; (3) Do the impacts of urbanization and the interactions among landscape 90 gradients differ with spatial scale?; and (4) Are stream habitat or landscape-level 91 variables better predictors of biological integrity? Biological Integrity in Urban Streams 92 Walton et al. -5 To accomplish this, we used principal components analysis to generate 93 statistically-independent combinations of the original land cover/population demographic 94 variables. Patterns of interaction among the original variables were interpreted from 95 correlations and the magnitude and direction of factor loadings on the principal 96 components. Multiple linear regression was used to test the value of the multivariate 97 components for predicting biological integrity. The question of scale was addressed by 98 comparing the relative influences of stream habitat and landscape-level effects on 99 biological integrity, and by comparing analyses that aggregated the predictor variables at 100 different scales, i.e., the catchment of the biological sampling point vs. 500 m radius 101 “local zone” surrounding the biological sampling point. 102 103 104 2. Study Area The analysis described herein focuses on small stream catchments, for the most 105 part 20-52 km2 drainage area, tributary to the Cuyahoga River in the Cleveland-Akron 106 metropolitan area, as well as a few small streams tributary to Lake Erie within the same 107 geographic area. The Cuyahoga River consists of 1963 km of stream miles draining 2100 108 km2 in northeastern Ohio, USA. The catchment lies within the Erie/Ontario Lake Plain 109 Ecoregion and parts of three different physiographic provinces: the Allegheny Plateau, 110 till plains, and lake plains. 111 The Cuyahoga River watershed is one of the most densely populated, urbanized, 112 and industrialized regions of Ohio. The basin accounts for 2% of total state land, but 17% 113 of Ohio’s population, approximately 1.9 million people (921 people km-2) (Rybka et al., 114 2001). While the upper reaches are dominated by agriculture, the middle and lower Biological Integrity in Urban Streams Walton et al. -6 115 reaches are heavily influenced by urbanization. Two of Ohio’s major cities, Akron and 116 Cleveland, occur within the mid- and lower segments, respectively. The river is 117 characterized by an unusual, U-shaped morphology, formed during the last glacial 118 recession through the merging of several formerly separate drainages. Hence, the 119 Cuyahoga flows southward from its upper reaches then changes direction near Akron, 120 Ohio and flows northward to its terminus at Lake Erie in Cleveland, Ohio (Fig. 1). 121 Because of this shape, eastward urban and suburban expansion is encroaching upon 122 headwater regions that are currently dominated by forest and agriculture. 123 The region has a long history of urbanization, accelerated by completion of the 124 Ohio and Erie Canal in 1825 (Cockrell, 1992). Subsequently, the Cleveland-Akron 125 corridor became one of the major commercial-industrial centers of North America 126 through the first half of the 20th century, with petroleum, steel, rubber, and manufacturing 127 among the major industries (Cockrell, 1992; Rybka et al., 2001). The Cuyahoga Valley 128 National Park, established in 1974, protects 134 km2 of the basin between Akron and 129 Cleveland. However, even this parkland has a history of substantial disturbance, 130 including agricultural, commercial, and industrial uses, as well as many contemporary 131 impacts within and along its boundaries (Cockrell, 1992). Additional environmental 132 challenges arise from economic and infrastructural decline and population loss from city- 133 centers and older, inner-ring suburbs, and out-migration to the urban fringe (Bier, 1993, 134 2001). 135 136 3. Methods 137 3.1 Biological Integrity and Stream Habitat Data Biological Integrity in Urban Streams 138 Walton et al. -7 Biological integrity and stream habitat data were extracted from a statewide 139 database maintained by the Ohio Environmental Protection Agency (OEPA). These data 140 have served as the basis for previously published analyses regarding the Cuyahoga and 141 other Ohio watersheds (Yoder, et al., 1999, 2000; Miltner, et al., 2004) and are central 142 component of the state-wide program for water quality assessment a. Biological data 143 consisted of multimetric indices of biological integrity (IBI) and well-being (modified 144 index of well-being, MIWB) based upon fish communities. The IBI is an aggregate index 145 based upon 12 sub-metrics characterizing the taxonomic composition, trophic structure, 146 abundance, and condition of the fish community (Karr, 1981; Karr, et al., 1986), as 147 modified for Ohio streams and rivers (Ohio EPA, 1987b, 1989b; Yoder and Rankin, 148 1995). The MIWB is an index that incorporates measures of abundance, biomass, and 149 diversity (Gammon, 1976; Gammon, 1980; Hughes and Gammon, 1987), as modified to 150 increase sensitivity to conditions particular to Ohio’s streams and rivers (Ohio EPA, 151 1987b, 1989b; Yoder and Rankin, 1995). Stream habitat data were also extracted from 152 the statewide database as the Qualitative Habitat Evaluation Index (QHEI). The QHEI is 153 a qualitative assessment of major features of stream habitats presumed to influence the 154 potential for healthy fish communities (Rankin, 1989, 1995). 155 156 157 3.2 Land Use and Census Data Land use data were extracted from a statewide land cover inventory of Ohio 158 produced by the Ohio Department of Natural Resources, based upon Landsat Thematic 159 Mapper Data collected in September and October 1994 (Schaal and Motsch, 1997), the 160 approximate mid-point of the time frame for the biological data used here. The data were Biological Integrity in Urban Streams Walton et al. -8 161 classified into seven land cover categories of urban, agriculture/open urban areas, 162 shrub/scrub, wooded, open water, non-forested wetlands and barren. Population density 163 and housing density data were obtained from the U.S. Census Bureau, 2000 Census of 164 Population and Housing, Summary File 1, 2001. Age of housing data were extracted from 165 the U.S. Census Bureau, 2000 Census of Population and Housing, Summary File 3, 2002. 166 167 168 3.3 Delineation of sample point catchments The basic spatial unit of analysis in this study was the catchment area for each of 169 the biological sampling points. The sample-point catchment area was defined using 170 digital elevation models (DEM), a vector hydrography database, and sample point 171 locations. The DEM used here was extracted from the 1:24,000-scale seamless US 172 Geological Service (USGS) National Elevation Dataset (NED) which was developed by 173 merging USGS’s highest resolution, best quality elevation data available (NED is 174 accessible on-line at gisdata.usgs.net/ned/default.asp). 175 To improve accuracy of stream and catchment delineation, we used a vector 176 hydrography database, the Valley Stream Segment (VST Rivers) file, to adjust the DEM. 177 The VST file is based upon a 1:100,000 base map from the National Hydrography 178 Dataset (NHD) from the USGS and the U.S. Environmental Protection Agency (USEPA). 179 The two main sources for information for this dataset are USGS digital line graphs and 180 the USEPA Reach File Version 3 (http://nhd.usgs.gov/). To a improve catchment 181 delineation, raster cells were adjusted in elevation at or near the VST vector layer 182 streams, thereby improving stream channel results. The adjusted raster elevation values 183 were then used to create a new vector-based stream network which included only the Biological Integrity in Urban Streams Walton et al. -9 184 streams recognized by the original hydrography stream layer but were precisely located 185 in reference to the DEM and the slope, flow direction, and related information necessary 186 to delineate catchment areas up stream and up slope from the sample points. 187 GIS surface analysis tools were used to create catchment area polygons. Figure 1 188 illustrates the points, VST Rivers hydrography layer, the revised DEM-based 189 hydrography network, and the catchment polygons in a portion of the study area. In 190 addition to the sample point catchments, we also delineated 500 m radius local zone sub- 191 polygons for each sample point. These were defined by inscribing a 500 m radius around 192 the sample point within the boundaries of the original catchment area (Fig. 1). 193 Population and housing unit counts are available at the census block level. 194 Because catchments and local zones split census blocks and block groups, these census 195 data were estimated by areal interpolation, specifically area apportionment. This method 196 apportions the data based on the relative areas of the block or block group that are 197 contained in each part split by catchments or zones. Geographic (polygon) boundary files 198 in computerized GIS database structure for census blocks and block groups are available 199 from the Census Bureau's Topologically Integrated Geographically Encoded Referenced 200 (TIGER) database. 201 202 203 3.4 Statistical Procedures The biological, land use, population demographic and stream habitat variables 204 used in this study are listed in Table 1. Biological data (IBI and MIWB) served as 205 dependent variables in these analyses. Also, IBI was decomposed into its 12 sub-metrics, 206 and these also were analyzed as dependent variables. QHEI served as both a dependent Biological Integrity in Urban Streams Walton et al. -10 207 and independent variable. QHEI was the dependent variable in regressions evaluating 208 land use and demographic gradients as predictors of stream habitat quality, whereas 209 QHEI was entered as an independent variable in regressions seeking to predict IBI, IBI 210 sub-metrics, or MIWB. 211 All variables were evaluated for conformance to normality and transformed, if 212 necessary, using appropriate transformations. Indices, counts, and density measures were 213 transformed according to log10X or log10(X+1), whereas proportion variables were arcsin 214 square-root transformed. Pearson-product moment correlations and linear regression 215 analyses were used to explore urbanization-stream quality relationships urbanization, as 216 measured using % urban land use and population or housing density. 217 To explore the contributions of landscape gradients in addition to urban land use 218 and to assess the potential for interactions among landscape and demographic gradients, 219 we employed principal components and multiple linear regression analyses. Principal 220 components analyses reduced the dimensionality among the independent variables and 221 produced gradients, i.e., principal components, which were independent and appropriate 222 for multiple regression analysis. Principal components were obtained for the land 223 use/land cover and population demography dataset. Distance of the sample point from 224 stream terminus and area of the sample-point catchments were also included within these 225 data since these two spatial variables were correlated with several of the landscape and 226 demographic features. Only components with eigenvalues > 1 were retained for 227 subsequent analyses. Principal components were interpreted based upon magnitude of 228 factor loadings and inspection of bivariate plots of components against the original 229 variables. Biological Integrity in Urban Streams 230 Walton et al. -11 The predictive value of principal components for biological integrity or habitat 231 quality was assessed using stepwise multiple regression (forward and backward selection 232 procedures, P=0.05 for entry or removal from the model). R2-change was used to assess 233 and rank the proportional contribution of each significant predictor to overall variance 234 explained by regression models. Julian date of the biological sample was entered into 235 these analyses to adjust for temporal changes in biological integrity. All statistical 236 procedures were conducted using SPSS for Windows, Version 11. 237 238 4. Results 239 4.1 Single Indices of Urbanization 240 The index of biological integrity (IBI) was significantly correlated with variables 241 that have been commonly used as measures of urbanization, e.g., % urban land use and 242 population density, although the magnitudes of these correlations were generally low, and 243 several other land use variables were more strongly correlated with IBI than urban land 244 use, e.g., grassland cover (Table 2). Percent urban land use and population density were 245 also found to be significant predictors, in combination with QHEI, of IBI by multiple 246 linear regression analysis, although only population density retained statistical 247 significance as a predictor of IBI in regressions that included two spatial covariates of 248 IBI, distance from terminus and catchment area (Table 3). 249 250 251 252 4.2 Multivariate Indices of Urbanization: Sub-catchment Scale Analyses Principal components based upon variables aggregated at the sample point subcatchment scale yielded four components with eigenvalues > 1, which accounted for Biological Integrity in Urban Streams Walton et al. -12 253 67.5% of total variation in the data set. Factor loadings for the components are shown in 254 Table 4, where principal components are labeled SP1, SPC2, etc., to indicate that they are 255 based upon sub-catchment scale data. SPC1 was most strongly influenced by catchment 256 area with lesser loadings associated with housing density, % barren, and % open water 257 cover. SPC2 increased with % wetland and shrub/scrubland cover. SPC3 described a 258 contrast between % forest and % grass/park/field cover (Fig. 2). SPC4 was strongly, 259 positively correlated with urban land cover. 260 No component was correlated significantly with QHEI (Table 4), whereas two 261 components, SPC1 and SPC3, were correlated significantly with IBI. Three principal 262 components (1, 3, and 4) were significant predictors IBI according to stepwise, multiple 263 linear regression (Table 5). IBI increased with SPC1, but declined with SPC3 and SPC4. 264 Multiple regression also indicated that IBI increased significantly with QHEI and julian 265 day (Table 5). The modified index of well-being (MIWB) was also significantly related 266 to principal components and IBI. MIWB also increased with SPC1, as well as SPC2 267 (Table 5). As in the case for IBI, MIWB increased with QHEI and julian day. 268 The multiple regression relating principal components to IBI had a substantially 269 higher coefficient of determination (R2) than any of the regressions relating principal 270 components to submetrics of the IBI (Table 5). Nevertheless, each of the principal 271 components was a significant predictor of at least two of the sub-metrics of the IBI. SPC3 272 showed the highest number of significant coefficients (it was a predictor of 6 of 12 sub- 273 metrics, range of P-values = 0.017 - <.001, Table 5). This component was negatively 274 related, generally, to sub-metrics representative of species richness and community 275 composition (e.g., number of native species, number of darter species, number of Biological Integrity in Urban Streams Walton et al. -13 276 sensitive species) and was positively related to a measure of fish condition, the proportion 277 of individuals with deformities, eroded fins, lesions, or tumors (Table 5). SPC2 was a 278 significant predictor of 5 of 12 submetrics, despite failing to emerge as a significant 279 predictor of overall IBI. SPC2 was positively related to two measures of community 280 composition (metrics of native and darter species), two measures of trophic structure (% 281 omnivores and insectivores), and was negatively related to % lithophile species. SPC1 282 was positively related to 4 of 12 submetrics which were indicative of abundance or 283 species richness and community composition (e.g., number of native species, number of 284 darter species, number of insectivores). SPC4 was negatively related to 2 submetrics (% 285 insectivores and number of individuals). QHEI was a significant predictor of 7 of 12 286 submetrics, which were largely measures of species richness and community 287 composition. Number of darter species increased during the time period within this data 288 set, while proportions of tolerant species, omnivores, and individuals with deformities, 289 eroded fins, lesions, and tumors declined (Table 5) 290 291 292 4.3 Multivariate Indices of Urbanization: Effects of Scale The addition of variables describing land use/land cover within the 500 m radius 293 neighborhood of the biological sampling point resulted in identification of eight principal 294 components with eigenvalues > 1, accounting for 80% of the total variance within the 295 dataset. Factor loadings for this analysis are shown in Table 6, where principal 296 components are labeled “LPC” to indicate that 500 m local zone data were included in 297 their calculation. Biological Integrity in Urban Streams Walton et al. -14 298 LPC1 was strongly related to population and housing density at both 500 m local 299 zone and the overall sample sub-catchment scales. LPC1 also increased with urban land 300 use at both scales. LPC2 was most strongly related to basin area at the sub-catchment 301 scale. LPC3 was strongly related to wetland cover at both the sub-catchment and 500 m 302 local scale. LPC4 was principally a descriptor of shrub/scrub cover at the two scales. 303 LPC5 decreased strongly with increasing forest cover in the 500 m zone, but increased 304 significantly with increasing grass/park/field cover and urban land cover in the 500 m 305 local zone. LPC6 was essentially a descriptor of barren lands at both scales. LPC7 was 306 strongly related to open water cover at the 500 m scale, and to a lesser extent to water at 307 the sub-catchment scale. LPC8 described a contrast between grass/park/field cover and 308 forest cover at the sub-catchment scale. 309 As in the analysis conducted at the sub-catchment scale exclusively, none of these 310 components were correlated with QHEI (Table 6). Six of the eight components (LPC1, 311 LPC2, LPC3, LPC4, LPC5, and LPC8) were correlated significantly with IBI (Table 6), 312 but only five of these (LPC4 excluded) was predictive of IBI based upon stepwise 313 multiple regression analysis (Table 7). LPC2 and LPC3 were predictive of MIWB (Table 314 7). IBI and MIWB also increased with QHEI and julian day. 315 Once again, the multiple regression predicting IBI had a substantially higher 316 coefficient of determinantion (R2 = .44) than regressions for each of the IBI sub-metrics 317 (range of R2 = .09 - .35), although each of the sub-metrics showed a significant 318 regression with at least one of the principal components. In general, sub-metrics of the 319 IBI responded to the same gradients as did overall IBI. However, LPC4, which did not 320 emerge as a significant predictor of IBI, was associated 6 of 12 submetrics. LPC4 was Biological Integrity in Urban Streams Walton et al. -15 321 negatively associated with measures of species richness and community composition and 322 trophic structure, but positively associated with percent of individuals with deformities, 323 eroded fins, lesions and tumors (Table 7). LPC7, which also showed no predictive value 324 for IBI overall, was positively associated with the proportion of insectivores in samples. 325 326 5. Discussion 327 5.1 Are multivariate measures more informative than univariate measures of 328 329 urbanization? Several authors have indicated that single measures of urbanization are unlikely to 330 be sufficient for assessing the ecological health of urban streams (Booth and Jackson, 331 1997; Karr and Chu, 1999; Yoder, et al., 2000; Morley and Karr, 2002), although no 332 previous study has conducted a comparison of the relative power of bivariate and 333 multivariate approaches for the same dataset. Previous analyses for northeast Ohio based 334 upon a subset of the data used here found significant, negative relationships between 335 urban land use and biological integrity based upon fish and invertebrate community 336 quality and that a threshold for significant degradation of fish communities, as measured 337 by IBI, occurred at 8% urban land use within the Cuyahoga River basin (Yoder, et al., 338 1999, 2000). Similarly, we found that IBI declined with % urban land use or population 339 density (Tables 2 and 3, Fig. 3). Further, sites for which the multivariate combination of 340 land use, population, housing, and stream habitat data predicted IBIs greater than 41 all 341 had observed IBIs exceeding the minimum value required for attainment of Ohio EPA 342 warm water habitat (WWH) use criterion (Fig. 4). The average % urban land use for this 343 group of sites, 6.5% ± 2.3% (N= 7) was essentially indistinguishable from the 8% Biological Integrity in Urban Streams Walton et al. -16 344 threshold value based on % urban land use alone. The majority of sites with predicted 345 IBIs lower than 41 failed to meet WWH use attainment and showed poor IBIs overall. No 346 sites with predicted IBIs below 26 achieved WWH status and the average % urban land 347 use for this group was 24.6% ± 4.4% (N = 50). In an analysis of streams in the Columbus, 348 Ohio area, Miltner et al. (2004) report a similar upper threshold of % urban land use 349 (27.1%), above which stream sites failed to achieve WWH status. 350 Hence, the multivariate approach used here identified management and 351 assessment thresholds largely equivalent to previous analyses based on bivariate 352 approaches. However, the multivariate approach revealed interactions among landscape 353 and demographic variables that could not be assessed with a single measure of 354 urbanization. In particular, the importance of urban land use recedes in multivariate 355 analyses, where other gradients and interactions among gradients emerge as more 356 important predictors. At the sub-catchment scale, the fourth principal component (SPC4) 357 most strongly represented urban land use. Although this component emerged as a 358 significant predictor of IBI (Table 5), SPC4 accounted for only 1.8% of variance in IBI. 359 In comparison, SPC3, which described a counter-gradient of forest versus grassland 360 cover, accounted for 9.1% of variance in IBI. 361 When 500 m local zone land use/land cover was entered into the analysis, urban 362 land use receded even farther into the background. Although urban land use at sub- 363 catchment or 500 m scale loaded significantly on several principal components, loadings 364 were relatively low (Table 6). In the local zone analysis, urban land use made its 365 strongest contribution to LPC1, which was even more strongly related to population and 366 housing density (Table 6). However, this component had predicted IBI only weakly, Biological Integrity in Urban Streams Walton et al. -17 367 accounting for only about 3.3% of variance in biological integrity. Components 1, 2, 3, 5, 368 and 8 describing other aspects of land use/land cover, including percent forest, wetland, 369 and grasslands in the sub-catchment and within the 500 m local zone, all accounted for 370 more variance in IBI (5.6-6.5%, accounting for 24% variance in total). 371 What accounts for the reduction in the influence of urban land use within these 372 analyses? Certainly one important factor is that much of the area is either heavily 373 urbanized or at least suburbanized to some degree, so that the effects of urbanization are 374 pervasive but the gradient of urbanization is relatively small. Mean urban land use 375 among the sites used in this study is relatively high (16%), even though many of the sites 376 are outside the urban core or are found within parklands or forested ravines. This level of 377 urbanization has been associated with substantial, and perhaps irreversible, biological 378 degradation (Steedman 1988, Booth and Jackson 1997, Yoder et al.1999). 379 It is also likely that these analyses reflect the long history of anthropogenic 380 disturbance within the region. Stream biota can reflect the historical legacy of past 381 stressors and land uses long after those factors have changed (Harding, et al., 1998). 382 Northeast Ohio has been a center for commerce and industry since early in the 19th 383 century, when development of the region was accelerated substantially with the 384 establishment of the Ohio and Erie Canal (Cockrell, 1992). Indeed, fish in many of the 385 region’s streams had shown evidence of substantial decline for decades prior to the 386 timeframe of the current study (Trautman, 1981). 387 388 389 Biological Integrity in Urban Streams Walton et al. -18 390 5.2 Are there interactive and scale effects among land use/demographic gradients? 391 Our findings reinforce the notion that the mix and spatial juxtaposition of land 392 uses within an urbanized basin are important determinants of biological integrity of 393 streams (Wang, et al., 2003). For example, the principal components that emerged as 394 most important in explaining variability in IBI in both the sub-catchment (SPC3, 9.1 % of 395 total variance in IBI) and the local zone analyses (LPC8, 6.5 % of total variance in IBI) 396 were components describing a spatial counter-gradient in forested and open, grassland 397 cover (Fig. 2). 398 Further, the nature of land use effects changed profoundly with spatial scale and 399 proximity to the biological sampling point. In particular, the polarity of forest cover 400 effects on biological integrity changed between sub-catchment and local zone scales. 401 Whereas high forest cover within the sub-catchment overall was associated with low IBI 402 value, high forest cover within the local zone was associated with high IBI (Fig. 3, Table 403 7). Since forest cover in the local zone may represent riparian vegetation in large part, 404 the positive effect on biological integrity at this scale is consistent with general findings 405 that riparian vegetation can buffer upland effects (Steedman, 1988; Horner, et al., 1997; 406 May, et al., 1997). On the other hand, the negative impact of forest cover at the sub- 407 catchment scale seems counterintuitive at first glance. 408 However, these findings are interpretable in light of current and historical patterns 409 of land use in northeast Ohio. Many sites characterized as having high forest cover at the 410 sub-catchment scale are also associated with high population and housing density, as well 411 as relatively high urban land use (Fig. 3). This combination of factors characterizes 412 older, inner-ring suburbs in the region. In these neighborhoods, the canopies of large Biological Integrity in Urban Streams Walton et al. -19 413 street trees overhang houses, streets and other impervious surfaces, and wooded parks are 414 interspersed within densely populated residential areas. These suburbs have a long history 415 of urban impact on local streams. By 1900, wealthy industrialists and merchants were 416 leaving an increasingly industrialized city-center of Cleveland to establish new suburban 417 neighborhoods just beyond the city limits (Cigliano, 1991). Out-migration from the city 418 center and from older inner-ring suburbs has continued and, in fact, has accelerated in 419 recent decades (Bier 1993, 2001). Within these older urban/suburban areas, population 420 loss and economic decline are associated with ageing and inadequate waste water 421 management infrastructure (Bier, 2001). 422 In addition, the counter-gradient of forest vs. grasslands at the sub-catchment 423 scale, in combination with the positive effect of local zone forest cover on IBI, defines a 424 landscape signature indicative of high biological integrity for the region. The sites with 425 highest biological integrity in our dataset were those characterized by high open 426 grassland cover and low forest cover at the sub-catchment scale, but high forest cover in 427 the local zone (Fig. 3). This nexus of land cover categories is most often found in areas 428 beyond the urban core where forest cover is associated with riparian strips adjacent to 429 open parkland and/or agricultural fields. Within cities and suburbs, similar landscapes are 430 found in protected and managed areas, including an extensive network of regional parks 431 and the Cuyahoga Valley National Park. 432 433 5.3 Are stream habitat or landscape variables better predictors of biological integrity? 434 Our measure of stream habitat quality in these analyses, the Qualitative Habitat 435 Evaluation Index (QHEI), was designed and calibrated as a measure of the potential for Biological Integrity in Urban Streams Walton et al. -20 436 stream habitat to support healthy, native fish communities (Rankin, 1989). Hence, this 437 variable was expected to covary significantly with IBI and MIWB. Indeed, QHEI was a 438 significant predictor of IBI, MIWB, and a majority of the IBI sub-metrics. In this regard, 439 the current findings are congruent with previous studies demonstrating that fish 440 community health is associated with habitat quality (Schlosser, 1982; Roth, et al., 1996). 441 However, QHEI was a poorer predictor of IBI than were landscape variables 442 overall. For the sub-catchment level analysis, QHEI accounted for 4.8 % of total variation 443 in IBI, whereas the principal components describing landscape and demographic features 444 combined to account for 19.5% of variance in IBI. In the analysis including variables 445 describing the 500 m radius local zone, the landscape components combined to explain 446 27.3% of the variance in IBI, in comparison to 2.6% attributable to QHEI alone. 447 Moreover, several landscape/demographic components explained more variability singly 448 than did QHEI. For example, LPC 8 alone explained 3-fold more variance in IBI (8.6%) 449 than did QHEI. Roth et al. (1996) also reported than habitat quality was no better as a 450 predictor of fish community health than features of land cover. Overall, these finding 451 emphasize the combined importance of both stream channel and conditions within the 452 uplands as determinants of biotic quality of streams (Booth and Jackson, 1997). 453 We also found that QHEI was unrelated to any of the land cover or demographic 454 variables, either alone or in combination as principal components, or when the land cover 455 or demographic variables were aggregated at sub-catchment or local-zone scales. Given 456 the links between landscape features and stream morphology, hydrology, and stream 457 habitat quality that have been documented in a variety of studies (Richards and Host, 458 1994; Roth, et al., 1996; MacRae and DeAndrea, 1999), this finding is noteworthy, but it Biological Integrity in Urban Streams Walton et al. -21 459 is not unique to the current analysis. Wang et al. (1997, 2003) reported little or no 460 correlation between habitat quality variables and effective impervious surface cover 461 among urban streams in Wisconsin. How then is it possible for stream habitat quality to 462 vary independently of land use/land cover and demography, while biological integrity 463 covaries significantly with both stream habitat quality and landscape level variables? 464 We suggest that the resolution of this apparent paradox lies in the interaction of 465 the geological and urbanization histories of the region. In many cases, streams in 466 northeast Ohio lie within ravines, often quite deeply incised, that were formed by the 467 retreat of the last glaciation (White and Totten, 1982). During early settlement of the 468 region, the deeply incised terrain made transportation and communication difficult, 469 isolated settlements, and the steep, unstable hillsides were largely unavailable for 470 building, cultivation or pasture land (Cockrell, 1992). Many of these ravines formed the 471 template for city and suburban parklands, including the Cuyahoga Valley National Park. 472 Hence, these areas preserved natural features precisely because they were not useful for 473 other purposes. Thus, the ravines, and associated parks, have provided some degree 474 protection from the worst effects of urbanization on the physical features of streams. 475 Nevertheless, biological degradation may proceed inexorably through a variety of 476 urbanization effects that degrade biota but have lesser impacts on stream habitat (Allan, 477 2004), including stressors that short-circuit the riparian zone, e.g., sewer outfalls, thermal 478 heat island effects, and atmospheric deposition. Further, stream biodiversity can reflect 479 the impacts of devastating pulse events that may not necessarily have discernable long 480 term effects on physical habitats. One local example is a large fire in a scrap tire yard in 481 1981 that released tens of thousands of liters of petroleum derivatives into the headwaters Biological Integrity in Urban Streams Walton et al. -22 482 of a small stream that was otherwise largely contained within the Cuyahoga National 483 Park (Cockrell, 1992). In these geological and historical contexts, contemporary 484 assessments of stream habitat for this region may provide only limited guidance 485 regarding the potential for streams to support healthy biological communities. 486 487 488 6. Conclusions While our findings are consistent with previous studies indicating that urban land 489 use has a negative association with biological integrity of streams (Klein, 1979; 490 Steedman, 1988; Roth, et al., 1996; Dreher, 1997; May, et al., 1997; Boward, et al., 1999; 491 Yoder, et al., 2000; Morse, et al., 2003; Roy, et al., 2003; Miltner, et al., 2004), this 492 analysis also demonstrates that spatial interactions with other aspects of the urban 493 landscape are important determinants of variability in stream biota. In fact, our results 494 suggest that in regions with long histories of urban development such as northeast Ohio, 495 other axes of landscape variability may emerge as even stronger predictors of variability 496 in biological quality among stream sites. Further, multiple landscape features may have 497 interactive effects on biological integrity which may vary both in magnitude and direction 498 with spatial scale, e.g., forest cover in the current case. 499 Our analysis also emphasizes that the influence of urbanization on streams is 500 shaped by regional geological and historical contexts. Within the Cuyahoga River basin, 501 unstable ravines of glacial origin have impeded agricultural and urban development in 502 some stream reaches, thereby preserving natural features of riparian zones and stream 503 habitats, but not necessarily biological integrity. Rather, in northeastern Ohio, land use Biological Integrity in Urban Streams Walton et al. -23 504 signatures indicative of parklands are better predictors of biological integrity of fish 505 communities than measures of stream habitat quality. 506 507 Acknowledgements 508 We a thank Stuart Schwartz, Director of the Center for Environmental Sciences, 509 Technology, and Policy (CESTP) at Cleveland State University, and his staff for their 510 administrative and data management assistance on this project. We thank Elizabeth 511 Whippo-Cline for her assistance with early stages of the project and Lester Stumpe of the 512 Northeast Ohio Regional Sewer District for his advice and support. The project has also 513 benefited from the work of the following graduate student assistants: Shawn Bleiler, 514 Sonya Steckler and Cari-Ann Hickerson. This project was financed through a grant from 515 the Ohio Environmental Protection Agency and the United States Environmental 516 Protection Agency, under the provisions of Section 319(h) of the U.S. Clean Water Act, 517 and through the U.S.E.P.A. National Risk Management Laboratory. 518 519 References 520 Allan, J.D., 2004. Landscapes and riverscapes: the influence of land use on stream 521 522 523 ecosystems. Annu. Rev. Ecol. Evol. Syst. 35, 257-284. Bier, T., 1993. Cuyahoga County Outmigration. Housing Policy Research Program, Levin College of Urban Affairs, Cleveland State University. 524 Bier, T., 2001. Moving Up, Filtering Down: Metropolitan Housing Dynamics and Public 525 Policy. The Brookings Institution, Center on Urban and Metropolitan Policy. Biological Integrity in Urban Streams 526 Walton et al. -24 Booth, D.B., Jackson, C.R., 1997. Urbanization of aquatic systems: degradation 527 thresholds, stormwater detection, and the limits of mitigation. J. Amer. Wat. 528 Resour. Assoc. 33, 1077-1090. 529 Boward, D., Kayzak, P., Stranko, S., Hurd, M., Prochaska, T., 1999. From the Mountains 530 to the Sea: the State of Maryland's Freshwater Streams. Maryland Department of 531 Natural Resources, Monitoring and Non-tidal Assessment Division. EPA 903-R- 532 99-023. Annapolis, Maryland. 533 534 535 Cigliano, J., 1991. Showplace of America: Cleveland's Euclid Avenue, 1850-1910. The Kent State University Press, Kent, Ohio. Cockrell, R., 1992. A Green Shrouded Miracle: The Administrative History of Cuyahoga 536 Valley National Recreation Area, Ohio. 537 http://www.cr.nps.gov/history/online%5Fbooks/Cuyahoga/. 538 Cohen, J.C., 2003. Human population: the next half century. 303, 1172-1175. 539 Coles, J.F., Cuffney, T.F., McMahon, G., Beaulieu, K.M., 2004. The effects of 540 urbanization on the biological, physical, and chemical characterisitcs of coastal 541 New England streams. U.S. Geological Survey Professional Paper 1695. 542 Dreher, D.W., 1997. 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Maryland stream biological survey: a state agency program to assess the impact of Biological Integrity in Urban Streams Walton et al. -26 571 anthropogenic stress on stream habitat quality and biota. Environ. Monitor. Assess. 572 51, 299-316. 573 574 575 Klein, R., 1979. Urbanization and stream quality impairment. Water Resour. Bull. 15, 948-963. MacRae, C., DeAndrea, M., 1999. Assessing the impact of urbanization on channel 576 morphology. 2nd International Conference on Natural Channel Systems. Niagara 577 Falls, Ontario. 578 May, C.R., Horner, J., Karr, B., Mar, B.W., Welch, E., 1997. Effects of urbanization on 579 small streams in the Puget Sound lowland ecoregion. Water. Protect. Tech. 2, 483- 580 494. 581 582 583 584 585 Miltner, R., White, D., Yoder, C.O., 2004. The biotic integrity of streams in urban and suburbanizing landscapes. Landscape Urban Plann. 69, 87-100. Morley, S.A., Karr, J.R., 2002. Assessing and restoring the health of urban streams in the Puget Sound basin. 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Ohio EPA, Division of Water Quality Planning and 597 Assessment, Ecological Assessment Section. Columbus, Ohio. 598 Rankin, E.T., 1995. The use of habitat assessments in water resource management 599 programs, in: Davis, W., Simon, T. (Eds.), Biological Assessment and Criteria: 600 Tools for Water Resource Planning and Decision Making. Lewis Publishers, Boca 601 Raton, Florida, pp. 181-208. 602 Richards, C., Host, G., 1994. Examining land use influences on stream habitats and 603 macroinvertebrates: a GIS approach. Water Resour. Bull. 30, 729-738. 604 Roth, N.E., Allan, J.D., Erickson, D.L., 1996. Landscape influences on stream biotic 605 integrity assessed at multiple spatial scales. Landscape Ecol. 11, 141-156. 606 Roy, A.H., Rosemond, A.D., Paul, M.J., Leigh, D.S., Wallace, J.B., 2003. Stream 607 macroinvertebrate response to catchment urbanisation (Georgia, U.S.A.). Freshwat. 608 Biol. 48, 329-346. 609 Schaal, G.M., Motsch, B.R., 1997. State of Ohio land cover, 1994. Database derived from 610 Landsat TM. Ohio Department of Natural Resources, Division of Real Estate and 611 Land Management, GIS and Remote Sensing Services, Columbus, OH. 612 613 614 615 Schlosser, I.J., 1982. Fish community structure and function along 2 habitat gradients in a headwater stream. Ecol. Monogr. 52, 395-414. Schuler, T.R., 1994. The importance of imperviousness. Watershed Protect. Tech. 1, 100111. Biological Integrity in Urban Streams 616 617 618 619 620 621 622 623 624 625 626 627 628 Walton et al. -28 Steedman, R.J., 1988. Modification and assessment of an index of biotic integrity to quantify stream quality in southern Ontario. Can. J. Fish. Aquat. Sci. 45, 492-500. Trautman, M.B., 1981. The Fishes of Ohio. Ohio State University Press, Columbus, Ohio. Wang, L., Lyons, J.L., Kanehl, P., 2003. Impacts of urbanization on stream habitat and fish across multiple scales. Environ. Manage. 28, 255-266. Wang, L., Lyons, J.L., Kanehl, P., Gatti, R., 1997. Influences of watershed land use on habitat quality and biotic integrity in Wisconsin streams. 22, 6-12. Weaver, L.A., Garman, G.C., 1994. Urbanization of a watershed and historical changes in a stream fish assemblage. Trans. Amer. Fish. Soc. 123, 162-172. White, G.W., Totten, S.M., 1982. Glacial Geology of Northeastern Ohio. Bulletin 68. Ohio Division of Natural Resources, Columbus, OH. Yoder, C.O., Rankin, E.T., 1995. The role of biological criteria in water quality 629 monitoring, assessment, and regulation. Ohio EPA Technical Report. Ohio EPA, 630 Monitoring and Assessment Section, Division of Surface Water. MAS/1995-1-3. 631 Columbus, Ohio. 632 Yoder, C.O., Miltner, R., White, D., 1999. Assessing the status of aquatic life designated 633 uses in urban and suburban watersheds, in: Kirschner, R. (Eds.), National 634 Conference on Retrofit Opportunities for Water Resource Protection in Urban 635 Environments. pp. 16-28. 636 Yoder, C.O., Miltner, R., White, D., 2000. Using biological criteria to assess and classify 637 urban streams and develop improved landscape indicators. National Conference on 638 Tools for Urban Water Resource Management and Protection. Chicago, Illinois. Biological Integrity in Urban Streams Walton et al. -29 639 Table 1. List and summary statistics for variables used in the current analysis. Also listed are shortened variable names used in subsequent tables. Sub-Catchment Scale 500 m Radius Local Zone Abbreviated Variable N Mean S.E.M. Min - Max Mean S.E.M. Min - Max Name Biological/Habitat Variables IBI 227 12.00 – 50.00 29.48 .6 MIWB 227 0 – 9.02 5.54 .12 IBI Sub-metrics 227 1. Number of native species 0 – 24 10.89 .29 2. Number of darter species 0–6 1.30 .10 3. Number of headwater 0–6 .97 .06 species 4. Number of cyprinid 0–9 4.04 .14 species 5. Number of sensitive 0–3 .23 .04 species 6. % of tolerant species 0 – 100 59.70 1.76 7. % omnivores 0 – 100 26.55 1.37 8. % insectivores 0 – 92.31 27.21 1.55 9. % pioneer species 0 – 100 33.04 1.48 10. Number of individuals 39 - 6523 731.89 56.86 11. % simple lithophiles 0 – 82.98 30.76 1.27 12. % of individuals with deformities, eroded fins, 0 – 15.43 1.00 .14 lesions, or tumors QHEI 165 61.66 .89 25.00 – 86.50 Land Use/Demographic Variables % Urban Urban 227 16.12 1.51 0-100 0 – 99.23 18.42 1.60 % Open Grassland / Parkland / Agricultural Grass 227 37.75 1.91 0-100 0 – 94.45 23.84 1.42 Fields % Shrub / Scrub Shrub 227 8.73 1.08 0-86.13 0 – 28.33 3.67 .35 % Non-forested Wetland Wetland 227 0-100 0 – 58.15 9.00 .96 4.93 .52 % Open Water Water 227 0-18.81 0 – 10.16 .43 .12 .11 .05 % Forested Forest 227 27.87 1.95 0-100 0 – 100 43.98 1.69 % Barren Barren 227 0-3.16 0 – 6.06 .07 .02 .05 .03 Population Density (per 5130.20 Pop 227 0 – 133772.57 326.73 29.14 0 – 2320.92 km2) 1157.28 2329.72 Housing Density (per km2) House 227 0 – 71217.05 39.92 4.31 0 - 378 544.61 Catchment Area (km2) Area 227 5.06 1.12 .10 – 84.50 Distance from Terminus Distance 227 0-46.67 7.97 .69 (km) Biological Integrity in Urban Streams Walton et al. -30 Table 2. Pearson product moment correlations for variables aggregated at sub-catchment scale. * .05>P>.01, **.01>P>.001, *** P<.001 IBI MIWB QHEI Barren Grass Shrub Urban Water Wetland Forest Pop House Distance MIWB .570*** QHEI .161* .153* Barren .093 .078 .015 Grass .312*** .069 -.026 .049 Shrub -.016 .086 -.044 .007 -.227** Urban -.153** -.037 .046 .063 -.329*** -.059 Water .263*** .119* -.034 .135* .123* -.097 -.115* Wetland .156** .221** .035 -.074 -.208** .329*** -.117* .092 Forest -.180** -.138* .041 .084 -.384*** -.351*** -.286*** .024 -.322*** Pop -.152** .087 -.021 .055 -.283*** -.203** .308*** -.035 -.139* .281*** House -.064 .046 -.025 -.107 -.235*** -.209** .404*** -.087 -.185** -.034 .315*** Distance -.338*** -.034 -.086 -.053 -.173** .213** -.028 -.012 .062 -.010 .072 -.097 Area .340*** .162** .040 .422*** .067 -.113* -.082 .396*** -.047 .355*** .101 .816*** -.158** Biological Integrity in Urban Streams Walton et al. -31 Table 3. Multiple linear regressions relating common measures of “urbanization” (% urban land use, housing density, population density) to IBI. Regressions are presented both with and without the spatial co-factors, distance from terminus and catchment area. Regression Statistics Variable Coefficient Std. Error P-value R2 Regressions excluding distance from terminus and catchment area Y-intercept 1.086 .141 <.001 .055 % Urban Land Use -.069 .031 .026 QHEI .238 .107 .028 Y-intercept Housing Density QHEI 1.024 -.022 .294 .235 .017 .129 <.001 .213 .024 .068 Y-intercept 1.181 .193 <.001 Population Density .039 .014 .005 QHEI .220 .014 .040 Regressions including distance from terminus and catchment area Y-intercept 1.028 .187 <.001 % Urban Land Use -.049 .030 .102 QHEI .202 .101 .046 Distance from Terminus -.074 .020 <.001 Catchment Area .026 .008 .001 .073 Y-intercept Housing Density QHEI Distance from Terminus Catchment Area .923 -.016 .279 -.084 .027 .224 .016 .119 .024 .010 <.001 .321 .022 .001 .005 .225 Y-intercept Population Density QHEI Distance from Terminus Catchment Area 1.085 -.041 .191 -.066 .032 .184 .013 .098 .020 .003 <.001 .002 .002 .001 <.001 .219 .184 Biological Integrity in Urban Streams Walton et al. -32 Table 4. Factor loadings, % of total variance, and correlation with QHEI and IBI for principal components obtained from sub-catchment scale analysis. Only loadings that were significant at P < 0.01 are shown. Pearson product-moment correlations with IBI and QHEI are shown with P-values in parentheses. Principal Components SPC1 SPC2 SPC3 SPC4 Original Variables Barren .616 ----.142 Grass/Parkland ---.292 -.766 -.320 Shrub/Scrub --.787 ----Urban ------.938 Water .597 .132 -.151 -.234 Wetland --.757 ----Forest .185 -.458 .706 -.381 Population Density .312 -.283 .558 .450 Housing Density .777 -.350 .404 .144 Catchment Area .894 -.132 .141 --Distance from Terminus -.178 .365 .414 --% of Total Variance Correlation with QHEI Correlation with IBI 26.7 16.1 13.5 11.2 .024 (.761) .353 (<.001) -.030 (.700) .035 (.595) .001 (.991) -.369 (<.001) .034 (.666) .112 (.093) Biological Integrity in Urban Streams Walton et al. -33 Table 5. Multiple regression results at sub-catchment scale relating IBI, MIWB, and sub-metrics of IBI to principal components describing land use/land cover and demographic data, QHEI and Julian day. Coefficients shown with P-values in parentheses; variables excluded by stepwise selection procedure are indicated by ns. Dependent Variables Regression Coefficients for Independent Variables SPC SPC SPC SPC Multimetric Indices R2 Y-intercept QHEI Julian Day 1 2 3 4 -59.528 .034 -.039 -.018 .322 11.666 IBI .39 ns (<.001) (<.001) (<.001) (.030) (<.001) (<.001) -57.996 .034 .033 .382 11.218 MIWB .17 ns ns (.002) (.005) (.010) (.006) (.002) IBI Sub-Metrics .015 .048 .051 -.035 .573 1. Number of native species .22 ns ns (.957) (.001) (.001) (.017) (<.001) -57.535 .051 .063 -.061 1.050 10.800 2. Number of darter species .30 ns (.035) (.003) (.001) (.001) (<.001) (.040) -.525 -.065 .434 3. Number of headwater species .13 ns ns ns ns (.079) (<.001) (.010) -.546 -.049 .671 4. Number of cyprinid species .13 ns ns ns ns (.085) (.003) (<.001) -.800 -.049 .488 5. Number of sensitive species .17 ns ns ns ns (.001) (<.001) (<.001) 105.988 -20.292 6. % tolerant species .05 ns ns ns ns ns (.004) (.004) 110.476 .050 -21.239 7. % omnivores .12 ns ns ns ns (<.001) (.010) (<.001) .499 .061 .096 -.051 8. % insectivores .18 ns ns ns (<.001) (.004) (<.001) (.024) 1.439 -.486 9. % pioneer species .03 ns ns ns ns ns (.001) (.044) 2.632 .121 -.121 10. Number of individuals .12 ns ns ns ns (<.001) (.001) (.002) -.223 -.053 .434 11. % simple lithophiles .07 ns ns ns ns (.552) (.008) (.040) 12. % of individuals with deformities, eroded 23.080 .018 -4.445 .12 ns ns ns ns fins, lesions, and tumors (.005) (.002) (.005) Biological Integrity in Urban Streams Walton et al. -34 Table 6. Factor loadings, % of total variance, and correlation with QHEI and IBI for principal components obtained from analysis incorporating variables aggregated at 500 m radius local zone scale. Only loadings that were significant at P < 0.01 are shown. Pearson product-moment correlations with IBI and QHEI are shown with P-values in parentheses. Principal Components Original Variables LPC1 LPC2 LPC3 LPC4 LPC5 LPC6 LPC7 LPC8 Sub-Catchment Scale Barren --.361 ------.783 .142 --Grass/Parkland -.328 .205 -.362 -----.173 .177 -.706 Shrub/Scrub ---.159 .235 .841 --------Urban .549 -.325 .152 ----.403 -.209 -.212 Water --.316 .213 -.141 ----.643 -.157 Wetland ----.852 .148 --------Forest --.472 -.241 -.192 ------.783 Housing Density .550 .729 ------------Population Density .896 .209 ----------.144 Catchment Area --.920 ------------Distance from Terminus ---.228 -.145 .553 .194 --.378 .268 Local Scale Barren @ 500 m ----------.795 ----Grass/Parkland @ 500 m .426 .213 --.156 .680 .190 -.157 -.211 Shrub/Scrub @ 500 m --.202 .238 .816 --------Urban @ 500 m .544 -.222 -.430 -.267 .392 ----.230 Water @ 500 m --.316 --------.808 --Wetland @ 500 m --.205 .786 .213 ----.251 --Forest @ 500 m ---------.953 -----.147 Housing Density @ 500 m .871 .247 -.150 ----------Population Density @ 500 m .912 --------------% of Total Variance 21.7 14.6 11.2 9.3 6.6 6.2 5.4 4.8 Correlation with QHEI -.019 (.807) -.012 (.875) .073 (.350) -.122 (.120) -.106 (.178) .067 (.394) -.005 (.953) .056 (.479) Correlation with IBI -.195(.003) .290(<.001) .216(.001) -.132(.048) -.265(<.001) .036(.592) .031(.639) -.319(<.001) Biological Integrity in Urban Streams Walton et al. -35 Table 7. Multiple regression results relating IBI, MIWB, and sub-metrics of IBI to principal components incorporating local zone scale aggregation. Coefficients shown with P-values in parentheses; variables excluded by stepwise selection procedure are indicated by ns. Dependent Variables Regression Coefficients for Independent Variables YLPC LPC LPC LPC LPC LPC Julian 2 Multimetric Indices R LPC 6 LPC 7 QHEI intercept 1 2 3 4 5 8 Day -47.625 -.025 .031 .027 -.029 -.038 .232 9.399 IBI .44 ns ns ns (<.001) (.004) (<.001) (.001) (<.001) (<.001) (.007) (<.001) -46.357 .027 .037 -.025 .311 8.995 MIWB .17 ns ns ns ns ns (.014) (.026) (.004) (.042) (.023) (.013) IBI Sub-Metrics .131 .032 .047 -.037 -.041 .509 1. Number of native species .21 ns ns ns ns ns (.638) (.024) (.001) (.008) (.001) (.001) -1.215 -.058 .043 .068 -.062 -.072 .838 2. Number of darter species .35 ns ns ns ns (.001) (<.001) (.013) (<.001) (<.001) (<.001) (<.001) 3. Number of headwater -.344 -.072 -.045 -.062 .338 .29 ns ns ns ns ns ns species (.212) (<.001) (.001) (<.001) (.030) -.423 -.043 -.048 .605 4. Number of cyprinid species .17 ns ns ns ns ns ns ns (.166) (.007) (.002) (.001) -.673 -.025 -.029 -.042 -.036 .417 5. Number of sensitive species .24 ns ns ns ns ns (.003) (.037) (.016) (<.001) (.001) (.001) 88.917 -.082 -16.998 6. % tolerant species .11 ns ns ns ns ns ns ns ns (.014) (.001) (.015) 89.532 -.040 .066 -17.196 7. % omnivores .17 ns ns ns ns ns ns ns (.001) (.027) (.001) (.001) .503 .045 .093 .062 .050 8. % insectivores .19 ns ns ns ns ns ns (<.001) (.034) (<.001) (.006) (.012) 1.513 .067 -.526 9. % pioneer species .09 ns ns ns ns ns ns ns ns (<.001) (.003) (.028) 2.647 .124 -.075 -.130 10. Number of individuals .16 ns ns ns ns ns ns ns (<.001) (.001) (.037) (.001) .534 -.065 -.036 -.042 11. % simple lithophiles .12 ns ns ns ns ns ns ns (<.001) (.001) (.035) (.020) 12. % of individuals with 18.054 .016 -.012 .016 -3.475 deformities, eroded fins, .16 ns ns ns ns ns ns (.025) (.007) (.026) (.004) (.025) lesions, and tumors Biological Integrity in Urban Streams Walton et al. -36 Figure Captions Fig. 1. A. Map of Cuyahoga River basin showing locations of IBI sample points. B. (Insert) Example of IBI sample point catchment and 500 m local zone delineations. Land cover categories are shown for local zones. Fig. 2. Percent forest cover (closed circles) and percent grassland cover (open circles), arcsin-square root transformed, as a function the third principal component at sample point sub-catchment scale (SPC3). Fig. 3. Characteristic land use and population densities associated with quartiles of IBI. Top panels A and B illustrate land use categories distributed among IBI quartiles at subcatchment and local zone scales, respectively. Bottom panels illustrate population densities associated with IBI quartiles at (C) sub-catchment and (D) local zone scales. IBI quartiles are arranged from lowest (quartile 1) through highest IBI score (quartile 4). Bars show means ± 1 standard error of the mean. Fig. 4. Plot of observed IBI as a function of predicted IBI values derived from multiple regression relating stream habitat quality (QHEI) and principal components combining land use and demographic variables to IBI. Symbols denote pentile groupings of residuals derived from the multiple regression. Stippled region indicates boundary region for warm water habitat (WWH) use attainment. Sites to the right of arrow labeled “max” all show IBIs sufficient for WWH attainment; sites to the left of “min” arrow failed to exceed the WWH boundary. Biological Integrity in Urban Streams Walton et al. -37 A. Lake Erie LAKE LAKE GEAUGA GEAUGA Cleveland Cleveland Cuyahoga Cuyahoga River River Basin Basin CUYAHOGA CUYAHOGA MEDINA MEDINA PORTAGE PORTAGE Akron Akron Legend IBI Sample Stream Cuyahoga Watershed 0 SUMMIT SUMMIT 5 10 kilometers County B. 0 500 1,000 meters Legend Land Use IBI Sample Stream IBI Catchment 500 Meter Local Zone Urban Agriculture/ Open Urban Area Shrub/ Scrub Wooded Open Water Non Forested Wetlands Biological Integrity in Urban Streams Walton et al. -38 Biological Integrity in Urban Streams Walton et al. -39 Biological Integrity in Urban Streams Walton et al. -40 min 60 max 54 OBSERVED IBI 48 42 36 30 24 18 18 24 30 36 42 48 PREDICTED IBI 54 60