American Fisheries Society Symposium , 2006 2006 by the American Fisheries Society

Effects of Small Dams on Cold Water Stream Fish Communities

DANIEL B. HAYES
Michigan State University, Department of Fisheries and Wildlife 13 Natural Resources Building, East Lansing, Michigan 48824 USA

HOPE DODD1
Illinois Natural History Survey, Center for Aquatic Ecology 607 East Peabody Drive, Champaign, Illinois 61820 USA

JOANNA LESSARD2
Michigan State University, Department of Entomology 243 Natural Science Building, East Lansing, Mighigan 48824 USA

Abstract.Dams provide many benefits to society, yet they have profound impacts on aquatic ecosystems. In addition to blocking fish migration, small surface-release dams on coldwater streams generally increase water temperature below the dam. The goal of this study was to evaluate how small, surface-release dams affect fish communities, separating the total effect into components related to fragmentation and habitat alteration. The approach taken in this study was to synthesize results of two large studies conducted in the Laurentian Great Lakes basin. The first study focused on low-head barriers (<0.5 m in height) built to prevent the upstream migration of sea lamprey Petromyzon marinus. In this study, 24 streams with sea lamprey barriers and 23 nearby reference streams without barriers were sampled. The second study evaluated 10 small hydroelectric dams. Dams that do not substantially warm downstream water temperature cause a moderate shift in fish community composition, whereas dams that warm downstream water temperature by more than 2C cause substantial shifts in fish community composition. The abundance of stream resident coldwater species such as brook Salvelinus fontinalis and brown trout Salmo trutta does not appear to be affected by the presence of a barrier. Dams that increase water temperature result in substantial reductions in abundance, however. These results suggest that the effect of habitat alteration caused by dams on coldwater streams should receive as much, if not greater attention, as the effect of habitat and population fragmentation.

Introduction
Dams have been built on many streams across the world, vastly altering stream ecosystems. The presCorresponding author:[email protected] Current address: Missouri State University, National Park Service, Department of Biology, 901 S. National Avenue, Springfield, Missouri 65897 USA 2 Current address: Tetra Tech Inc., 400 Red Brook Boulevard, Suite 200, Owings Mills, Maryland 21117 USA
1 *

ence of a dam and impoundment in the course of a free flowing river creates a discontinuity in the natural structure and function of a stream (Ward and Stanford 1983), leading to changes in physical, chemical, and biological conditions both upstream and downstream of the dam. The effects of dams vary depending on their size and ecological and environmental setting (Poff and Hart 2002). Some of the dominant impacts include 1) reduc-

587

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tion in sediment and woody material transport; 2) change in water temperature and flow regime; and 3) creation of a barrier to upstream and downstream movement of organisms and nutrients (e.g., Poff and Hart 2002). Dams affect stream fish communities in many ways, but two of the primary mechanisms of direct impact are the alteration of stream habitats above and below dams and fragmentation of stream fish habitat and populations. It is commonly viewed that the combined effect of dams on stream fish communities is to reduce biodiversity and reduce productivity of valued fisheries (Heinz Center 2002). These effects are often obvious where large, bottom-release dams are built on rivers containing anadromous fish populations (e.g., Dauble et al. 2003). The effect of smaller dams is less clear, however. Restoring habitat and population connectivity is often used as a justification for dam removal or developing fish passage, but little quantitative evidence on the benefit is available in the literature (Heinz Center 2002). This presents problems for policy decisions where the benefits of dam removal or fish passage must be weighed against the cost of these management actions. Further, many watersheds contain multiple dams, but the lack of information makes it difficult to prioritize restoration activities. The impact of dams on population fragmentation has received considerable attention, and has often been used as an argument for dam removal (Morita and Yokota 2002). Fragmented populations experience reduced gene flow, resulting in lower effective population sizes and eventually deleterious effects of inbreeding. The impact of habitat alteration on fish populations and communities has received less attention, however. This is particularly true for small dams (e.g., <10 m in height) which have received less study than larger dams, despite the vastly greater number of small dams worldwide. The goal of this study was to evaluate how small,

surface-release dams affect fish communities, separating the total effect into components related to fragmentation and habitat alteration. The specific objectives were to 1) compare differences in fish community composition among reference stream reaches without dams, reaches containing dams with minimal habitat alteration, and reaches with substantial habitat alteration; and 2) compare the density of two valued game fishes (brook trout Salvelinus fontinalis and brown trout Salmo trutta) among stream reaches in these different situations.

Methods
The approach taken in this study was to synthesize results of two large studies conducted in the Laurentian Great Lakes basin. The first study focused on low-head barriers (<0.5 m in height) built to prevent the upstream migration of sea lamprey Petromyzon marinus (Dodd 1999; Dodd et al. 2003). In this study, 24 streams with sea lamprey barriers, and 23 nearby reference streams without barriers, but similar in size and habitat conditions were sampled in 1996. In each stream, three sites approximately 100 m in length were sampled above the barrier and three sites below the barrier (or below a comparable position in reference streams). These barriers were designed to have little or no impact on stream habitat conditions, but are intended to be a barrier to fish migration. The second study evaluated 10 streams with small (<5 m) hydroelectric dams (Lessard 2000; Lessard and Hayes 2003). In this study, three sites approximately 75 m in length were sampled above and below each dam. Two of these streams had a temperature increase of less than 2C below the dam. We had defined this a priori to be lowimpact based on regulations on allowable water temperature increases. In both studies, fish were collected with barge or backpack electrofishing units. In the lamprey bar-

Table 1. Mean ( 1 SE) stream habitat conditions in study streams. Substrate sizes were coded as follows: 1=clay, 2=silt, 3=sand, 4=gravel, 5=cobble, 6=boulder, 7=bedrock.

Mean water temperature (C) Mean stream width (m) Up 8.60.9 10.51.0 503.7 Down Up Up 17.30.6 17.60.6 Down Down 584.8

Mean maximum water depth (cm)

Mean coded substrate size Up 4.00.2 Down 3.90.2

Reference streams

EFFECTS OF DAMS ON FISH COMMUNITIES

Low-head lamprey barriers

16.80.5 9.90.9 12.51.1

16.90.5

593.4

755.6

4.00.2 3.80.2

Low-impact hydrodams

17.81.0 10.91.8

17.51.7

12.40.8

682.6

576.9

3.70.3 4.30.2

High-impact hydrodams

16.61.2

20.00.9

10.51.6

13.51.7

485.8

668.1

3.50.2 3.80.2

589

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rier study, a single pass at each site was performed, providing a measure of relative abundance for each species. A multiple pass depletion method was used in the hydrodam study, allowing for population estimates of large-bodied species. Total catch was used as a measure of relative abundance for species that did not show clear depletion across electrofishing passes. Stream habitat conditions were documented at all sites using the same methods in both studies. Three transects were located at each site, and wetted stream width was measured to the nearest 1 m, maximum water depth was measured to the nearest 1 cm. Substrate size composition was evaluated using the pebble count method (Kondolf and Li 1992). Water temperature was recorded only on the day of fish sampling in the lamprey barrier study, whereas in the hydrodam study, water temperature was collected from 1 May to 30 August using Onset temperature loggers. More detailed specifications of fish and habitat sampling are available in Dodd (1999) and Lessard (2000). Differences in habitat conditions among stream types (e.g., high-impact hydrodam streams) and position (i.e., above or below barrier) were evaluated using a general linear model (Searle 1987). Preplanned contrasts were used to evaluate specific comparisons, such as above lamprey barriers to below lamprey barriers. Fish community composition was compared among stream types and position using correspondence analysis (Manly 1994). The first two dimensions resulting from the correspondence analysis were graphed to visually evaluate the presence of distinct groupings and each dimension was compared across stream groups and position using a general linear model. As with habitat conditions, preplanned contrasts were used to evaluate specific comparisons. The correspondence analysis was limited to a total of 29 species, which were most abundant in the catch and widely distributed across the region sampled. Similarity in fish community composition (i.e., species occurrence )

within a stream was evaluated using Srensons similarity index (Srenson 1948). Similarity in fish community structure (i.e., proportional abundance) was evaluated using Morisitas similarity index (Morisita 1959).

Results
Stream Habitat Conditions

On average, habitat conditions in reference streams (Table 1) showed no significant change in water temperature or substrate size (P > 0.05) between upstream and downstream reaches, but showed a significant increase in stream width (P = 0.003) and a nearly significant increase in depth (P = 0.06). Habitat conditions in streams with lowhead lamprey barriers showed a similar pattern (Table 1), except that water depth showed a significant increase below the barriers (P < 0.001). In the two low-impact hydrodam streams, habitat conditions other that water temperature showed a similar upstream and downstream pattern to reference streams (Table 1), but none of the differences were significant due to the low sample size. Habitat conditions in the eight high-impact hydrodam streams also followed a similar pattern, except for the large difference in water temperature between upstream and downstream reaches (Table 1). Comparing across stream groups, mean stream width and mean substrate size were not different (P > 0.05) among any of the above barrier reaches, nor among any of the below barrier reaches. Mean water depth differed only for reaches below lowhead lamprey barriers relative to below sections of the reference streams (P = 0.01). Water temperature was similar across all stream groups except below the high-impact hydrodams.
Fish Community Composition and Structure

Across both studies, a total of 25,561 individual

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Table 2. Coordinates for each fish species for the first two dimensions of the correspondence analysis. Results are sorted by first dimension coordinates. Species scientific names are given in Appendix 1.

Dimension 1
Rainbow darter Lake chub Blacknose shiner Rosyface shiner Blackside darter Pumpkinseed Hornyhead chub Common shiner Logperch Johnny darter Bluntnose minnow Longnose dace Creek chub Rock bass Green sunfish Eastern blacknose dace Brook stickleback Central mudminnow White sucker Redbelly dace Mottled sculpin American brook lamprey Rainbow trout Coho salmon Brown trout Brook trout Slimy sculpin -0.585 -0.573 -0.563 -0.554 -0.542 -0.536 -0.532 -0.499 -0.486 -0.469 -0.461 -0.383 -0.370 -0.362 -0.345 -0.301 -0.267 -0.239 -0.200 -0.115 -0.091 0.047 0.313 0.770 1.822 2.199 2.952

Dimension 2
0.603 -1.923 1.483 0.692 1.104 0.939 1.097 1.117 0.706 0.309 0.400 -0.411 0.090 0.884 0.584 -0.215 -0.597 0.034 0.408 -0.334 -0.687 -0.274 -1.193 -1.648 -0.035 0.378 0.856

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1.5 1 0.5 0 -0.5 -1 -1.5 -1 0 1 2 3 4

Figure 1. Plot of the first two dimensions from a correspondence analysis of fish species - site associations. For clarity, only the locations of sites are plotted.

Barrier Above Barrier Below Reference Above Reference Below High-Impact Above High-Impact Below Low-Impact Above Low-Impact Below

fish were captured and identified, comprising 87 species (Appendix 1). The top 11 species, including eastern blacknose dace Rhinichthys atratulus, mottled sculpin Cottus bairdii, longnose dace R. cataractae, creek chub Semotilus atromaculatus, rainbow trout Oncorhynchus mykiss, common shiner Luxilus cornutus (also known as Notropis cornutus), johnny darter Etheostoma nigrum, brown trout, white sucker Catostomus commersoni, slimy sculpin Cottus cognatus, and rainbow darter E. caeruleum contributed more than 75% of the number caught. Preliminary analysis of fish community composition using correspondence analysis placed unusually high loadings on fantail darter E. flabellare and black bullhead Ameiurus melas, each of which occurred at a limited number of sites, but in high abundance. Because of their low prevalence, analysis proceeded excluding these species. Results of the correspondence analysis indicated that the first

dimension was positively related to salmonids and slimy sculpin (Table 2), all of which are coldwater specialists. Negative loadings were observed for the remaining species, but none showed loadings as high in absolute value as the salmonid species. The second dimension did not show any apparent pattern related to taxonomic affiliation or temperature preference. A plot of the first canonical dimension versus the second canonical dimension did not reveal any obvious groupings related to position relative to barriers or to the stream group (e.g., reference streams; Figure 1). Comparison of the canonical dimensions within a stream group across the barrier indicated no significant difference within reference streams (dimension 1 P = 0.997; dimension 2 P = 0.784), low-head lamprey barrier streams (dimension 1 P = 0.931; dimension 2 P = 0.724), or the low-impact hydro dams (dimension 1 P > 0.99; dimension 2 P > 0.99), but highly

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significant differences were observed in the first dimension (P = 0.002) for high-impact hydrodam streams, but not in the second dimension (P = 0.761). Fish species richness averaged slightly higher in downstream sampling sites than upstream sites in reference streams (Table 3), resulting in a net increase of 1.7 species from upstream to downstream reaches. This difference was not statistically significant, however (P = 0.19). Stream reaches without dams showed a high degree of community similarity between upstream and downstream sampling sites (Table 3), with Srensons index (SI) averaging 0.69 and Morisitas index (MI) averaging 0.75. Fish species richness in upstream reaches of the streams with low-head lamprey barriers was similar to that in reference streams (Table 3). Species richness was much higher in downstream sections of streams with low-head lamprey barriers (P < 0.001; Table 3), producing a net difference of 4.3 species between upstream and downstream reaches. Both Srensons and Morisitas index indicated a lower degree of similarity in fish community between upstream and downstream reaches in streams with low-head lamprey barriers (Table 3). In the two low-impact hydrodam streams, mean fish species richness decreased from an average of 17.5 species in upstream reaches to 13.0 species in downstream reaches, yielding a net difference of 3.5 species. Given the small sample size, this difference was not significant (P = 0.31). Further, SI and MI indicated that fish community composition and structure were similar between upstream and downstream reaches (Table 3). In the eight streams with hydrodams that had a large temperature impact, mean species richness showed a pattern similar to that in streams with low-head lamprey barriers. Fish species richness increased from an average of 12.0 species in upstream reaches to 16.6 species in downstream reaches (P = 0.038), for a net difference of 4.6

species. Despite a similar pattern in species richness, the structure and composition of the fish community showed a much lower degree of similarity between upstream and downstream reaches in streams with substantial water warming (Table 3). There was a strong, but variable, relationship between temperature change and community structure (r = -0.62, P = 0.032; Figure 2). For hydrodams, MI declined from high values when the temperature difference was less than 2C, to low values (e.g., <0.5) with progressively greater temperature differential. Morisitas index varied substantially among reference streams, but averaged 0.75, a value very close to the linear regression line prediction for hydrodams with a temperature difference of zero (Figure 2). The mean MI for streams with low-head lamprey barriers was 0.52, a value substantially below the regression line for hydrodams (Figure 2), but within the range of variability observed.
Impact on Game Fishes

In reference streams, the frequency of occurrence and relative abundance catch per unit effort (CPUE) of brook trout and brown trout was similar between upstream and downstream reaches (Table 4; P > 0.20). In streams with low-head lamprey barriers, the frequency of occurrence for both species was similar between upstream and downstream reaches, and was similar to reference streams. Catch per unit effort of brook trout was higher in upstream reaches (P = 0.004), but CPUE was similar among reaches for brown trout (P = 0.68). Brook trout only occurred in one of the two low-impact hydrodam rivers, and density of brook trout in the downstream reach was approximately 17% of that in the upstream reach (Table 4). Brown trout occurred in both low-impact hydrodam rivers, and their density averaged approximately 400% higher in downstream reaches than in upstream reaches, but this difference was not significant (P = 0.74) due to small sample

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Table 3. Mean species richness (1 SE), Srensons similarity index and Morisitas similarity index in study streams.

Mean Species Richness Upstream Downstream 11.40.7 0.690.03 9.70.8 Srensons Similarity Index Morisitas Similarity Index 0.750.06

Reference streams

Low-head lamprey barriers 10.50.9 14.81.0

0.610.03

0.520.04

HAYES ET AL.

Low-impact hydrodams 17.53.5 13.02.0

0.760.14

0.870.06

High-impact hydrodams 12.02.4

16.61.7

0.490.08

0.370.10

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1 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0 -2 0 2 4 6 Temperature (C) change

Community Similarity

Reference Streams Low-head dams Hydro dams

Figure 2. Relation between temperature change caused by hydrodams and community similarity as measured by Morisitas Index. Low-head and reference streams data points are means 2 SE, and are included for reference relative to results for the hydrodams.

size. The frequency of occurrence of brook trout and brown trout was lower in downstream reaches of rivers with high-impact hydrodams than in upstream reaches. Mean density of brook trout and brown trout was also substantially lower in downstream reaches (P = 0.016), averaging only 1% of upstream density for brook trout and 33% for brown trout (P = 0.017; Table 4).

Discussion
Results in our reference streams provide a strong basis for evaluating the patterns observed in streams with low-head lamprey barriers and hydrodams. The slight increase in species richness from upstream to downstream reference reaches is consistent with the river continuum concept (RCC), and with other studies showing increases in species richness in larger streams (Lyons 1996). Both

measures of fish community composition (i.e., SI) and fish community structure (i.e., MI) show that, on average, fish communities are highly similar over the scale of our sampling (typically 1020 river km). Results of the correspondence analysis support this conclusion, indicating no significant difference in community composition of the numerically dominant species. It is important to note that measures of similarity are sensitive to the level of sampling effort and sampling variability. In this study, nearly all streams were sampled at three upstream and three down stream sites, and site length and width were similar, enabling comparisons among stream groupings. Streams with low-head lamprey barriers generally showed a greater discontinuity in species richness between upstream and downstream reaches than was observed in reference streams. It is interesting

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Table 4. Measures of abundance ( 1 SE) and frequency of occurrence for brook trout and brown trout in study streams.

Brook Trout Upsteam Upsteam 1.90.9/pass 6 streams 0.10.04pass 6 streams 11136/ha 2 streams 672332/ha 8 streams 53/ha 3 streams 0.90.3/pass 7 streams 0.50.2/ pass 5 streams 0.10.04/pass 8 streams 4242/ha 1 stream 1.00.3/pass 8 streams 248248/ha 1 stream 370290/ha 6 streams Downstream

Brown Trout Downstream 2.31.1/ pass 5 streams

HAYES ET AL.

Reference streams n=23 streams

Low-head lamprey barriers n=24 streams

0.10.04/pass 5 streams 402347/ha 2 streams 220112/ha 6 streams

Low-impact hydro dams n=2 streams

High-impact Hydro-dams n=8 streams

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to note, however, that this discontinuity occurred due to higher species richness below dams rather than due to reduced richness above dams. In fact, the point estimate for species richness in upstream sections of streams with low-head lamprey barriers was higher (but not significantly so) than the point estimate for reference streams. More detailed analysis on patterns of species richness in streams with low-head lamprey barriers (Dodd 1999) suggest that the heightened species richness below low-head lamprey barriers is due primarily to an accumulation of species below these dams. Interestingly, this accumulation of species richness is not accompanied by higher overall density of fishes below dams (Dodd 1999). These observations indicate that low-head lamprey barriers tend to achieve their intended goal (i.e., prevent access to upstream spawning sites by sea lamprey) without fragmenting the populations and habitats of other fishes to such an extent that species richness in upstream reaches is compromised. The high degree of similarity indicated by the SI for lamprey barrier streams suggests that the composition of fish communities perched above lowhead lamprey barriers is not strongly altered by the presence of the dam. Morisitas index suggests, however, that the upstreamdownstream structure of the fish community differs to a greater extent than it does in reference streams. Thus, few species are lost above these barriers, but the relative abundance of some species is altered. Because of the high variability in density and sporadic occurrence of many species, it is difficult to point to individual species that are particularly susceptible to the fragmentation effect of low-head lamprey barriers (Dodd 1999). Results of the correspondence analysis likewise do not indicate any directional change in species composition. In streams with hydrodams causing substantial increases in downstream water temperature, upstream species richness was similar to both reference streams and streams with low-head lamprey barriers, providing additional evidence support-

ing the idea that fragmentation by itself had relatively little influence on this attribute of fish communities. The increase in species richness below hydrodams is consistent with the observations in streams with low-head lamprey barriers, but we suspect that it is also partially due to increases in water temperature. Other studies (e.g., Brooker 1981) have shown that species richness in streams typically increases with increasing water temperature. This is one area where differences in the study designs may have substantial influence on the results. The low-head lamprey barrier streams and the references streams were selected from streams with direct connections to the Great Lakes themselves. The streams in the hydrodam study, on the other hand, generally did not have direct, unimpeded access to the Great Lakes. Thus, the presence of potadromous fishes in the lamprey barrier streams may create a situation where more migratory species are available to accumulate below these barriers. The substantially lower values of Srensons and Morisitas indices in streams with high-impact hydrodams strongly suggest that alteration of downstream habitat conditions (in this case, water temperature) has much greater influence than does fragmentation by itself. In fact, the plot of Morisitas similarity index versus water temperature change (Figure ) suggests that hydrodams with no impact of water temperature have longitudinal patterns in community structure that is similar to reference streams. The significant difference in the first dimension, which was related to coldwater specialists, produced by the correspondence analysis is further evidence of a directional shift in species composition below the high-impact hydrodams. Although the thermal niche of brook trout, brown trout, rainbow trout, and slimy sculpin differ somewhat, the increase in temperature below the high-impact dams was enough to reduce the abundance of one or more of these species. The abundance and frequency of occurrence of

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HAYES ET AL. of the Great Lakes. Masters thesis. Michigan State University, East Lansing. Dodd, H. R., D. B. Hayes, J. R. Baylis, L. M. Carl, J. D. Goldstein, R. L. McLaughlin, D. L. G. Noakes, L. M. Porto, and M. L. Jones. 2003. Low-head lamprey barrier effects on stream habitat and fish communities in the Great Lakes basin. Journal of Great Lakes Research 29(Supplement 1):386402. Heinz Center. 2002. Dam removal: science and decision making. H. John Heinz Space Center for Science, Economics, and the Environment. Washington, DC. Kondolf, G. M., and S. Li. 1992. The pebble count technique for quantifying surface bed material size in instream flow studies. Rivers 3:8087. Lessard. J. L. 2000. Temperature effects of dams on coldwater fish and macroinvertebrate communities. Masters thesis. Michigan State University, East Lansing. Lessard, J. L., andD. B. Hayes. 2003. Effects of elevated water temperature on fish and macroinvertebrate communities below small dams. River Research & Applications 19:721 732. Lyons, J. 1996. Patterns in the species composition of fish assemblages among Wisconsin streams. Environmental Biology of Fishes 45:329341. Manly, B. F. J. 1994. Multivariate statistical methods a primer. Chapman and Hall/CRC, Boca Raton, Florida. Morisita, M. 1959. Measuring of interspecific association and similarity between communities. Memoires of the Faculty of Science, Kyushu University, Series E Biology 3:6580. Morita, K., and A. Yokota. 2002. Population viability of streamresident salmonids after habitat fragmentation: a case study with white-spotted charr (Salvelinus leucomaenis) by an individual based model. Ecological Modeling 155:8594. Poff, N. L., and D. D. Hart. 2002. How dams vary and why it matters for the emerging science of dam removal. Bioscience 52:659668. Searle, S. R. 1987. Linear models for unbalanced data. Wiley, New York. Srenson, T. 1948. A method of establishing groups of equal amplitude in plant sociology based on similarity of species content. Kongelige Danske Videnskabernes Selskabs Skrifter 5:134. Ward, J. V., and J. A. Stanford. 1983. The serial discontinuity concept of lotic ecosystems. Pages 2942 in T. D. Fontaine and S. M. Bartell editors. Dynamics of lotic ecosystems. Ann Arbor Science, Ann Arbor, Michigan.

coldwater game fish (brook trout and brown trout) showed little difference between upstream and downstream reaches in reference streams. Point estimates of brook trout abundance in low-head lamprey barrier streams and low-impact hydrodam streams suggest that the downstream abundance of this species may be reduced due to fragmentation effects, but variability among streams and sites make this relationship uncertain. Brown trout show no clear pattern of decline from upstream to downstream in either the lamprey barrier streams or low-impact hydrodam streams, suggesting that fragmentation affects this species to a lesser extent than brook trout. The large decline in overall trout abundance below high-impact hydrodams demonstrates the degree to which thermal habitat alteration can affect these species. As such, dams on coldwater streams can pose a serious threat to these fisheries, but the principal effect of dams on these species is through habitat alteration rather than habitat and population fragmentation.

Acknowledgments
The support of the Great Lakes Fishery Commission, the Michigan Department of Natural Resources, and Michigan State University is gratefully acknowledged.

References
Brooker, M. P. 1981. The impact of impoundments on the downstream fisheries and general ecology of rivers. Advances in Applied Biology 6:91152. Dauble, D. D., T. P. Hanrahan, D. R. Geist, and M. J. Parsley. 2003. Impacts of the Columbia River hydroelectric system on main-stem habitats of fall Chinook salmon. North American Journal of Fisheries Management 23:641659. Dodd, H. R. 1999. The effects of low-head lamprey barrier dams on stream habitat and fish communities in tributaries

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Appendix 1. List of species and number of individuals in descending order of total catch.

Common name

Scientific Name

Total Catch

Eastern blacknose dace Mottled sculpin Longnose dace Creek chub Rainbow trout Common shiner Johnny darter Brown trout White sucker Slimy sculpin Rainbow darter Central mudminnow Rock bass Brook trout Fantail darter Hornyhead chub Blacknose shiner Bluntnose minnow American brook lamprey Logperch Black bullhead Brook stickleback

Rhinichthys atratulus Cottus bairdii Rhinichthys cataractae Semotilus atromaculatus Oncorhynchus mykiss Luxilus cornutus Etheostoma nigrum Salmo trutta Catostomus commersoni Cottus cognatus Etheostoma caeruleum Umbra limi Amblophites rupestris Salvelinus fontinalus Etheostoma flabellare Nocomis biguttatus Notropis heterolepis Pimephales notatus Lampetra appendix Percina caprodes Ameiurus melas Culaea inconstans

3184 2621 2477 2130 1800 1740 1450 1314 1203 854 748 556 479 419 387 302 292 248 246 241 217 212

600

HAYES ET AL.

Appendix 1 continued.

Common name Rosyface shiner Northern redbelly dace Pumpkinseed Lake chub Blackside darter Northern hog sucker Coho salmon Green sunfish Bluegill Burbot Cutlips minnow Yellow perch Smallmouth bass Warmouth Threespine stickleback Spotfin shiner Brassy minnow Blackchin shiner Largemouth bass Shorthead redhorse Common carp Fathead minnow

Scientific Name

Total Catch 199 166 157 154 151 118 100 100 98 92 88 86 77 72 67 63 57 53 49

Notropis rubellus Phoxinus eos Lepomis gibbosus Couesius plumbeus Percina maculata Hypentelium nigricans Oncorhynchus kisutch Lepomis cyanellus Lepomis macrochirus Lota lota Exoglossum maxilingua Perca flavescens Micropterus dolomieui Lepomis gulosus Gasterosteus aculeatus Cyprinella spilopterus Hybognathus hankinsoni Notropis heterodon Micropterus salmoides

Moxostoma macrolepidotum 45 Cyprinus carpio Pimephales promelas

44 43

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Appendix 1 continued.

Common name Northern pike Pearl dace Sea lamprey Stonecat Northern brook lamprey Iowa darter River chub Trout-perch Central stoneroller Yellow bullhead Brown bullhead Fallfish Emerald shiner Chinook salmon Bowfin White crappie Golden Shiner Spottail shiner Golden redhorse Lake trout Mimic shiner Sauger

Scientific Name

Total Catch 35 33 29 29 20 20 20 19 14 14 12 10 10 9 9 9 6 6 5 5 5 4

Esox lucius Margariscus margarita Petromyzon marinus Noturus flavus Ichthyomyzon fossor Etheostoma exile Nocomis micropogon Percopsis omiscomaycus Campostoma anomalum Ameiurus natalis Ameiurus nebulosus Semotilus corporalis Notropis atherinoides Oncorhynchus tshawytscha Amia calva Poxomis annularis Notemigonus crysoleucus Notropis hudsonicus Moxostoma crythrurum Salvelinus namaycush Notropis volucellus Sander canadense

602

HAYES ET AL.

Appendix 1 continued.

Common name Silver shiner Striped shiner Redfin pickerel Black crappie Atlantic salmon Greater redhorse Ninespine stickleback River darter Ruffe Silver redhorse White bass American eel Channel catfish Chestnut lamprey Finescale dace Flathead catfish Pugnose minnow Red shiner Southern redbelly dace Walleye Longnose sucker

Scientific Name

Total Catch 4 4

Notropis photogenis Luxilus chrysocephalus

Esox americanus vermiculatus 4 Poxomis nigromaculatus Salmo salar Moxostoma valenciennesi Pungitius pungitius Percina shumardi Gymnocephalus cernuus Moxostoma anisurum Morone chrysops Anguilla rostrata Ictalurus punctatus Ichthyomyzon castaneus Phoxinus neogaeus Pylodictis olivaris Opsopoeodus emilie Notropis lytrensis Phoxinus erythrogaster Sander vitreus Catostomus catostomus
3 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1