AN ABSTRACT OF THE THESIS OF Michelle L. LaRue for the degree of Master of Science in Fisheries Science presented on July 24, 2001. Title: Characterization of Stream Fish Assemblages and Land Use Associations within a Southern Ohio National Forest. Abstract approved: Redacted for Privacy Li Redacted for Privacy Hiram W. Li Seasonally, in 1998 and 1999, I examined spatial and temporal variation in fish assemblages of agricultural, forested, and acid mine drainage tributaries within the Wayne National Forest (WNF) in southern Ohio. Land use and natural disturbance explained patterns in stream fish assemblages. Creek chub and green sunfish were present in all land use types. Generally, with the exception of creek chub and green sunfish, species most abundant in one land use type occurred infrequently in the other land use types sampled. For example, redbelly dace and blacknose dace dominated forested assemblages but rarely occurred in mining or agricultural assemblages. Agricultural sites consisted of higher order streams, located at lower elevations with reduced canopy cover. Forested sites included intermittent streams associated with higher elevation, low stream order, and high canopy cover. Acidic conditions characterized mining sites, which otherwise remained physically similar to forested sites. Stream order, elevation, and canopy cover explained the majority of the variance in assemblage structure within 1998, and pH was also important. In 1999, water quality, specifically dissolved oxygen, and seasonal variation became important. Assemblages changed following drought in 1999. Forested assemblages remained most similar following drought, while agricultural assemblages exhibited less similarity (i.e., greater variability). These results suggest that large-scale reach characteristics and chemical signals related to land use are important to fish assemblage structure, but in times of environmental fluctuation, water chemistry of other site-specific variables may be of even greater importance due to physiological tolerances and limitations of fishes. Characterization of Stream Fish Assemblages and Land Use Associations within a Southern Ohio National Forest by Michelle L. LaRue A Thesis Submitted to Oregon State University In Partial Fulfillment of the requirements for the degree of Master of Science Presented July 24, 2001 Commencement June 2002 Master of Science thesis of Michelle L. LaRue presented on July 24. 2001. APPROVED: Redacted for Privacy , reprèienting Fisheries Science Redacted for Privacy Professor, representing Fisheries Science Redacted for Privacy Head of Department and Wildlife Redacted for Privacy Dean of Grâduafe School I understand that my thesis will become part of the permanent collection of Oregon State University libraries. My signature below authorizes release of my thesis to any reader upon re est. / Redacted for Privacy Michelle L. LaRue, Author ACKNOWLEDGEMENTS I offer my most sincere gratitude to my mentor, Deanna Stouder, for long term encouragement and friendship. I would like to thank Judy Li and Hiram Li for their wonderful assistance and for serving as my advisors at Oregon State University. I also thank Cliff Pereira for his role on my committee. I thank the staff at the Wayne National Forest for their friendship, support, and expertise, especially Rebecca Ewing and Ralph Miller. Field assistants are essential to any project: I thank Josh Harper for his help and patience, and for his creative field accessories. I also thank A. Butterworth for her assistance. In addition, I'd like to recognize D. Stouder, R. Miller, S. Micucci, and R. Miller for being cheerful volunteers. I thank T. Cavendar at the Ohio Museum of Biological Diversity, Ohio State University for his help with specimen identification. I also thank M. White, F. McCormick, R. Ewing, and J. Grow for knowledgeable conversations regarding the local geography, fauna, and histomy. Funding was provided by the Ohio Cooperative Fish and Wildlife Research Unit and the Wayne National Forest (CCS-09- 14-98-03), and from the USDA Forest Service, Pacific Northwest Research Station, Aquatic and Land Interaction Program. I'd also like to thank Hecla Water Association, Oregon Cooperative Fish and Wildlife Research Unit, and USGS for field equipment loans. There are many individuals who have served as mentors and friends, providing me with confidence in my blossoming career; thank you so much to all of you who have ever had, and continue to have, faith in me. Much love to my family who all believe I am crazy, but believe nonetheless. TABLE OF CONTENTS Page INTRODUCTION ............................................................................... 1 STUDYAREA ..................................................................................4 METHODS ......................................................................................9 SiteSelection ........................................................................... 9 Habitat Characteristics ................................................................ 14 Fishes .................................................................................. 16 DataAnalysis .......................................................................... 16 RESULTS ......................................................................................20 1998 and Land Use Patterns ........................................................ 20 Habitat Characteristics .......................................................... 20 Assemblage Characteristics .................................................... 25 Integrating Assemblage Composition with Habitat Characteristics...... 29 1999 and Examination of Drought .................................................. 31 Habitat Characteristics .......................................................... 31 Assemblage Characteristics .................................................... 36 Integrating Assemblage Composition with Habitat Characteristics...... 37 Response of Fish Assemblages Following Drought ..................... 41 Spatial and Temporal Variation in Fish Assemblages..............................45 DISCUSSION ................................................................................... 52 BIBLIOGRAPHY .............................................................................. 60 APPENDIX ...................................................................................... 68 LIST OF FIGURES Figure Page Schematic map of the state of Ohio highlighting the location of the Wayne National Forest ................................................................. 5 2 Monthly precipitation received in south central Ohio during 1998and 1999 ........................................................................... 7 3 Schematic of a typical stream reach (50m following thaiweg), depicting the start and end of the study site ................................................... 15 4 Comparison of means for stream temperature (a), dissolved oxygen (b), pH (c), conductivity (d), turbidity (e), and canopy (1) for 1998 and 1999 bylanduseandseason ................................................................ 21 5 Substrate composition for 1998 (a) and 1999 (b) by land use and season. . . .23 6 Temporal patterns in number of species captured for all sites .................. 27 7 Temporal patterns in catch per unit effort (CPUE) for all sites ................. 28 8 NMS ordination of 1998 fish assemblages. Overlaid on the ordinations are joint plots of environmental variables (a) and individual fish species (b) strongly correlated (r2> 0.5) with the ordination axes ............... 30 9 NMS ordination of 1999 fish assemblages. Overlaid on the ordinations are joint plots of environmental variables strongly correlated (r2 > 0.5) with ordination axes one and two (a) and one and three (b)..................... 38 10 NMS ordination of 1999 fish assemblages. Overlaid on the ordinations are joint plots of individual fish species strongly correlated (r2 > 0.5) with ordination axes one and two (a), and one and three (b) .................... 39 11 Composition of fish assemblages following drought in autumn 1999 .......... 42 12 HIerarchical agglomerative cluster analysis of autumn 1998 and 1999 fish assemblages for agricultural and forested sites ............................... 44 LIST OF FIGURES (Continued) Figure Page 13 Comparison of autumn 1998 and autumn 1999 fish assemblages in forested and agricultural sites for number of species captured (a) and catch per unit effort (CPUE)(b) .....................................................46 14 NMS ordination, based on fish assemblage data, projected onto axes one and two (a), and onto axes one and three (b), for 1998 (filled symbols) and 1999 (empty symbols) .................................... 50 LIST OF TABLES Table Page 1 Sampling locations and landscape characteristics within Wayne National Forest, Ironton District, Ohio ............................................. 10 2 Seasonal samples collected at each site ........................................... 13 3 Percent total catch of the most common fishes ( 1% of total) in 1998 and 1999 by land use ..................................... 26 4 Correlations between axes coordinates from 1998 NMS ordination and species abundance and physical parameters .................................. 32 5 Percent catch of all fishes captured annually and for entire study ............. 33 6 Correlations between axes coordinates from 1999 NMS ordination and species abundance and physical parameters.................................. 40 7 Morisita's Index values for assemblage comparisons between autumn 1998 and autumn 1999 within a site (a) and for autumn 1998 and autumn 1999 assemblage comparisons within agricultural and forested land uses (b) .................................................................. 43 8 Composition of autumn 1998 and autumn 1999 fish assemblages in forested and agricultural sites ................................................... 47 9 Correlations between axes coordinates from 1998 and 1999 NMS ordination and species abundance and physical parameters .................... 51 CHARACTERIZATION OF STREAM FISH ASSEMBLAGES AND LAND USE ASSOCIATIONS WIT}IIN A SOUTHERN OHIO NATIONAL FOREST INTRODUCTION Fish assemblages may reflect patterns of land use and natural disturbances, thus it is important to understand which environmental characteristics, from landscape level to microhabitat, are strongly associated with assemblage structure. Streams are hierarchically organized systems consisting of multiple spatiotemporal scales (Frissell et al. 1986) with physical and biological connections linking streams to their landscapes (THynes 1975, Gregory et al. 1991, Stanford and Ward 1992, Montgomery and Buffington 1999); therefore landscape alterations potentially influence streams at different spatial scales. Headwater streams are particularly sensitive to, and often the most directly impacted by, landscape alteration Karr and Schlosser 1978, Beschta and Platts 1986). Low order streams are also important because they comprise the greatest proportion of streams in a watershed. Natural disturbances (e.g., drought or floods) vary in duration, frequency, and timing, so impacts may also be hierarchically scaled. Land use activities may alter stream fish communities (Schlosser 1991). For example, changes in fish assemblages have been documented with increased agricultural activity (Smith 1971, Trautman 1981, Menzel et al. 1984) and water quality degradation (TKarr 1991). In North America, 33% to 75% of aquatic 2 organisms are rare or extinct (Master 1990, Naiman et al. 1995), including 34% of fish species (Doppelt 1993); habitat alteration is a contributing cause in 73% of fish species extinction (Miller et al. 1989). Natural disturbances may also influence fish assemblages by temporarily altering stream environments. The primary objective of this study was to describe patterns in fish assemblage composition within forested, agricultural, and acid mine drainage tributaries of the Wayne National Forest in south central Ohio. Ohio has a legacy of landscape alteration and modification, including the extraction of natural resources. The state was once 95% forested, but much of the land has been converted for agricultural and residential purposes. The first land purchase to create a national forest in Ohio was in 1934, but the Wayne National Forest (WNF) did not become official until 1951. Stewardship and management of watersheds are priorities of national forests and a current goal of WNF is the development of an ecological aquatic classification system and determination of ecological potential of the aquatic resources. The land within WNF is a fragmented landscape, containing multiple land uses, and both private and public lands. Current land use practices include timber harvest, agriculture, oil and gas development, and commercial coal, clay and limestone mining. While the USDA Forest Service is not currently using timber harvest as a management tool on WNF lands, it maintains an active recreation program which includes hiking, mountain biking, off road vehicles (ORV), and horse trails (R. Ewing, USDA Forest Service, personal communication). Agricultural land use in the watersheds is limited to 3 private lands where channel modification is a common practice. Pastures, hay fields and row crops generally occur along mainstem reaches and higher order tributaries, although some pasture land is found on ridges and sideslopes where the first order streams originate. Intennittent and first order reaches primarily occur in forested areas, while mining areas exist throughout the watersheds. I used descriptive patterns in fish assemblage composition within forested, agricultural, and acid mine drainage tributaries of the Wayne National Forest to ask the following questions: 1) Do fish assemblages vary over time? 2) Are there associations between fish assemblage composition and physical habitat variables? 3) Do fish assemblages within different land use types respond differentially to drought? and 4) Based on land use and temporal variability, what recommendations can be made regarding classification and restoration of these tributary streams? 4 STUDY AREA The Wayne National Forest (WNF) covers over 68,394 ha in unglaciated south central Ohio within the Western Allegheny Plateau physiographic region (Figure 1). This region is unglaciated and hilly (Hammond 1964) and consists mainly of forested patches interspersed with agricultural land (Anderson 1970) with very few urbanized areas. As a consequence of the hilly terrain, agricultural lands are found primarily in the valley lowlands. When nomadic native Indians inhabited the area the land was relatively undisturbed. Later, native people established seasonal camps and villages, and cultivated river valleys and terraces. In the late 1700's colonists came to Ohio and began to settle along the Ohio River and other major waterways; the hills of WNF were some of the last areas to be settled in Ohio. Land was initially cleared for agricultural purposes. Commercial lumbering peaked in 1849; less than 100 years later most marketable trees had been harvested. Southern Ohio was also an active producer of oil, coal, and iron ore. The furnaces used to smelt iron ore required vast amounts of timber for charcoal fuel, and by the early 1900's, most furnaces became non-functional, partly due to the lack of wood. Given these many landscape changes, few undisturbed streams or areas exist in the region today. This study was conducted in two of the major drainage basins located within the WNF, Symmes Creek and Pine Creek; both are tributaries to the Ohio River. Symmes Creek has a drainage area of 921.3 km2 (Krolczyk 1954) and is A Forested sites added in 1999 Figure 1. Schematic map of the state of Ohio highlighting the location of the Wayne National Forest (three districts in stippled locations). Locations of sample sites within the Wayne National Forest are indicated. located in Jackson, Gallia, and Lawrence counties. The mean gradient is low at 0.6 rn/km (Krolczyk 1954). Previous surveys in Symmes Creek (Holeski etal. 1992, 1993 and 1995) reported approximately 44 different fish species including the eastern sand darter (Ammocryptapellucida), a federally monitored species in Ohio. Pine Creek is located west of Symmes Creek and drains 478.4 km2 with an average gradient of 1.6 ni/km (Krolczyk 1954). The watershed is located in Scioto and Lawrence counties and flows through the western edge of the WNF. In the late 1970's 21 species of fish were collected in the Pine Creek basin (Barnes and Canine 1978). South central Ohio normally experiences the most precipitation in spiing and early summer followed by a dry autumn. In 1998, the average precipitation for Ohio was 102.44 centimeters (107% of normal; 7.Olcentimeters above normal) and precipitation in south central Ohio was also above normal (107%) with an average of 112.57 centimeters (7.70 centimeters above normal) (Ohio Department of Natural Resources (ODNR) 1998). However, in 1999 precipitation was consistently below normal for most of the year (Figure 2) and 1999 was ranked as the 1 8 driest year in 117 years of record (ODNR 1999). Overall precipitation during the 1999 calendar year was below normal for all of Ohio (89% of nonnal with average precipitation of 85.06 centimeters; 10.41 centimeters below normal) and for south central Ohio (87% of normal with average precipitation of 90.73 centimeters; 14.15 centimeters below normal). The Palmer Drought Severity Index rated precipitation levels in south central Ohio during 1998 (+7.70 centimeters) as a 7 25 -. 20 1998 Regional Average Departure from Normal * 15 10 E 5 0 -5 25 1999 Regional Average 20 Departure * from Normal 15 10 U 5 0 -5 -10 Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dcc Figure 2. Monthly precipitation received in south central Ohio during 1998 and 1999. *Depflje from normal is based on 50 year record (1931-198 1). 8 vely moist spell; the region was rated as an extreme drought in 1999 centimeters) (ODNR In April 1999, 1998 and (-14.15 1999). south central Ohio began experiencing mild drought conditions (ODNR 1999); flows were considered below normal for this region, but instream water volumes were not noticeably affected. Severe drought conditions were reached by June 1999, and in July 1999 stream flows were deficient in the southern half of Ohio. Sixty-six counties were declared agricultural disaster areas including Gallia, Jackson, Lawrence, and Scioto counties (ODNR 1999). Severe drought conditions continued into autumn 1999. The south central region had greater than normal precipitation for August 1999 (Figure 2) largely due to showers and thunderstorms occurring on August 24-25 (ODNR 1999). After September 1999, south central Ohio received above normal precipitation for the remainder of the year. METHODS Site Selection Six tributaries in the Symmes Creek basin and six tributaries in the Pine Creek basin within the Ironton district of the Wayne National Forest were sampled seasonally (i.e., spring, summer, autumn) in 1998 and 1999 to assess fish assemblage structure (Figure 1, Table 1). Four sites (three in Symmes Creek and one in Pine Creek) were added in spring 1999 to include larger order forested sites for direct comparison among land use sites of similar stream order. Study reaches varied in stream order and were chosen to represent three common land uses of the region; forested areas, agricultural croplands, and acid mine drainage (Table 1). Land use categories were determined from a land cover inventory map for the state (ODNR 1994) and verified in the field. Forested and agricultural reaches were selected so that land use upstream and in adjacent areas was consistent with land use at the study reach. Acid mine drainage streams were directly influenced by point-source pollution from abandoned coal mines. Acid mine drainage occurs when sulfur and suffitic materials are oxidized by air, water, and bacteria into sulfuric acid and ferrous sulfate (Warner 1971), and under neutral pH conditions, ferric hydroxide will precipate onto the stream bottom (Letterman and Mitsch 1978). Sampling reaches were chosen to be representative of each land use category and located away from bridges, major tributaries, agricultural drains, pastures, and feed lots. Habitat information and fish were sampled concurrently 10 Table 1. Sampling locations and landscape characteristics within Wayne National Forest, Tronton Distict, Ohio. Land use codes: A = Agricultural, F = Forested, M = Mining. A indicates sites where cross-section data were not collected for Rosgen Stream Classification. * indicates sites added in 1999. M.a.s.l = meters above sea level. Table 1. LATTITUDE & A DRAINAGE BASIN Pine COUNTY Scioto Turkeyfoot Run A Pine Scioto Camp Creek A Symmes Gallia Long Creek A Symmes Lawrence Cooney Branch F Pine Scioto Howard Run* F Pine Lawrence Wolcott Hollow F Pine Lawrence Asbuxy Creek F Symmes Lawrence Bakers Fork* F Symmes Lawrence Jenkins Ridge F Symmes Gallia Meyers Hollow F Symnies Gallia STREAM NAME Sugarcamp Hollow LAND USE RockyFork F Symmes Gallia Brushy Fork M Pine Lawrence Negro Creek M Pine Lawrence Sand Fork M Symmes Lawrence Webster Creek M Symmes Lawrence LONGfl1Th)E 38°48.07N 82°42.21W 38°43.83N 82°44.58W 38°45.59N 82°27.49W 38°39.26N 82°28. lOW 38°41.57N 82°44.86W 38°44.09N 82°41 .72W 38°42.57N 82°42.04W 38°44.60N 82°34.40W 38°44.02N 82°31.32W 38°43.3 iN 82°26.45W 38°4 1 .05N 82°24.41W 38°48.96N 82°26.59W 38°46.88N 82°38.72W 38°44.56N 82°37.29W 38°37.36N 82°22.48W 38°43. 18N STREAM ORDER SITE DRAINAGE AREA (KM2) ELEVATION (m.a.s.L) STREAM 2 11.1 195 0.5 ROSGEN STREAM TYPE Bc 2 12.2 189 0.8 A 1 14.3 189 0.8 G 3 28.1 183 0.5 Bc Intermittent 2.2 207 4.5 Bc Intermittent 1.0 224 4.2 Bc Intermittent 2.3 189 1.6 C/Cc 1 3.4 213 1.8 B 2 5.8 189 0.3 C/Cc Intermittent 3.2 232 2.7 C/Cc Intermittent 1.6 219 2.1 B 1 6.6 201 0.1 A Intermittent 2.4 219 4.2 C/Cc 1 3.4 207 0.8 C/Cc 1 5.8 224 23.0 B Intermittent 4.2 183 0.3 A KM 12 within each 50-rn study reach. Spring sampling occurred during April, May, and June; summer sampling occurred in July and early August; autumn sampling occurred in very late August and September. Due to intermittent conditions and drought, not all sites were sampled in every season (Table 2). At least one site in all land use types exhibited intermittent conditions. In autumn 1998, four sites (Cooney Branch, Wolcott Hollow, Turkeyfoot Run, and Webster Creek) were dry due to intermittent conditions (Table 2); these same sites were also dry in autumn 1999. The remaining forested sites (Jenkins Ridge and Meyers Hollow) and agricultural sites (Camp Creek, Long Creek, Sugarcamp Hollow) did not desiccate in autumn 1998; these same sites were recharged with enough water to allow sampling in autumn 1999 following drought (Table 2). During drought in summer 1999, streams in only four sites (Brushy Fork, Negro Creek, Sand Fork, and Sugarcamp Hollow) contained enough water to allow sampling; streams within all forested sites, three agricultural sites, and one mining site were dry in summer 1999 (Table 2). Stream flow was too negligible to be measured in most of these sites. Autumn showers recharged study sites, filling them with enough water to allow sampling from August 31 to September 10, 1999. Prior to the late August showers, four of the six sites sampled contained no water. Table 2. Seasonal samples collected at each site. Land use codes: A = Agricultural, F = Forested, M = Mining. Letters in parentheses indicate basin: P = Pine Creek and S = Symmes Creek. * indicates sites added in 1999, / indicates sites sampled, and - indicates dry sites. STREAM NAME Sugarcamp Hollow (P) Turkeyfoot Run (P) Camp Creek (S) Long Creek (S) Cooney Branch (P) Wolcott Hollow (P) Jenkins Ridge (S) Meyers Hollow (S) Asbury Creek* (S) Bakers Fork* (S) Howard Run* (P) Rocky Fork* (S) Brushy Fork (P) Negro Creek (P) Sand Fork(S) Webster Creek (S) LAND USE A A A A F F F F F F F F M M M M SPRING SUMMER AUTUMN SPRING SUMMER AUTUMN 1998 1998 1998 1999 1999 1999 1 1 1 1 1 1 1 1 1 I 1 1 - (n=4) (n=4) (n=3) (n=4) / / II I (n1) - (n =4) (n =4) (n =2) I II I I / - (n=4) (n=4) (n=3) / I 1 - I I / I I II I I I I I (n I I I/ = 8) (n=4) 1 (n =0) 1 / I (n=3) / I - I / I (n =2) - / - (n=3) (n=1) 14 Habitat Characteristics I evaluated landscape features, stream reach, and site attributes, including water quality and instream habitat, at each site. Habitat transects were located perpendicular to stream flow every 5-rn for the entire 50-rn reach, resulting in a total often transects in 1998. In 1999, I reduced the number of transects from ten to five, placed at 10-rn intervals along the stream reach. This reduction in total transects was made to reduce time and effort in the field after assessment of the 1998 data indicated that five transects would adequately describe the 50-rn stream reach. Channel and bank characteristics were measured at each transect with a combination of transect-line and transect-point methods similar to those described in Simonson et al. (1994) (Figure 3). For each transect-line, wetted channel width, bank erosion, and riparian width (up to 10-rn) were measured and at each transect- point, water depths, substrate composition (visually estimated for a .25 x .25 m quadrat), embeddedness (visually estimated to the nearest 5% for gravel and cobble), and canopy cover (estimated to the nearest 10% using a spherical forest densiometer) were measured. Habitats were classified as riffle, run, or pool habitat. Water quality measurements collected at one location within each reach included water temperature (°C, YSI Model 80'), dissolved oxygen (mg/i, YSI Meter 80), ambient conductivity (pS/cm, YSI Meter 80), turbidity (ntu, Hach 2100A meter'), and pH (Oakton pH meter'). Discharge (calculated using stream velocity as measured across one transect per reach with a Pygmy current meter'), air 'The use of trade or firm names in this publication is for reader information and does not imply endorsement by the U.S. Department of Agriculture of any product or service. 15 Figure 3. Schematic of a typical stream reach (50m following thaiweg), depicting the start and end of the study site. A detailed view of the transect describing transect lines and transect points is shown. (Modified after Simonson et al. 1994). 16 temperature (using a handheld thermometer), and gradient (using a surveyor rod and level) were also measured. Stream order (Strahier), elevation, and drainage area for each site were determined from 7.5-minute USGS topographic maps. Thirteen of the 16 study sites were surveyed for Rosgen stream classification purposes (Rosgen 1994), and an additional 38 sites within WNF were also surveyed (Appendix I). Fishes For each season at each site, fishes were collected from a seine-blocked, 50- m reach on two to four successive passes with a battery operated backpack electrofisher and two fishnetters. On one occasion, only one pass was performed due to equipment failure. Electrofishing proceeded in an upstream direction; all habitats were sampled to ensure that a representative sample of the assemblage was collected, and substrates were disturbed to ensure benthic fishes were dislodged. Standard length, weight, and species of all individuals were recorded before fishes were returned to the stream. Any unknown or uncertain specimens were preserved in 10% buffered formalin for later identification and stored at the Ohio Museum of Biological Diversity in Columbus, Ohio. Data Analysis Multivariate analyses were used because environmental variables (and potentially biological assemblages) operate together and not in isolation (Hawkes et 17 al. 1986, Frenzel and Swanson 1996). Multivariate methods are species dependent methods and are more sensitive in discriminating spatial and temporal patterns than univariate or graphical/distributional methods that are species independent (Warwick and Clarke 1991). Non-metric multidimensional scaling (NMS) (Kruskal 1964, Mather 1976) is a multivariate ordination method based on an iterative rank-ordering procedure resulting in an optimal solution, and is well suited to non-normal data (McCune and Mefford 1999). In particular, NMS has been used in other aquatic ecological studies to examine community patterns of phytoplankton (Salmaso 1996), zooplankton (Sprules 1980), macroinvertebrates (Stephenson etal. 1993, Wright 2000), and fish (McCormick etal. 2000, Rose 2000, Wright 2000). This method positions sites in species space, so those sites with similar assemblages are placed closer together than sites with dissimilar ones. Relationships between assemblages and environmental data were examined using joint plots (McCune and Mefford 1999). To examine spatial and temporal variability in fish assemblage structure, three site-by-species matrices for species catch data were analyzed using NMS (PC-ORD, version 4.14). Streams in which no fish were captured during sampling were not included in the multivariate analyses. For each analysis, I used a Sorensen's (Bray-Curtis coefficient) distance measure at the following settings: 40 runs with original data, 50 Monte Carlo simulations, 0.20 step length, and a 400 iteration maximum. Dimensionality (i.e., number of axes) of the final ordination configurations was determined by choosing only those dimensions that provided 18 the greatest reduction in stress. Stress is an inverse measure of fit to the data and is reported for the final ordination on a 0-100 scale (McCune and Mefford 1999). Final stress for the ordination was less then the final stress in 95% of the randomized Monte Carlo simulations (i.e., Monte Carlo test p-values <0.05). Stability of the final solution was reported by PC-OR]) as the standard deviation in stress over the preceding 15 iterations (McCune and Mefford 1999). All final ordinations were oriented to the same variable to ease interpretation and communication of results. Primary, secondary, and tertiary axes were identified using r2 values for proportion of variance explained by each axis so that the primary axis (Axis 1) accounts for the greatest proportion of variance, or the greatest r2 value. Logarithmic transformation (log 10 (x+1)) of the species catch data was used to include contribution by both common and rare species. Conductivity values were also log transformed prior to analysis, and matrices were checked for outliers (i.e., ±2 standard deviations of the mean average distance) prior to ordination. To express the linear association between environmental variables or fish species and the ordination scores for each axis, I used a Pearson product-moment correlation coefficient (r). The correlation coefficient (r) ranges from +1 to 1 for perfect positive association to perfect negative association. When the correlation coefficient equals zero there is no linear association (Sokal and Rohlf 1987). The coefficient of determination (r2) measures the proportion of variation of one variable determined by the variation of the other. It is a measure of strength and 19 will range from zero to one (Sokal and Rohlf 1987). I used r2 to rank the relative importance of correlations with the ordinations and considered r2 values> 0.500 (r > 0.700) to indicate strong correlation. I used the mean ± one standard deviation to describe habitat variables (e.g., agricultural sites had a mean stream temperature of 20.3 ± 2.6 °C). To evaluate assemblage changes following drought conditions, I compared 1999 autumn assemblages to the 1998 autumn assemblages in agricultural and forested sites for catch per unit effort (CPUE), number of species captured, and species composition. I used Morisita's index (Morisita 1959) and hierarchical agglomerative cluster analysis, with Euclidean distance measure and Ward's method, to compare assemblages among sites and land use for autumn 1998 and autumn 1999. Agglomerative methods sequentially merge objects or groups of objects with other objects or groups, and hierarchical classifications optimize the route through which the groups are attained; Ward's method is a space conserving linkage method (i.e., the original space properties are preserved) compatible with Eucidean distance measure (MeCune and Mefford 1999). Morisita's index has been used in other studies of stream fish assemblages (Ross et al. 1985, Matthews 1986, Matthews et al. 1988, Bart 1989, Matthews 1990, Meador and Matthews 1992, Stewart et al. 1992), and it is not affected by diversity or sample size (Wolda 1981). Index values range from zero (no similarity between samples) to slightly greater than one for identical collections; values 0.40 indicate low similarity (Ross et al. 1985). 0.74 indicate high similarity and 20 RESULTS 1998 and Land Use Patterns Habitat Characteristics In 1998, agricultural sites had a mean stream temperature of 20.3 ± 2.6 °C (Figure 4a), and mean pH of 7.6 ± 0.3 (Figure 4c). Dissolved oxygen values ranged from 1.75 to 9.40 mg/l (Figure 4b). Agricultural sites had low conductivity (X = 249.8 ± 128.8 .tS/cm) (Figure 4d) and widely ranging turbidity values (i.e., from 4.0 to 24.5 ntu; X = 13.3 ± 14.3 ntu) (Figure 4e). Percent canopy cover was reduced in agricultural sites (X =22±27) (Figure 4f). Substrate composition included sand, silt, and gravel (Figure 5a). Agricultural sites ranged in elevation from 183 to 195 meters above sea level. Agricultural sites were classified as Bc and Ge channel types; Bc channel types are moderately entrenched with low gradient and Ge channel types are entrenched with low gradient, sinuosity, and width/depth ratio (Rosgen 1994) (Appendix I). Forested sites had a mean stream temperature of 15.8 ± 3.0 °C (Figure 4a), and a mean pH of 7.3 ± 0.4 (Figure 4c). Dissolved oxygen values ranged from 2.62 to 9.70 mg/I (Figure 4b). Forested sites had low conductivity (X = 148.1 ± 60.1 iS/cm) (Figure 4d) and low turbidity values (X = 3.4 ± 2.8 ntu) (Figure 4e). Percent canopy cover was high in forested sites (X =90±7) (Figure 4f). Substrate composition included gravel, sand, cobble, and detritus, although detectable amounts of detritus were present only in autumn (Figure 5a). Forested 30 a. Stream Temperature 25 . 20 . 15 I 10 5. 12 10 b. Dissolved Oxygen 8 6 4. 2 10 C.PH I 8 6 4 2 0 n4 n4 n3 n=4 n=4 n2 Agricultural Forested n=4 n=4 n=3 Mining I Spring n=4 n1 n=3 Agricultural A Summer n=8 n=0 n=2 Forested it4 n3 n=1 Mining S Autunm Figure 4. Comparison of means for stream temperature (a), dissolved oxygen (b), pH (c), conductivity (d), turbidity (e), and canopy (f) for 1998 and 1999 by land use and season. Error bars indicate one standard deviation of the mean. 1998 3000 1999 d. Conductivity 2000 I. 1000 A K 40 f U e. Turbidity 30 20 10 I 100 80 U I fCanopy 60 4o 2O -- ____________________ ir4 n=4 n=3 n=4 n=4 n=2 Agricultural Forested n=4 n=4 n3 n4 n=1 n3 Mining U Spring Figure 4. Continued Agricultural £ Summer Autumn n=8 n=0 n=2 n=4 n=3 n=1 Forested Mining 24 sites ranged in elevation from 189 to 224 meters above sea level. Forested sites were classified as Bc, C or Cc, and E type channels; C or Cc channel types are slightly entrenched and meandering with moderate to low gradient and a high width/depth ratio, and E channel types are moderately sinuous with low width/depth ratios and low to moderate gradient (Rosgen 1994) (Appendix I). Mining sites were characterized by acidic conditions, with pH ranging from 2.8 to 6.7 (X = 4.7 ± 1.2) (Figure 4c). Mean stream temperature was 17.9 =E 2.1 °C (Figure 4a), and dissolved oxygen values ranged from 3.12 to 9.00 mg/I (Figure 4b). Mining sites had variable conductivity (X =811 ± 775 itS/cm) (Figure 4e) and turbidity (X = 11.5 ± 8.2 ntu) (Figure 4d). Percent canopy cover was high in mining sites (X =91 ±6) (Figure 4f), and substrate composition included gravel, sand, silt, precipitate, and detritus (Figure 5a). As in forested sites, detectable amounts of detritus were present only in autumn. Precipitate only occurred in mining sites. Mining sites ranged in elevation from 183 to 224 meters above sea level. Mining sites were classified as C or Cc and E type channels (Appendix I). All land use types demonstrated a strong seasonal trend in dissolved oxygen values (Figure 4b). Agricultural sites tended to contain greater amounts of sand and silt, while forested and mining sites typically had larger proportions of gravel (Figure 5). Percent canopy cover was most variable in agricultural sites; one site, Turkeyfoot Run, had several large deciduous trees along its banks while the banks of the other agricultural sites were dominated mainly by herbaceous vegetation, bare soil, and/or a few small willows. Also, in autumn 1998, not all sites were 25 sampled due to intermittent conditions (see Table 2); this and/or deciduous leaf drop may have contributed to the trend toward decreasing canopy cover in autumn. Assemblage Characteristics In 1998, agricultural assemblages were dominated by Pimephales notatus, Notropis buccatus, Semolilus atromaculatus, Luxilus chrysocephalus, and Etheostoma nigrum which together comprised greater than 75% of the catch for this land use (Table 3). Pimephales notatus, N buccatus, N. stramineus, Ameiurus natalis, A. me/as, Esox americanus, Hypentelium nigricans, Lampetra aeyptera, Lepomis humilis, L. macrochirus, L. megalotis, L. chrysocephalus, Lythrurus umbratilis, and Micropteruspunctulatus all occurred exclusively in agricultural sites. Agricultural assemblages tended to contain a greater number of species (Figure 6) and a higher catch per unit effort (CPTJE) (Figure 7) compared to assemblages in forested or mining sites. The range was greatest among agricultural sites for these measures; seven to 19 species were captured in spring 1998, and CPT.JE ranged from 36 to 642 in autumn 1998. Forested assemblages were dominated by S. atromaculatus; Phoxinus erythrogaster, Rhynichthys atratulus, L. cyanellus, andE. fiabellare were also common in forested sites. Together these five species comprised 98% of the total catch for forested sites in 1998 (Table 3). Both S. atromaculatus and L. cyanellus occurred in all land use types, but the two dace species rarely occurred in sites other than forested ones. Species composition of forested assemblages was similar ( Table 3. Percent total catch of the most common fishes 1% of total) in 1998 and in 1999 by land use. An asterisk (*) indicates species contributing <1% of the total catch by land use. Numbers in parentheses indicate total catch. Land use codes: Ag = Agricultural, For = Forested, and Mm = Mining. Family Catostomidae Centrachidae Scientific Name Moxostoma erythrurum Lepomis cyanellus Lepomis macrochirus Lepomis megalotis Common Name Golden Redhorse Green Sunfish Bluegill Sunfish Longear Sunfish Campostoma anomalum Central Stoneroller Cyprinidae Luxilus chrysocephalus Striped Shiner Redfin Shiner Lythrurus umbratilis Silveijaw Minnow Notropis buccatus Spotfin Shiner Notropis spilopterus Notropis stramineus Sand Shiner Phoxinus erythrogaster Redbelly Dace Pimephales notatus Bluntnose Minnow Rhynichthys atratulus Blacknose Dace Semotilus afromaculatus Creek Chub Amelurus natalis Yellow Bullhead Ictaluridae Fantail Darter Etheostomaflabellare Percidae Johnny Darter Etheostoma nigrum Blackside Darter Percina maculata Lampetra aepyptera Least Brook Lamprey Petromyzontidae 1998 % Total Catch For Mm Species Ag Code (2827) (1963) (29) MOER 12 14 LECY 1 0 LEMA 1 0 LEME 1 0 CAAN 3 0 0 LUCH 6 * LYUM 0 2 0 20 0 NOBU NOSP 0 0 NOST 4 * 17 3 PHER 0 34 0 PINO 13 RHAT 1 0 14 51 76 SEAT 0 0 AMNA 3 5 0 ETFL 2 * ETNI 5 7 PEMA LAAE 1999 % Total Catch Ag For Mm (930) 1 (633) * 2 2 2 0 0 8 3 1 2 0 0 0 0 20 1 5 15 1 0 16 49 45 1 10 5 1 1 * 2 0 (62) 0 40 0 0 0 0 0 0 0 0 0 50 0 0 0 0 27 2 ---- Camp O. Long -- Sugarcamp Agricultural 2 2( - 1 Turkeyfoot --V \ 14 ic v 4 S V 2 2 22 O Forested -dv-- Meyer 20 5 0 v Wolcott 18 16 0 4- 14 0) 12 C) 0) 10 Cooney Jenkins Asbury Bakers Rocky -- Howard - 8 0 6 0.. ... 4 2 -0- Negro 22 22 Brushy Mining -- Sand --c' Webster 20 18 16 14 12 10 8 6 ---v 4 2 -----V----V 0 SP 98 SU 98 AU 98 SP 99 SU 99 AU 99 Season Figure 6. Temporal patterns in number of species captured for all sites. 700 --- Camp 600 0 Long Agricultural v Sugarcamp Turkeyfoot 500 I\ /\ 400 \ 300 \ \ 200 \ \o 100 / 700 -- Cooney 600 ';;' O Jenkins -Y-- Meyer Forested -ci Wolcott Asbury -0-- Bakers 500 -U--- 400 -- Rocky 0 300 -a-- Howard 0 ................... 200 100 700 Brushy Mining -0- Negro -Y- Sand -V- Webster 650 40 30 / Q. / w _______ SP 98 SU 98 AU 98 SP 99 SU 99 AUO9 Season Figure 7. Temporal patterns in catch per unit effort (CPUE) for all sites. 29 among sites and sites contained between three and nine species (Figure 6). Catch per unit effort was consistently lower at two sites desiccated in autumn (Cooney Branch and Wolcott Hollow) than the two sites persistently wetted (Meyers Hollow and Jenkins Ridge) (Figure 7). Mining sites contained few to no fish, with few species (Figure 6). The number of species captured from any site was typically between zero and two during 1998. Semotilus atromaculatus and L. cyanellus were the dominant species in mining assemblages, accounting for 90% of the fish catch (Table 3); however, CPUE was rarely greater than ten (Figure 7). Catch per unit effort fluctuated minimally during 1998, ranging from zero to 17 in spring. Integrating Assemblage Composition with Habitat Characteristics The ordination of 1998 fish assemblages displayed a land use pattern with three clusters of samples; agricultural sites were located on the left half of the ordination, forested sites in the mid-upper right, and mining sites on the mid-lower right (Figure 8) (Stress of 3-dimensional ordination = 8.94; instability = 0.0000 1; cumulative r2 = 0.9 15). Large-scale reach characteristics (i.e., stream order, elevation, and canopy cover) explained 58% of the variance, while pH accounted for 33%. Only pH seemed to distinguish between assemblages in forested and mining sites (axis two), but stream order, elevation, and canopy cover were the differing characteristics between agricultural and forested/mining assemblages (axis one) (Figure 8a). Low pH was associated with mining sites. Higher 30 a. A AA Land Use o Agricultural A Forested Mining O A aim lull 111i IIJIIJ U Axisi Intermittent r2=.343 o 1Order 2Order b. o AA 0 0 0 A Seat 3"Order A A A Nobu OPino Esam A A 0 .- 11111 11111 U Axis! 111111 r=.343 Figure 8. NMS ordination of 1998 fish assemblages. Overlaid on the ordinations are joint plots of environmental variables (a) and individual fish species (b) strongly correlated (r2 > 0.5) with the ordination axes. Species codes consist of first two letters of genus and species (see Table 5). Each symbol represents one site/date. Final stress of 3-dimensional ordination = 8.94; cumulative r2 = .9 15. 31 elevation, greater percent canopy, and lower stream order were associated with forested and mining sites. Lower elevation, lower percent canopy, and higher stream order were associated with agricultural sites. Species negatively correlated with axis one, including P. notatus, N. buccatus, and L. chrysocephalus, were found in agricultural assemblages and were associated with large-scale reach characteristics (i.e., higher stream order, lower elevation, and less canopy cover) (axis one) (Table 4, Figure Sb). Esox americanus was captured only from agricultural sites, in low abundance overall (Table 5), but this species was also associated with large-scale reach characteristics (Table 4, Figure Sb). Semotilus atromaculatus was a ubiquitous species, occurring in high abundance in all sites regardless of the associated land use (Table 3). However, in 1998 only 22 S. atromaculatus were captured in mining sites compared to 403 in agricultural sites and 1,003 in forested sites. Semotilus atromaculatus abundance increased along a pH gradient from mining sites to agricultural and forested sites (axis two) (Figure 8b,Table 4). 1999 and Examination of Drought Habitat Characteristics In 1999, agricultural sites had a mean stream temperature of 16.2 ± 5.4 °C (Figure 4a), and a mean pH of 8.2 ± 0.6 (Figure 4c). Dissolved oxygen values ranged from 1.34 to 10.48 mg/I (Figure 4b). Agricultural sites had low Table 4. Correlations between axes coordinates from 1998 NMS ordination and species abundance and physical parameters. Correlations greater than r = ± 0.600 are shown, and r2 values> 0.500 indicate strong correlation. Fish Species Axis! r r2 Pimephales notalus Notropis buccatus Esox americanus Luxilus chrysocephalus Lepomis macrochirus Etheostoma nigrum Lampetra aepyptera Ameiurusnatalis Phoxinus erythrogaster -0.832 -0.739 -0.737 0.692 0.546 0.543 -0.718 -0.683 -0.647 -0.628 -0.615 0.515 0.466 0.611 0.373 Axis2 r Semotilus atromaculatus 0.867 Rhynichthys atratulus 0.680 Axis3 0.752 0.462 0.4 18 Etheostoma nigrum Pimephales notatus Ameiurus natalis Luxilus chrysocephalus Notropis buccatus Esox americanus r r2 0.797 0.786 0.709 0.63 6 0.699 0.68 5 0.489 0.469 0.645 0.4 16 0.618 0.503 0.395 0.378 Physical Parameters Axis! Stream Order % Canopy Elevation Land Use r -0.843 0.797 0.757 0.691 r2 0.711 0.636 0.573 0.478 Axis2 r pH 0.730 r2 0.532 Axis3 r % Canopy -0.82 1 Land Use Stream Order Stream Temperature Elevation Dissolved Oxygen -0.740 0.674 0.654 -0.610 -0.600 r2 0.674 0.562 0.455 0.428 0.372 0.360 t'J 33 Table 5. Percent catch of all fishes captured annually and for entire study. *indicates species contributing <1% of total fish catch. Species codes are the first two letters of genus and species. 34 Table 5. % % % 1998 1999 Total Common Name Species Code Catostomus commersoni Hypentelium nigri cans Moxostoma eryfhrurum White Sucker Northern Hogsucker Golden Redhorse CACO RYNI MOER * * * * 0 * * 1 * Lepomis cyanellus Lepomis cyanellusX Lepomisgibbosus Lepomis gulosus Lepomis humilis Lepomis macrochirus Lepomis megalotis Green Sunfish Green Sunfish Hybrids Pwnpkinseed Warmouth Orangespotted Sunfish Bluegill Longear Sunfish Spotted Bass LECY LECX LEGI LEGU LEHU LEMA LEME MIPU 5 3 5 * * * * 0 * 0 * * * 0 * Central Stoneroller Steelcolor Shiner Common Carp Striped Shiner Rosefin Shiner Redfln Shiner Golden Shiner CAAN CYWH CYCA LUCH LYAR LYUM NOCH NOBU NOSP NOST Family Scientific Name Catastomidae Catch Catch Catch Centrachidae Micropteruspunctulatus 1 1 * * 1 * * * * 2 0 * 1 * * * 0 * 4 6 4 * 0 * 1 Campostoma anomalum Cyprinella whipplei Cyprinus carpio Luxilus chrysocephalus Lythrurus ardens Lythrurus umbratilis Notemigonus chrysoleucas Notropis buccatus Notropis spilopterus Notropis stramineus Phoxinus erythrogaster Pimephales notatus Rhynichthys atratulus Semotilus atromaculatus Spotfin Shiner Sand Shiner Redbelly Dace Bluntnose Minnow Blacknose Dace Creek Chub Esox americanus Grass Pickerel ESAM * 0 * Ameiurusme!as Ameiurus natalis Black Bullhead Yellow Bullhead AMME AMNA * * * 2 * 1 Fantail Darter Johnny Darter Blackside Darter ETFL ETNI PEMA 3 5 3 3 3 3 * 1 * Least Brook Lamprey LAAE * 1 * Silveçjaw FIlER PINO RHAT SEAT 1 * * 1 * 12 3 9 0 2 7 20 6 30 1 * * 2 7 8 9 6 48 17 6 34 Esocidae Ictaluridae Percidae Etheostomaflabellare Etheostoma nigrum Percina maculata Petromyzontidae Lampetra aepyptera 35 conductivity (X = 253.3 ± 213.8 jiS/cm) (Figure 4d) and low turbidity values (X = 5.11 ± 1.78 ntu) (Figure 4e). Percent canopy cover was low in agricultural sites (X =3.2±7.1) (Figure 4f), and substrate composition included sand and silt (Figure 5b). Forested sites had a mean stream temperature of 16.2 ± 2.0°C (Figure 4a), and a mean pH of 7.7 ± 0.3 (Figure 4c). Dissolved oxygen values ranged from 1.65 to 8.42 mg/I (Figure 4b). Forested sites had low conductivity (X = 167.9 ± 67.9 j.tS/cm) (Figure 4d) and low turbidity values (X = 3.23 ± 3.78 ntu) (Figure 4e). Percent canopy cover was high in forested sites (X =66±29) (Figure 4. Substrate composition included gravel, sand, and silt (Figure 5b). Mining sites were characterized by acidic conditions, with pH ranging from 5.4 to 6.7 (X = 6.2 ± 0.6) (Figure 4c). Mean stream temperature was 16.8 ± 5.3 °C (Figure 4a), and dissolved oxygen values ranged from 3.72 to 9.72 mg/i (Figure 4b). Mining sites had variable conductivity (X = 1368 ± 923 .tS/cm) (Figure 4e) and turbidity (X 11.9 ± 9.1 ntu) (Figure 4d). Percent canopy cover was high in mining sites (X =60±34) (Figure 4f), and substrate composition included gravel, sand, and precipitate (Figure Sb). Spring canopy cover was variable in forested and mining sites and may have been associated with the timing of deciduous trees leafing out (Figure 4f). Silt was the dominant substrate in summer at Sugarcamp Hollow, the only agricultural site sampled in summer 1999 due to drought (Figure Sb). The proportion of gravel in forested sites seemed to decrease in autumn and therefore the relative proportion 36 of sand and silt increased (Figure 5b), but some forested sites were not sampled in this season due to drought. Substrate composition in agricultural and forested sites exhibited a trend toward finer particle size in 1999 (Figure 5). Agricultural sites exhibited less range in turbidity than in 1998 (Figure 4e), and mining sites were not as highly acidic as in 1998 (Figure 4c). Assemblage Characteristics In 1999, agricultural assemblages were dominated by S. atromaculafus, P. notatus, L. chrysocephalus, E. nigrum, and N. buccatus, comprising 82% of the total catch for agricultural sites (Table 3). These five species were also dominant in 1998 agricultural assemblages, although species ranking changed in 1999. Cyprinella whipplei, L. gulosus, and N. spilopterus were three new species captured in 1999. As in 1998, agricultural assemblages tended to contain a greater number and range of species (Figure 6), higher CPUE (Figure 7), and greater range in CPUE compared to assemblages in forested or mining sites. In Sugarcamp Hollow, the only persistent agricultural site in 1999, fish species numbers ranged from six to 11, and CPUE fluctuated from 20 to 508. Forested assemblages in 1999 were similar to 1998 assemblages, except that L. cyanellus was not a dominant species in 1999 (Table 3). Etheosromaflabellare was rarely found in sites other than forested ones (Table 3). The number of species captured in forested assemblages was between two and eight (Figure 6). Catch per 37 unit effort was relatively low in spring and autumn 1999, ranging from 22 to 110, and was remarkably constant among sites and between seasons (Figure 7). Semoilus atromaculatus and L. cyanellus were once again the dominant species in mining assemblages, accounting for 90% of the fish catch (Table 3). Catch per unit effort fluctuated more in 1999 with as many as 35 fish captured from Sand Fork in autumn (Figure 7). Species numbers in mining sites remained low, but increased marginally in one creek, Sand Fork, during autumn (Figure 6). Integrating Assemblage Composition with Habitat Characteristics The 1999 ordination of fish assemblages revealed a strong seasonal influence (Figure 9b); regardless of land use all spring samples formed one cluster, and a second cluster contained summer and autumn samples (Stress of 3dimensional ordination = 7.73; instability = 0.00001; cumulative r2 0.944). No trend associated with land use was detected in the ordination. Dissolved oxygen and stream order were also correlated with the ordination axes (Figure 9a). Dissolved oxygen explained 40% of the variance; stream order and seasonal patterns explained 37% and 17% of the variance, respectively (Figures 9a and 9b). Luxilus chrysocephalus, E. nigrum, P. notatus, N buccatus, L. megalotis, and L. macrochirus were associated with agricultural sites; L. chrysocephalus was associated with low dissolved oxygen (axis one) while the other five species were associated with stream order (axis two) (Figure lOa,, Table 6). Etheostoma a. 0 0 o Stream Order N 2nd Order 0 0 G A 1 Intermittent i Order 3'' Order Dissolved Oxygen N .- A Ilifi AAA Axis! AAA Land Use o r2=.400 L O b. Agricultural Forested Mining Sugarcamp Sand Camp Season 0 Long Sand Sugarcamp II 0 Sugarcanip I A AAJenkins Long Camp AMeyers A Axis! AA A Dissolve Oxygen Season Spring Summer Autumn r2=.400 Figure 9. NMS ordination of 1999 fish assemblages. Overlaid on the ordinations are joint plots of environmental variables strongly correlated (r2> 0.5) with ordination axes one and two (a) and one and three (b). Each symbol represents one site/date. Only sites sampled more than once per year are identified by name (b). Final stress of 3-dimensional ordination = 7.73; cumulative r2 = .944. 39 Intermittent o 1Order 8 2n Order Etni N Plnoj.eL:ma A 3Order dIED A ALA Axis! 0 AAA r2=.400 Land Use o b. O Agricultural Forested Mining N Season p.- 0 Axisi Spring Summer Autumn r2=.400 Figure 10. NMS ordination of 1999 fish assemblages. Overlaid on the ordinations are joint plots of individual fish species strongly correlated (r2 > 0.5) with ordination axes one and two (a), and one and three (b). Species codes consist of first two letters of genus and species (see Table 5). Final stress of 3-dimensional ordination = 7.73; cumulative r2 = .944. Table 6. Correlations between axes coordinates from 1999 NMS ordination and species abundance and physical parameters. Correlations greater than r = ± 0.600 are shown, and r2 values> 0.500 indicate strong correlation. Fish Species Axis! r Luxilus chrysocephalus Semotilus afromaculatus Pimephales notatus -0.730 0.534 -0.696 -0.667 0.484 0.445 r2 Axis2 r Etheostoma nigrum Pimephales notatus Notropis buccatus Lepomis megalotis Lepomis macrochirus Phoxinus erythrogaster Notropis stramineus Rhynichthys atratulus Lampetra aepyptera 0.796 0.773 0.738 0.634 0.598 0.545 0.7 17 0.5 14 0.5 10 0.714 -0.676 0.663 -0.649 0.606 r2 r Axis3 Etheostomaflabellare -0.762 Axis3 r Season 0.753 r2 0.580 0.457 0.440 0.42 1 0.368 Physical Parameters Axis! Dissolved Oxygen Stream Temperature r 0.782 -0.660 r2 0.612 0.436 Axis2 Stream Order Mean Stream Width Elevation r 0.840 0.695 -0.663 r2 0.706 0.484 0.439 r2 0.567 0 41 flabellare, a dominant fish in forested assemblages, was associated with season (axis three) (Figure lob, Table 6). Response offish assemblages following drought In 1999 fish assemblages from all land use types exhibited seasonal variation, as spring assemblages differed from autumn assemblages. This seasonal variation was not apparent in 1998, and one of the biggest differences between the two years was the occurrence of drought in 1999. Even with a dominant seasonal pattern in 1999, differences in fish assemblages persisted among the different land use types (Figure 11). Because forested sites were classified as intermittent and agricultural sites were perennial streams, I contrasted these sites in autumn 1998 before drought conditions with the same sites in autumn 1999 following drought conditions. Overall, fish assemblages were similar within a location between autumn 1998 and autumn 1999 for both land uses, with the exception of Camp Creek (Table 7a). However, when assemblages were compared by land use, differences between forested and agricultural sites emerged. Comparison of autumn 1998 and autumn 1999 samples revealed higher similarity among forested sites than among agricultural sites; forested sites were grouped together with 90% of the information remaining, while only 40% of the information remained after agricultural sites were grouped (Figure 12). Table 7*. Morisita' s Index values for assemblage comparisons between autumn 1998 and autumn 1999 within a site. Forested_Sites Autumn 1998 - Autumn 1999 0.76 0.63 Jenkins Meyer Agricultural Sites Autumn 1998 - Autumn 1999 0.29 0.88 0.83 Camp Long Sugarcainp Table 7b. Morisita's Index values for autumn 1998 and autumn 1999 assemblage comparisons between sites within agricultural and forested land uses. Sites were compared for assemblage similarity within a season. Agricultural Sites Forested Sites Autumn 1998 Autunin 1999 Jenkins-Meyer 0.51 0.81 Autumn 1998 Autumn 1999 Camp-Long 0.85 0.35 Camp-Sugarcamp 0.82 0.22 Long-Sugarcamp 0.78 0.67 q.j Distance (Objective Function) I 3.IE+01 1.6E+01 2.SE-01 I I I I 4.6E+01 I I 6.2E+01 I I Information Remaining (%) I 50 75 100 I I I I 0 25 I I I Camp 98 Long 98 Sugar 98 Camp 99 Sugar 99 Long 99 Jenkin 98 Jenkin 99 Meyer 99 Meyer 98 Figure 12. Hierarchical agglomerative cluster analysis of autumn 1998 and 1999 fish assemblages for agricultural and forested sites. 45 Forested assemblages were more similar following drought conditions in autumn 1999 than in autumn (Table 7b, Figure 1998 12). Assemblages in all forested sites had reduced CPUE and reduced species numbers following drought in autumn 1999 (Figure 13). Species composition of both forested assemblages in 1999 was reduced to three dominant species: S. atromaculatus and the two dace species (Table 8). These were the same three species that accounted for over 91% 1999 catch in forested sites (Table 3). Less abundant species in of the absent from the assemblages in 1999, 1998 were with the exception of Campostoma anomalum (Table 8). Agricultural assemblages in autumn 1998 and autumn 1999 were highly variable (Figure 12, Table 8), and agricultural assemblages were more similar prior to drought than after (Table 7b). Three new species (N. and L. gulosus) were spilopterus, C. whipplei, captured only in autumn 1999 (Table 8). Rare species (i.e., < 1% of total catch for a given location) in 1998 remained rare, were absent, and/or were common in 1999 assemblages (Table 8). Catch per unit effort decreased in two out of three agricultural sites in autumn 1999 (Figure 13b). Spatial and Temporal Variation in Fish Assemblages The ordination of combined 1998 and 1999 fish assemblages revealed strong differences between years, and an association with land use (Figure 14) (stress of 3-dimensional ordination = 9.90; instability = 0.00001; cumulative r2 0.9 16). Total catch was highest in 1998 with 4,819 fishes compared to 1,625 = fishes 18 a. C' 16 14 I Jenkins 0 Meyer XMean 12 10 8 4 OCamp 0 Long 2 A Sugarcamp 6 0 900 * Mean b. ___________________________________________ 0 Jenkins 0 Meyer Z Mean o Camp 800 700 I] A Sugarcamp Long * Mean I 600 500 400 300 200 100 0 A_____________ 1998 Agricultural 1999 1998 Forested 1999 Figure 13. Comparison of autumn 1998 and autumn 1999 fish assemblages in forested and agricultural sites for number of species captured (a) and catch per unit effort (CPUE) (b). Error bars indicate one standard deviation of the mean. 47 Table 8. Composition of autumn 1998 and autumn 1999 fish assemblages in forested and agricultural sites. Species are listed in descending order based on overall percent catch (see Table 5). A "x" indicates >1% of the total catch for a given location; a "+" indicates 1% of the total catch for a given location. Species indicated in bold are species captured only in autumn 1999. + x maculata Percina spiopterus Notropis x erythrurm Moxostoma ardens Lythrurus megalotis Lepomis macrochirus Lepomis x x x x x + x + + + + + x x x gulosus Lepomis x + x + + x + + x x x x + + + + x x x x x + x + + + x x x x x x x x x x x x x x x x x 1999 1998 1999 Autumn 1998 1999 Autumn 1998 x + x x x x x x x x 1999 1998 Autumn 1999 Autumn 1998 Autumn Autumn Sugarcamp Long Sites A2ricultural + + Autumn Autumn Camp Autumn + Meyer aepyptera Lampetra americanus Esox whipplei Cyprinella commersoni Catastomus melas Amelurus umbratilis Lythrurus anoinalum Cainpostoma natalis Ameiurus stramineus Notropis nigrum Etheostoma Etheostomaflabellare chrysocephalus Luxilus cyanellus Lepomis atratulus Rhynichthys erythrogaster Phoxinus buccatus Notropis notatus Pimephales atromaculatus Semotilus Species Autumn Jenkins Sites Forested 8. Table 49 in 1999. Twenty-nine species were captured in 1998 versus 26 species in 1999. The overall pattern of land use was somewhat muted by seasonal patterns and lowwater conditions in the second year, but land use was a strong correlate of axis one (Table 9, Figure 14) and explained 47% of the variance. Fish assemblage patterns were not distinguishable among basins in the ordination. Pimephales notatus, L. chrysocephalus, N. buccatus, and E. nigrum were associated with agricultural sites (Table 9, Figure 14), and together these four species accounted for 58% of the overall catch for agricultural sites. The two dace species, P. erythrogaster and R. atratulus, were associated with forested sites (Table 9, Figure 14b), and they accounted for 31% of the overall catch for forested sites. Semotilus atromaculatus were the most abundant species in forested sites (5 0%), but they occurred in high abundance in assemblages of all land use types. 50 a. ° 0 Agricultural D o 0 cP 0 Luch Etni Forested . A A A Pino L (#2 A . 1' . aD U A Axis 1 Mining AA A r2.466 b. . S 0 r cPiIio U U 0 SD 00 a U A tni OLuch A (#2 LA Rhat Axis 1 iPher A A A r2=.466 Figure 14. NMS ordination, based on fish assemblage data, projected onto axes one and two (a), and onto axes one and three (b), for 1998 (filled symbols) and 1999 (empty symbols). Overlaid on the ordinations are joint plots of species strongly correlated (r2> 0.5) with the ordination; species codes consist of first two letters of genus and species (see Table 5). Each symbol represents one site/date. Final stress of 3-dimensional ordination = 9.90; cumulative r2 = .9 16. Table 9. Correlations between axes coordinates from 1998 and 1999 NMS ordination and species abundance and physical parameters. Correlations greater than r = ± 0.600 are shown, and r2 values> 0.500 indicate strong correlation. Fish Species Axis! r Pimephales notatus Luxilus chrysocephalus Notropis buccatus Etheosloma nigrum Ameiurus natalis Lepomis macrochirus -0.890 -0.825 -0.821 -0.754 -0.609 -0.606 r2 Axis2 r r2 0.791 Axis3 Phoxinus erythrogaster Rhynichthys atratulus Semotilus atromaculatus 0.68 1 0.673 0.569 r -0.877 -0.876 -0.690 r2 0.770 0.768 0.476 0.37 1 0.368 Physical Parameters Axis! Land Use Stream Order Elevation %Canopy r 0.802 -0.671 0.640 0.639 r2 Axis2 r 0.642 Season 0.921 r2 0.848 Axis3 Stream Order r 0.617 r2 0.380 0.45 1 0.409 0.408 (fl 52 DISCUSSION Both land use and natural disturbance (i.e., drought) were important in explaining fish assemblage patterns during a two-year study of tributary streams within the Wayne National Forest in southern Ohio. With the exception of the ubiquitous creek chub (S. atromaculatus) and green sunfish (L. cyanellus), fish assemblages within agricultural and forested tributaries were dominated by species that rarely occurred in the other land use type. Acid mine drainage tributaries contained few to no fish. Large-scale reach characteristics (i.e., stream order, elevation, and canopy cover) explained the majority of the variance in fish assemblage structure within important. But in 1999, 1998; water chemistry, specifically pH, was also large-scale reach characteristics were secondary to water chemistry (i.e., dissolved oxygen), and seasonal variation was important. These results suggest that large-scale reach characteristics and chemical signatures related to land use are important to fish assemblage structure, but in times of environmental fluctuation, water chemistry or other site-specific variables may also be of importance due to physiological tolerances and limitations of fishes. It is common for fish assemblages to be associated with land use characteristics within a drainage basin. As indicators that land use influences fish assemblages, percent land use in the watershed was correlated with biotic integrity in Wisconsin (Wang et aL 1997), in a Michigan watershed (Roth et in Nebraska streams (Frenzel and Swanson 1996). al. 1996), In New Zealand, fish and 53 distribution has been associated with land use, including habitat and elevational differences (Hanchet 1990, Jowett and Richardson 1996). Riparian deforestation has also been linked to changes in assemblage structure in southeastern streams (Jones et al. 1999). Even so, most studies have focused on environmental degradation rather than patterns in fish assemblage structure within various land use types. Increased species richness in agricultural sites may be associated with stream order, a natural gradient (Sheldon 1968, Vannote et al. 1980, Rahel and Hubert 1991), or with open (i.e., lack ol) canopy, caused by landscape alteration (Murphy etal. 1981). While trends in stream order and elevation were associated with fish assemblages in WNF, neither variable could fully describe assemblage groupings. All sites were categorized as headwater streams (sensu Vannote et al. 1980), and other measurements of stream size, including channel width and depth, were not strongly correlated in the analyses. Land use often incorporates elements of natural landscape gradients (e.g., stream order and elevation) with human impacts (e.g., riparian zone alterations). Therefore, land use and natural gradients were confounded, and that may explain why stream order and elevation were associated with the fish assemblages. Given the small difference in stream orders and minimum elevation change, fish assemblage patterns in small tributary streams of WNF can be described best by land use patterns. The distribution of common species in tributaries in WNF is similar to other studies. For example, P. erythrogaster is commonly found only in headwaters (Trautman 1981, Stewart et al. 1992) and Hawkes etal. (1986) reported that S. 54 atromaculatus in the Kansas River system preferred intermittent streams. In a midAtlantic classification study, McCormick etal. (2000) also found that headwater streams were dominated by two ubiquitous species, R. atratulus and S. atromaculatus, as did Pinder and Morgan (1995) in Maryland Appalachian plateau streams. Additionally, both S. atromaculatus and R atratulus were more abundant in smaller streams in Iowa than in larger ones, and although P. notatus is often depicted as a colonizing species, the species is generally more abundant in larger streams (Starrett 1950). The high tolerance of S. atromaculatus to acid mine drainage conditions was also documented in a Pennsylvania study (Letterman and Mitsch 1978). Forested streams within the WNF are often intermittent and prone to annual desiccation. Such streams may be considered environmentally harsh, containing assemblages of limited species composition. In Illinois streams, Smith (1971) included desiccation as a factor responsible for changes in fish populations, and also in reducing the range of some species. In the Des Moines River, Starrett (1950) attributed limited distribution of certain fishes to their inability to cope with harsh physiochemical and crowded conditions in intermittent streams. Species common in intermittent streams may exhibit greater tolerances to temperature, oxygen, and pH extremes (Matthews and Styron 1981) and high vagility (Poff and Ward 1989). Wide tolerances to physiochemical extremes, or eulytropy, are helpful to organisms that remain in isolated pools during desiccation as these organisms may face suffocation, heat stress, crowding, starvation, and increased 55 predation. In the WNF, intennittent assemblages were dominated by few species, and dominance patterns changed very little over time. Meador and Matthews (1992) also observed that intermittent assemblages were temporally consistent. The consistency of assemblages is likely due to the hardiness of headwater species like S. atromaculatus (Shelford 1937) and P. erythrogaster (Matthews and Styron 1981) combined with the ability to rapidly recolonize desiccated areas. Phoxinus oreas is a congener ofF. erythrogaster and occupies similar habitat (Jenkins and Burkhead 1994). Assemblages in all sites changed following drought. Agricultural sites within the WNF were perennial streams and they exhibited greater variability following drought. Intermittent assemblages in WNF were most similar following drought, implying that assemblages experiencing desiccation on a regular and perhaps predictable basis may be better suited to cope with drought than those assemblages within perennial systems. But not all intermittent streams became desiccated every year, and in those years additional species, such as L. cyanellus, may enter the assemblages of non-desiccated streams; however, these additional species were eliminated during periods of desiccation. A similar phenomena occurs in southwestern montane streams where floods eliminate introduced mosquitofish and allow native topminnows to persist; in streams without floods, introduced mosquitofish have extirpated the native topminnows (Meffe 1984). Moyle and Light (1996) also attributed the success or failure of species invasions to abiotic conditions. Even though short-term changes may be dramatic, faunas usually recover to nonnal levels, and drought typically does not have long term effects on streams (Lanmore et al. 1959, Bayley and Osborne 1993). A number of studies have indicated that small stream assemblages are persistent and stable over time (Moyle and Vondracek 1985, Matthews etal. 1988, Ross et al. 1985, Meffe and Berra 1988) Minckley and Meffe 1987, in spite of short term changes resulting from drought and/or floods. Some physical habitat differences among land use types did not influence fish assemblage structure in this study. Substrate in agricultural sites was finer than in forested and mining sites, yet substrate was not associated with fish assemblages. However, dominant fishes in agricultural sites prefer sandy creek beds (N buccatus), or are silt-tolerant (P. notatus and L. chrysocephalus); dominant fishes in forested sites are usually associated with coarser substrates (Trautman 1981). Conductivity was greatest in mining sites while forested sites were typically less turbid, but neither variable was strongly correlated with fish assemblage structure. In acid mine drainage studies, pH and conductivity are negatively correlated due to increased leaching under low pH conditions (Dills and Rogers 1974), but conductivity may also reflect underlying geology and soils (Frenzel and Swanson 1996). Low turbidity is common in mining streams due to precipitation by acid effluents (Parsons 1955), but turbidity also may be increased by human induced landscape alterations (Karr 1991). Stream habitats are hierarchically nested from the watershed or landscape level down to the microhabitat, and each level of habitat is controlled by characteristics of the higher level, ultimately constrained by 57 watershed characteristics. While patterns in substrate, conductivity, and turbidity are evident, the larger scale variables revealed by the analyses (e.g., stream order, elevation, canopy cover) apparently have greater influence on fish assemblage structure because they help define lower level characteristics. Classification of tributary streams within the WNF will be more accurate when both physical and biological components are incorporated. Multiple environmental variables at multiple scales influenced fish assemblage structure within the WNF, so channel, or geomorphologic, classification must be tightly linked with the biota. Classifiers recognizing the importance of both spatial and temporal variation will gain a better understanding of healthy stream function. A hierarchical channel classification is strongly recommended because the hierarchical nature of streams influences fish assemblage structure over multiple scales (Hawkins et 1999). al. 1993, Maxwell et The approach of Maxwell et al. 1995, al. (1995) Montgomery and Buffington is appropriate for initial classification efforts because it incorporates important measurements to hierarchically map and define characteristics for an overall landscape perspective rather than opting for a single classification system. Fishes are good indicators of environmental integrity and can be used to monitor ecosystem health and changes (Karr 1981, Moyle 1994). In addition, macroinvertebrates are good indicators especially where fishes are not abundant (e.g., mining streams), and may be used in conjunction with fish assemblages. Canopy cover, an important variable explaining fish assemblage patterns, and riparian zones are known to be important 58 to stream function (Karr and Schiosser 1996); 1978, Gregory etal. 1991, Roth etal. therefore, vegetation surveys can provide additional perspective of the aquatic-terrestrial linkage. Land management must incorporate the hierarchical nature of streams into a holistic landscape perspective. Dale etal. (2000) proposed five ecological principles necessary for land use management: time, species, place, disturbance, and landscape. These principles recognize the following: 1) ecosystems are dynamic and potentially exhibit seasonal and annual changes in response to weather (sensu Odum 1969), 2) species composition and diversity may change in response to land use alterations, 3) local environmental conditions may constrain land use patterns, 4) disturbances (both natural and man-induced) influence populations, communities, and ecosystems and, 5) site characteristics may be explained by both local and landscape attributes. In this study I have incorporated these principles by 1) documenting interannual variation in fish assemblages, 2) identifying patterns in fish assemblages within agricultural, forested, and mining land use types, 3) identifying patterns in environmental variables among agricultural, forested, and mining land use types, 4) examining the response of stream fish assemblages to drought, and 5) identifying enviromnental characteristics strongly correlated with fish assemblages. By integrating fish and physical variables in a multivariate gradient analysis I interpreted fish assemblage structure in association with environmental correlates. Descriptive results from this type of multivariate analysis can be used to generate hypotheses for future experimental studies and also provide a template for future monitoring programs or restoration activities. For example, in acid mine drainage tributaries water quality and fish assemblages fluctuated throughout the two years of this study. The variation in these sites along with similarity in physical habitat to forested sites suggests that remediation of pollution in acid mine drainage tributaries could allow recovery of fishes. Because not all portions of a stream network are affected by landscape alteration, unimpacted segments may serve as reffigia (Sedell et al. 1990), allowing fishes to recolonize adjacent areas. 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Ph.D. Thesis, Oregon State University, Corvallis, OR. 68 APPENDIX APPENDIX I collected cross-section data for the purpose of Rosgen stream classification (Rosgen 1994) at 13 fish sites and at an additional 38 sites within the WNF. Bankfull widths ranged from 2.2 to 16.5 meters; entrenchment values ranged from 1.2 to 72; width/depth ration ranged from 6.6 to 87.9. Values for gradient were generally low, ranging from 0 to 2%. Sinuosity was atypically low with values usually from 1.0 to 1.2 with only three exceptions. These low values are associated with "A" type channels and do not concur with the other delineative measurements. However, sinuosity carries the least weight of all the delineative measurements and is one of the most frequently measured incorrectly (Rosgen 1994). By excluding sinuosity, 40 of 51 sites could be classified by entrenchment, w/d ratio, and gradient; three B, nine Bc, 11 C or CC, 13 E, and four F type channels resulted. Disagreement between entrenchment and w/d ratio values was the reason 11 sites could not be classified. However, classification was successful with 10 of the remaining 11 sites when the "fudge factors" were applied: two Bc, one E, and six Gc type channels resulted, and one channel could be either a Bc or Gc channel. B channel types are moderately entrenched with moderate gradient; Bc channel types are moderately entrenched with low gradient. C channel types are slightly entrenched meandering channels with moderate to low gradient and a high width/depth ratio; Cc channel types are C channels with a gradient less than .001. 70 E channel types are moderately sinuous with low width/depth ratios and low to moderate gradient. F channel types are entrenched and meandering with low to moderate gradient, and Gc channel types are entrenched with low gradient, sinuosity, and width/depth ratio (Rosgen 1994). APPENDIX (Continued) Delineative measurements for Rosgen stream classification at 51 sites within the Wayne National Forest boundary. Dominant substrate was used at sites where a pebble count was not performed. Lattitude and longitude are also included. S = Symmes Creek and P = Pine Creek. Bankfull D50 or Pebble Site Basin Width (m) Entrenchment WID ratio Gradient Sinuosity Dominant Count? Lattitude Longitu Asbury/Fish 4.48 3.5 E,C 9.9 A,G,E 0.01 Gc,F,E,Bc,C,Cc 1.2 A 99 S Gravel Y 3844.60 8234.40 99/00 BakersFork/Fish 5.79 3.0 E,C 13.6 B,C,F 0.01 Gc,F,E,Bc,C,Cc 1.0 A S Gravel Y 3844.02 8231.32 Brushy-lower/Fish 99 P 11.19 2.5 E,C 87.9 B,C,F 0 Gc,F,E,Bc,C,Cc 1.0 A Silt Y 3846.88 8238.72 2.56 99 Brushy-upper P 1.9 B 13.0 B,C,F 0.01 Gc,F,E,Bc,C,Cc 1.0 A Gravel Y 3846.96 8238.56 98 Brushyl (CR34) P 4.94 5.3 E,C 7.7 A,G,E 0.01 Gc,F,E,Bc,C,Cc 1.2 A Clay N BrushyBuckeyel 5.85 98 2.1 B 24.2 B,C,F S 3842.57 8233.53 N 98 BrushyBuckeye2 3.29 2.6 E,C S 14.3 B,C,F 0.01 Gc,F,E,Bc,C,Cc 1.0 A Sand N 3843.04 8233.58 98 BrushyBuckeye3 7.04 1.2 A,G,F 31.1 B,C,F 0.01 Gc,F,E,Bc,C,Cc 1.1 A S Gravel N 3842.50 8234.10 98 BrushyBuckeye4 3.05 2.8 E,C S 14.9 B,C,F 0.01 Gc,F,E,Bc,C,Cc 1.1 A Gravel 3843.26 8234.34 N 98 Buffalol 6.40 1.5 B S 17.0 B,C,F 0.01 Gc,F,E,Bc,C,Cc 1.1 A Gravel N 3845.38 8233.50 98 Buffalo2 5.06 S 1.4 B 10.0 A,G,E 0 Gc,F,E,Bc,C,Cc 1.0 A Gravel N 3845.26 8233.45 98 Buffalo3 8.50 7.2 E,C 19.2 B,C,F 0.01 Gc,F,E,Bc,C,Cc 1.0 A S Gravel N 3845.23 8233.40 10.15 99 1.3 A,G,F 15.1 B,C,F S 0 Gc,F,E,Bc,C,Cc 1.2 A Sand Buffalo@Bakers Y 3843.00 8231.18 Buffalo@Bridge 6.70 99 1.5 B 7.4 A,G,E S 0 Gc,F,E,Bc,C,Cc 1.1 A Sand Y 3844.30 8241.37 99 Camp/Fish 4.81 S 1.4 B 6.8 A,G,E 0 Gc,F,E,Bc,C,Cc 1.0 A Saud Y 3845.59 8227.49 Caulley 8.17 99 3.6 E,C 13.3 B,C,F S 0 Gc,F,E,Bc,C,Cc 1.1 A Gravel Y 98 Cooneyl 3.99 P 4.4 E,C 12.0 B,C,F 0.01 Gc,F,B,Bc,C,Cc 2.3 E Gravel N 3841.30 8244.53 98 Cooueyl/Fish P 6.16 1.9 B 18.4 B,C,F 0.01 Ge,F,E,Bc,C,Cc 1.1 A Gravel N 3841.40 8244.54 98 Cooney3 P 2.96 1.9 B 8.5 A,G,E 0.02 B,G,Fb,Eb,Cb 1.2 A Gravel N 3841.54 8244.37 98 Handley 4.66 1.6 B 9.0 A,G,E 0.01 Gc,F,E,Bc,C,Cc 1.0 A S Gravel N 3842.15 8230.23 5.67 99 Howardl/Fish P 1.8 B 31.1 B,C,F 0.01 Gc,F,E,Bc,C,Cc 1.3 G,B Gravel Y 3844.09 8241.72 99 Howardil P 3.29 1.9 B 11.2 A,G,E 0.01 Gc,F,E,Bc,C,Cc 1.0 A Gravel Y 3844.11 8241.68 Year -4 I- APPENDIX (Continued) Bankful Year 98 98 Site Indianl Indian2 98 Indian3 98 Indian4 98 Indian5 98 Indian6 98/99/00 Jenkins/Fish 98 Johnsl 98 Johns2 98 Little Buffalo! 98 Little Buffalo2 Little Pine 99 99 LongfFish 98/00 Meyer/Fish 99 Negro/Fish 99 Pine above Young's 98 Pine2 98 Pine3 99 Pine522 99 Pine@LyraLickRun PineYoung'sBr. 99 98 Pine Creek Chapel SandFork/Fish 99 99 Sugarcamp/Fish 99 Symmes I 99 Symmes II 99 TnbtoBakers 99/00 WolcottlfFish 99 Wolcotifi Basin Wldth(m) Entrenchment WID ratio S S S S S S S S S S S P S S P P P P P P P P S P S S S P P 6.06 6.22 5.55 3.17 4.60 7.04 7.25 10.45 13.53 4.45 2.16 11.46 9.54 3.53 6.77 9.17 11.55 11.25 15.18 11.64 10.00 10.2! 7.71 3.78 9.23 16.49 5.18 5.91 6.10 2.0 B 1.2 1.6 1.4 2.1 A,G,F B B B 2.0 B 3.6 E,C 4.3 E,C 3.1 E,C 4.6 E,C 5.6 E,C 2.6 E,C 1.6 B 3.2 E,C 2.6 E,C 1.5 B 3.6 E,C 2.1 B 2.4 E,C 1.5 B 1.3 A,G,F 4.6 E,C 2.7 E,C 1.4 B 3.0 E,C 1.8 B 1.4 B 5.0 E,C 3.7 E,C Gradient 17.6 B,C,F 26.1 B,C,F 27.3 B,C,F 17.3 B,C,F 22.0 B,C,F 38.9 B,C,F 20.1 B,C,F 0.02 B,G,Fb,Eb,Cb 0.01 Gc,F,B,Bc,C,Cc 0.01 Gc,F,E,Bc,C,Cc 0.01 Gc,F,E,Bc,C,Cc 0.02 B,G,Fb,Eb,Cb 0.01 Gc,F,E,Bc,C,Cc 0 Gc,F,E,Bc,C,Cc 0 Gc,F,E,Bc,C,Cc 6.9 A,G,E 9.2 A,G,E 0.01 GC,F,E,BC,C,CC 0 Gc,F,E,Bc,C,Cc 9.5 A,G,E 6.6 A,G,E 0 Gc,F,E,Bc,C,Cc 10.2 A.G,E 13.8 B,C,F 7.6 A,G,E 13.9 B,C,F 9.7 A,G,E 10.! A,G,B 9.1 A,G,E 10.5 A,G,E 11.8 A,G,E 12.4 B,C,F 10.5 AG,E 11.7 A,G,E 12.! B,C,F 11.5 A,G,E 11.9 A,G,E D50 or Pebble Sinuosity Dominant Count? Lattitude Longitude 1.1 1.0 1.0 1.0 1.1 1.1 1.0 1.1 1.1 1.0 1.2 1.0 1.1 1.2 1.1 A A A A A A A A A A A A A A A A A 0 Gc,F,E,Bc,C,Cc 0 Gc,F,E,Bc,C,Cc 0.01 Gc,F,E,Bc,C,Cc 0.01 Gc,F,E,Bc,C,Cc 0 Gc,F,E,Bc,C,Cc 1.! 0 Gc,F,E,Bc,C,Cc 1.! 0 Gc,F,E,Bc,C,Cc 1.3 G,B 0 Gc,F,E,Bc,C,Cc 0 Gc,F,E,Bc,C,Cc 0 Gc,F,E,Bc,C,Cc 0 GC,F,E,BC,C,CC 2.0 E 0 Gc,F,E,Bc,C,Cc 1.1 A 0 Gc,F,E,Bc,C,Cc 1.0 A 0 GC,F,E,BC,C,CC 0 GC,F,E,BC,C,CC 8.7 A,G,E 0.01 GC,F,E,BC,C,CC 1.0 A 16.0 B,C,F 0.01 Gc,F,E,Bc,C,Cc 1.4 G,B 0 Gc,F,E,Bc,C,Cc 1.0 A 28.8 B,C,F Cobble Gravel Gravel - Cobble Gravel Sand Sand Sand Bedrock Gravel Silt Sand Sand Gravel Sand Sand Sand Saud Sand Sand Sand Gravel Silt Sand Sand Gravel Gravel Saud N N N N N N N N N N N Y Y N Y Y N N Y Y Y N Y Y Y Y Y Y Y 3845.25 8234.11 - 3845.03 3845.36 3845.38 3845.22 3843.02 3842.09 - - 8234.36 8234.51 8235.09 8233.52 8226.47 8230.10 - - 3837.97 3839.26 3841.05 3844.56 3845.30 3845.49 3845.19 3840.66 3846.25 3845.42 3846.16 3837.36 3848.09 3848.69 3848.07 3844.02 3842.57 3842.59 - 8242.82 8228.10 8224.41 8237.29 8238.84 8241.06 8239.03 8242.45 8242.49 8239.67 8241.55 8222.48 8242.39 8226.05 8226.59 8231.54 8242.04 8242.27 -.4 N)