SOME RESPONSES OF A WARM WATER STREAM FlSH COMMUNITY T0 DIFFERING LEVELS OF PERTURBATION AND ENRICHM‘ENT 1' basis for the Degree 0f M. S. MICHEGAR STATE UNSVERSITY CHRESTOPHER J. SCHMITT 1974 3 1 358 3989 ' flfl\\$\1|1\\\2\||\9“|\“\\|WM “PM H \\ HM v! WQ ABSTRACT SOME RESPONSES OF A WARM WATER STREAM FISH COMMUNITY TO DIFFERING LEVELS OF PERTURBATION AND ENRICHMENT 'By Christopher J. Schmitt The fish community of the Red Cedar River, a warm water stream in south-central Michigan, was investigated above and below an urban area known to be a source of perturbation and enrichment. Although growth was generally faster for all fishes from the lower, more enriched area than the upper, among-location differences in back-calculated total length at annulus II were significant only for rock bass (Ambloplites rupestris). Species composition and diversity index, a, were similar at both locations but the distribution of biomass and numerical abundance, as indicated by non-parametric tests of rank, was significantly different. The consistent absence of the rainbow darter (Etheostoma caeruleum) at the lower site was noted. Fish standing crop estimates computed for both locations were greater than corresponding estimates made in previous years. Rock bass biomass demonstrated the greatest increase in the last 10 years, approaching 80 lbs acre"l (89.7 kg -1 — ' ha ) at the upper site and 25 lbs acre 1 (28.0 kg ha i) at the lower. Smallmouth bass (Micropterus dolomieui) and nOrthern pike (Esox lucius), the major game species, com- Prfised 2-12% and 0.3—2% of the total fish biomass of 671.3 Christopher J. Schmitt lbs acre_l (752.“ kg ha-l) and 436.3 lbs acre.l (A89.0 kg ha-l) at the upper and lower sites, respectively. Fish standing crop appears unrelated to net primary productivity in the enriched warm water stream. Trophic changes and physical-chemical stream alterations associated with population growth and watershed urbanization are dis- cussed as causative factors which determine the standing crop and species composition of the fish community. SOME RESPONSES OF A WARM WATER STREAM FISH COMMUNITY TO DIFFERING LEVELS OF PERTURBATION AND ENRICHMENT By Christopher J. Schmitt A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Fisheries and Wildlife 197A at? 3 C (7%) ACKNOWLEDGEMENTS I extend my sincere gratitude to all the people with whom I was associated during my course of study for the educa— tional fulfillment derived from many hours of stimulating discussion. I am particularly indebted to Dr. Thomas G. Bahr for his overall guidance, continued encouragement, and assistance in the preparation of this manuscript. He, along with the other members of my guidance committee, Dr. Eugene W. Roelofs, Dr. Walter H. Conley, and Dr. Donald J. Hall, gave freely when advice and time were requested. I would also like to acknowledge the continued assistance of Dr. Terry A. Haines, project director, and Dr. Robert C. Ball for making available the facilities of the Institute of Water Research. Sincerest thanks are extended to my fellow graduate students, Drs. Wayne L. Smith and Adam T. Szluha and Scott Reger, along with the rest of our field and laboratory personnel, for their assistance in gathering and preparing the materials utilized in this investigation. This study was supported, in part, by funds from Grant lA-Bl-OOOl-3153, provided by the United States Department of the Interior, Office of Water Resources Research, as authorized by thr Water Resources Research Act of 196A and, in part, by the National Science Foundation Institutional Grant for Science, both administered by the Institute of Water ii Research, Michigan State University. Funds and equipment were also provided by the Department of Fisheries and Wild- life and Agricultural Experiment Station, Michigan State University. Use of Michigan State University computing facilities was made possible through support, in part, from the National Science Foundation. 111 TABLE OF CONTENTS Page INTRODUCTION . . . . . . . . . . . . . . . 1 SITE DESCRIPTION . . . . . . . . . . . . . 6 MATERIALS AND METHODS . . . . . . . . . . 1“ Fish Sampling . . . . . . . . . . . . 1A Age and Growth . . . . . . . . . . . 15 Population Estimates . . . . . . . . 16 Standing Crop Estimates . . . . . . . 19 Diversity . . . . . . . . . . . . . . 19 Water Quality . . . . . . . . . . . . 20 Fish Kill . . . . . . . . . . . . . . 20 RESULTS . . . . . . . . . . . . . . . . . 21 Length—Weight Relationships . . . . . 21 Growth Study . . . . . . . . . . . . 21 Species Composition . . . . 38 Estimates of Population Density and Standing Crop . . . . . . . . . . 39 Diversity . . . . . . . . . . . . . . 56 DISCUSSION . . . . . . . . . . . . . . . . 58 SUMMARY AND CONCLUSIONS . . . . . . . . . 73 LITERATURE CITED . . . . . . . . . . . . . 76 APPENDIX . . . . . . . . . . . . . . . . . 81 iv Table 10 LIST OF TABLES Physical and chemical water quality parameters at the Red Cedar River study sites, 1971-72, expressed as means and ranges ( ). From Ball and Kevern, 1973 except where indicated Maximum and minimum water temperatures (C) and dissolved oxygen concentrations (mg 1'1) at the Red Cedar River study sites, 1971-72. From Ball and Kevern (1973) . . . . . . . . . . . . Length-weight regressions of Red Cedar River fishes from 2 locations, 1971-72 ~Scale radius-total length regressions and correlation coefficients of Red Cedar River fishes from 2 locations, 1971-72 . . . . . . . . . . . . Age and growth computations for Red Cedar River rock bass from 2 locations, 1971 . . . Age and growth computations for upper Red Cedar River smallmouth bass, 1971 Age and growth computations for Red Cedar River white suckers from 2 loca- tions, 1971 . . . . . . . Age and growth computations for Red Cedar River golden redhorse from 2 locations, 1971 . . . . . . . . . . . Standing crop, numerical abundance, and % composition, by family, of the Red Cedar River fish community at 2 locations, 1971-72 . . . . . . . Estimated numerical abundance and standing crop of Red Cedar River fishes by species or species group, at 2 locations, 1971-72 . . . Page 12 13 23 3A 35 36 37 A0 A2 List of Tables (con't.) Table ll 12 13 1A 15 A-3 A-A A-S A-6 Contribution of the major Red Cedar River fish groups to the total stand- ing crop at 2 locations after subtract- ing the contribution of minnows and darters, 1971-72 . . . . . . Species composition of the Red Cedar River Catostomidae at 2 locations, 1971-72 . . Species diversity (5) and equitability of the Red Cedar River fish community at 2 locations, 1962, 1971, 1972 (1962 data from Horton, 1969) . . . . . An energy budget for the Red Cedar River at 2 locations, 1971-72. All values, except where indicated, in gC m"2 day'l. From Ball and Kevern (1973) Characteristics of the Red Cedar River in 3 stages of hypothetical succession Species collected at the upper station, lower Red Cedar site, August 3-6, 1971 Species collected at the lower station, lower Red Cedar site, August 3—6, 1971 Species collected at the lower station, lower Red Cedar site, July 23-25, 1972 Species collected at the lower station, upper Red Cedar site, August 3-6, 1971 Species collected at the upper station, upper Red Cedar site, August 3-6, 1971 Species collected at the lower station, upper Red Cedar site, July 23—25, 1972 vi Page 50 52 57 62 63 81 83 85 87 9O 93 Figure LIST OF FIGURES Map of the Red Cedar River watershed Map of the Red Cedar River showing upper and lower study sites . . . Back—calculated total length (TL) (mm) at annulus formation 1 l S.E. and instantaneous growth rate (G) of Red Cedar River rock bass at two locations, 1921, 1971, 1972 (1961 data from Linton, 19 A) . . . . . . . . . . . . . . . . . Back-calculated total length (TL) (mm) at annulus formation i 1 S.E. and instantaneous growth rate (G) of upper Red Cedar River smallmouth bass, 1971-72 Back-calculated total length (TL) (mm) at annulus formation i 1 S.E. and instantaneous growth rate (G) of Red Cedar River white suckers at 2 locations, 1971-72 . . . . . . . . . . . . . . Back-calculated total length (TL) (mm) at annulus formation 1 l S.E. and instantaneous growth rate (G) of Red Cedar River golden redhorse at 2 loca— tions, 1971-72 Relative numerical and standing crop abundance of the most frequently en- countered Red Cedar River fish families at 2 locations, 1971-72 . . . . . Species composition of the Red Cedar River Catostomidae at 2 locations 1961, 1962, 1971, 1972 (1961 data from Linton and Ball, 1965; 1962 data from Horton, 1969) . . . . . . . . . . . . . . . . vii Page 10 27 29 31 33 A8 5A List of Figures (con't.) Figure 9 Page Hypothetical temporal-longitudinal stream succession. A "natural" wood- land stream with relatively stable water levels and low sediment loads in degraded to an autotrophic stage char- acterized by high nutrient levels, sediment loads, and primary productivity and greater discharge variability be- cause of deforestation and intensive agriculture. Further degradation re- sults in a heterotrophic state based on organic wastes where urban runoff and resultant storm drain overflow along with water removal, cause severe water level fluctuations and high sediment loads. 1 - complex, heterotrophic, shade adapted, cool water system charac- teristic of Red Cedar River tributary headwaters; 2 - simplified, auto- trophic, light adapted, warm water sys- tem represented by the Red Cedar River in the vicinity of the upper site; 3 — simplified, heterotrophic, warm water system represented by the Red Cedar River at the lower site. Re- drawn from Cummins (1972) . . . . . . . 6O viii INTRODUCTION The enrichment and perturbation of streams through man's activities occurs at an accelerated rate as industrial centers and associated urban developments become typical character- istics of an increased number of watersheds. Urbanization is a complex process with poorly understood physical, chemical, and biological interactions. Industrial processes subject streams to point sources for the introduction of toxic substances while population centers also downgrade the quality of natural waterways through organic and inorganic enrichment. Organic materials, largely the remains of human waste materials, enter streams through combined sewer overflow, stormdrains, and inefficient or non-existent sewage treatment causing heterotrophic production and associ- ated high respiratory oxygen demand (King and Ball, 1967). Autotrophic production is stimulated by the introduction of plant nutrients, nitrogen and phosphorus, produced in domestic wastewater and as a byproduct of the intensified agriculture necessary to sustain the urban communities and by the impoundment of flowing waters. Additionally, the process of urbanization itself involves denuding of the watershed, alteration of natural flow regimes, and mis- managed construction projects resulting in erosion, sedimen— tation, and siltation (King and Ball, 196“). These practices, 1 2 along with wetland drainage and agricultural, domestic, and industrial water removal reduce the stream's ability to absorb the impact of enrichment. Concurrent with the degradation of streams through urbani— zation and industrialization, however, has come the high standard of living associated with the urban, technologically advanced society. This standard involves considerable lei- sure time, resulting in a high recreational demand being placed upon natural waterways. Recently there has been con- siderable monetary expenditure towards the rehabilitation of streams for recreational pursuits. While Hooper (1969) advocates the adoption of common biological indices to eutrophication and enrichment, most efforts in this area have been directed towards the study of benthic macroinvertebrate communities. Larkin and Northcote (1969) point out, however, that the avoidance reactions of fishes, because of their well developed sensory abilities and mobility, may provide extremely sensitive indicators of subtle environmental change. Additionally, one of the final measurement of any rehabilitation program will be the quantity and quality of the fish fauna a stream supports. Unlike cold-water stream systems, warm-water stream communities are generally highly productive complex assemblages of many species. While evidence of change in the relative abundance of the few species sought by sportsmen is generally available, subtle, long—term responses of a warm-water stream fish community as a whole, rather than the responses of the few gamefish present in the community, are largely unknown. 3 Simple artificial cold-water stream communities have been shown to respond positively to organic enrichment (Warren, 32 31., 196A). McFadden and Cooper (1962) found generally fertile streams to produce larger brown trout (Salmo trutta) than infertile streams. Quick (1971) observed more rapid growth of cold-water fishes in enriched Michigan streams than in unenriched streams. Ball and Tanner (1951) demonstrated increased growth and standing crop of pond fishes after fertilization and many investigators (see Larkin and Northcote, 1969) have shown similar positive responses in lakes. This relationship, however, has never been successfully demonstrated in the warm-water stream environment. The Red Cedar River, because of its proximity to Michigan State University has been intensively studied, especially during the period 1959—1964. During this period the river was receiving untreated and primary treated sewage effluent from all the municipalities upstream of East Lansing (Figure 1). A metal plating plant in Fowlerville (Figure 1) produced toxic effluents which caused frequent fish kills (Garton, 1968). The Lake Lansing Drain, the outlet of an eutrophic lake with a surrounding community, joins the Red Cedar River between Okemos and East Lansing and represented a major source of phosphorus and organic enrichment (Vannote, 1961). Con- struction of an interstate highway along much of the river's length constituted a major perturbation from 1961 to 196“ (King and Ball, 1964) because of erosion and subsequent sedimentation. Presently, however, all the communities in the Red Cedar drainage are equipped with at least secondary A- sewage treatment facilities. Toxic effluents are produced much less frequently in Fowlerville. The Lake Lansing Drain is no longer a major source of enrichment since the treated effluent of the surrounding community, along with that of Okemos and Meridian Township, East Lansing, and Michigan State University enter the stream at a point downstream from the University. A considerable catalog of information exists pertaining to the physical, chemical, and biological characteristics of the Red Cedar River since the intensive investigations of 1959-196A. Physical-chemical and toxicological investigations include Ball, Linton, and Kevern (1968), Stevens (1967), Garton (1968), Jensen (1966) and Talsma (1972). Waybrant (1971) and Vannote (1961) studied nutrient loading. King and Ball (1967), Rawstron (1961), Grzenda and Ball (1968), and Ball, Kevern, and Linton (1969) provide quantitative measures of primary productivity as do Vannote and Ball (1972). King and Ball (1967), King (1962), and Reger (1973) estimated benthic macroinvertebrate standing crop. Linton (196A) and Linton and Ball (1965) estimated the standing crOp and production of fishes in five Red Cedar River loca- tions. King and Ball (1967) and Vannote and Ball (1972) assessed community energetics and productivity. In addition, Linton (1967) described the population dynamics of 5 Red Cedar River rock bass (Ambloplites rupestris) populations and Horton (1969) detailed the species compositions of the fish fauna of the Red Cedar River drainage. 5 The purpose of this investigation was to utilize the considerable background of information compiled on Red Cedar River ecology, along with intensive field investigation, to determine the long-term responses of the Red Cedar River fish community to changes in perturbation and enrichment levels in recent years. Additionally, the response of a warm water stream fish community to organic enrichment and stimulated primary productivity was investigated. SITE DESCRIPTION The Red Cedar River originates at the outlet of Cedar Lake, Marion Township, Livingston County (TlN, R3E) in the south-central portion of Michigan's Lower Peninsula. It flows in a northwesterly direction some 79 km, ending in confluence with the Grand River within the City of Lansing (Figure l). The portion of the watershed investigated com- prises an area of 92,000 ha (355 miZ) which is 57% cropland, 18% forest, 1A% urbanized, and 11% other uses (Waybrant, 1971). The urbanized portion includes the municipalities of Fowlerville, Webberville, Williamston, Okemos, and Haslett and the City of East Lansing and Michigan State University. Two study sites were selected: an upper site, located just downstream of the Dobie Road bridge 3 river km upstream from Okemos, and a lower site, located between the Harrison Road and Kalamazoo Street bridges on the campus of Michigan State University some 10 river km downstream from the upper site (Figure 2). The effects of the urbanized areas of Okemos, Lake Lansing, East Lansing, and Michigan State University on the Red Cedar River fish community were evaluated by this site selection. The stream bottom at the upper site is composed of 23% gravel and rubble, 6A% sand, 10% silt, and 3% logs. Macrophytes (mostly Potemogeton sp.) are present on 18% of the substrate (Reger, 1973) The downstream bottom composition 6 Figure 1. Map of the Red Cedar River watershed. m¥<._ . (Dmo J. 3'1OOIW Id ‘- m._.__>mm._ on. k. .3. ‘31 MW"! )1 HO $70M ‘93 13m mm NVOCI .330 E $1 91» .Pm2<.._.=>> HO Mvn a QM? mozmxo . O, Oz_m2<._ 9?... ad mbGaOEg _v_ 11 o o 00:5 a 1 u u d m m NouuaH @2524: .w 025243 Figure 2. Map of the Red Cedar River showing upper and lower study sites. 10 NC]. N91 N31 II wa—t — I lurw‘Oa-I — B—l x :3 >133. .230 108 so“: . C .. $55.5 i. «ostxw «.r :2m J. 1...! . x! I x. 2; :35. .0100 to. .250; KNEE ¢ 3N and may be considered absent when r 1 7. The standard error of the estimate is computed as 18 (fi-m> (fi-c> S.E.(N) = N A mc(N-l) (Robson and Regier, 1968). In 1972 multiple census mark and recapture population estimates were computed in addition to Peterson and Bailey estimates. The Schnabel method, where A k N: M raw mc/ i=1 i=1 when k = the number of sampling periods is particularly useful since it yields an unbiased estimate when catches, c, are small relative to the population number, N (Robson and Regier, 1968). The standard error of a Schnabel estimate is A A k S.E.(N) = N/ M 6 i=1 (Robson and Regier, 1968) The Schumacher—Eschmeyer method, where k M2? N: M m c/ i=1 i=1 mr is not as sensitive to nonrandom distribution of individuals as the other mark and recapture estimation methods (Hundley, 1954). The standard error of this estimate is k k A N k-l r /c - mr/N S.E.(N) = i=1 i=1 k 2 mr i=1 (McCann and Cruse, 1969) 19 Hundley (1954) found the Schnabel method more reliable when r i 5% N, and the Schumacher-Eschmeyer method better when r > 5% N. Differences between methods were small. All population estimates were computed using a CDC 6500 computer and programs modified from McCann and Cruse (1969)- Standinngrqp Estimates Standing crop estimates for each species were computed as the product of the mean weight of the individuals sampled and the estimated number of that species per unit area of stream (Linton and Ball, 1965). For those species used in age and growth determinations, total standing crop was com— puted as the sum of the standing crop in each age class. Diversity The Shannon-Weiner index of species diversity (Wilhm and Dorris, 1968) and total equitability (MacArthur, 1965) based on numbers of individuals and biomass per unit area (Wilhm, 1968) were computed for each location—year combination using a CDC 6500 computer. The Shannon-Weiner index, al- though the subject of substantial criticism regarding its ecological significance (Hurlbert, 1971), was selected because it is dimensionless and relatively independent of sample size, (Sanders, 1968) allowing comparisons between locations, times, and authors (Mathis, 1968). Since the index is based upon relative abundance, it is not severely affected if rare species are overlooked in the sampling procedure (Reger. 1973). 20 Water Quality Concurrent with the present investigation a detailed study of benthic macroinvertebrate diversity and standing crop (Reger, 1973) and an investigation of water quality and community energetics were performed (Ball and Kevern, 1973). Water quality parameters including alkalinity, hardness, pH, total solids, nitrate nitrogen, total phosphorus, and chlorides were determined on the basis of monthly water samples. Sediments and fish were analyzed for pesticide residues bi- annually by the Pesticide Research Center, Michigan State University. Twenty-four hour continuous monitoring of dis- solved oxygen and temperature was performed once monthly, except during winter months, to determine the magnitude of diurnal temperature and oxygen fluctuations and to estimate primary productivity. Determinations of dissolved and fine particu- late organic matter and 24 hour organic drift were also accomplished. Fish Kill On August 11, 1971, a major fish kill took place at the lower site. Many fishes were seen at the surface and, on subsequent days, numerous dead fish were found in and along the stream. The lethal conditions have been attributed to low dissolved oxygen levels during and immediately after a heavy rainfall following a prolonged dry spell (Talsma, 1972). RESULTS Length-Weight Relationships Log-log length-weight relationships were computed for the species utilized in the age and growth study using study using least—squares procedures (Sokal and Rohlf, 1969). The coefficients of the transformed relationships are presented in Table 3. The appropriate length-weight relationship, along with back-calculated lengths at annulus formation, were used to compute estimated weight at annulus formation (Tables 5-8). Growth Study Mean total length at annulus formation for rock bass, redhorse, white sucker, and smallmouth bass were computed utilizing the appropriate scale radius-total length relation- ship (Table 4). Back-calculated total lengths, utilizing 1972 collections and coefficients, were computed to serve as an independent indicator of replicability for each location- species combination (Figures 3-6). These replicates were not included in testing for differences among mean lengths because of generally smaller sample sizes and the use of different scale radius—total length relationship intercept values (Table 4). The unmeasureable effect of the intercept value could alter the significance level of a test for dif- ferences. Testing was therefore restricted to among location comparisons utilizing only 1971 samples. Paired t tests (Sokal and Rohlf, 1969) were used for all growth comparisons. 21 22 . . emmm.m asaa.mn mama . u oomo.m mmem.e- aama asoasoaoo moaoomoaoaz mmmm.m mo:©.:u u u mama . :aeo.m mmaa.m- mmmo.m maeo.mu aama anomaoEEoo moaoonooeo amaa.m amaa.eu mmme.m .am:a.:- mama aama.m memm.mu mamo.m mamo.mu aama soaanoaao osoomoaoz amo:.m mm:e.mu some.m ommo.eu mama am:o.m aaaa.e- ooom.m mmme.:u aama manomomma mooaamoaoaa c o moa : o moq amom moaooqm pmeoo com aozoq ameoo pom Lona: .malaama amcoapmooa m Soap mocmam ao>am amooo pom mcoammoawoa unwaozlcpmcoq .m manme 23 . u I amm.o amam.m mm.mm mama I u - omm.o ammm.m am.mm aama asoaaoaoo moaoomoaoaz mam. meem.m aa.ea I u . mama mam. sema.m mm.ma amm.o ommo.m mm.am aama anomaoeaoo masoonooeo mam. omam.m mm.ma mam.o aoaa.m :m.am mama amm. ammm.a 03.3: mmm.o ammm.a mm.ma aama soasaepaeo maooooaoa amm.o ammm.a mm.mm mam.o ammm.a mo.am mama mam.o momm.a ma.am aam.o mmmm.a mm.mm aama maaomoosa mooaaaoaosa a p m a o m amow moaooam mmpoo pom aozoa ameoo pom Lemma .mauaama .mcoaomooa m aoam nonnam ao>am ameoo pom mo mucoaommmooo CoapmaoLLOO one mcoammoamoa camcoa aspOBImSHUmm oamom .3 oaome 24 Instantaneous growth rate, G, was computed using the formula loge Yt - loge Yo t where Yt = total length at the end of the time period, t, Y = the total length at the beginning of the time period, 0 t, and t = the time interval between Y0 and Y (Tesch, 1968). t Using t = 1 year, back—calculated lengths at consecutive annuli were substituted for Y0 and Y to compute the specific t growth rate (Figures 3-6). Initial length at time of hatch- ing, necessary for the estimation of C during the first year of growth, was selected for each species. Twenty-five mm was selected for white suckers and redhorse (Carlander, 1969) and 10 mm was selected for rock bass and smallmouth bass. These lengths approximate the initial total length of these species. Since the loge of any value utilized will be subtracted from the loge of the back-calculated length at annulus I, the actual value selected for the initial length is somewhat irrelevant as long as it is consistent for a species and specific growth rate comparisons are restricted or adjusted to that value. 10"3 g was assumed to be the initial weight of all fry for weight increment computations (Quick, 1971). Mean back-calculated length at annulus II for lower Red Cedar rock bass was found to be significantly greater at the 0.05 level than that of rock bass captured at the upper site (Figure 3, Table 5). Among location differences for the sucker species were found to be not significant. 25 Among location comparisons for smallmouth bass were precluded by the paucity of specimens from the lower site; only two were captured in the entire investigation (Tables Al, 2, 3). Comparisons of computed weight at annulus formation indicate that all species investigated gain weight more rapidly at the lower site than at the upper (Tables 5-8). Rock bass at the lower site are a full year's growth in weight ahead of those from the upper site by annulus II (Table 5). While annual increments of length and weight are greater at the lower site throughout the life of the fish (Table 5), the specific growth rate of lower Red Cedar rock bass is very much greater than that of the upper during the first year of life (Figure 3). The growth results obtained for the sucker species support the conclusions of Linton and Ball (1965) who found sucker growth to be highly variable both within and among locations. Differences in redhorse scale radius-total length regression coefficients (Table 4) are probably due to the small number and limited size range of some samples (Linton, 1964). Smallmouth bass growth at the upper site was found to be virtually the same as that computed by Linton (1964) for this area in 1961 and slightly faster than that computed for an enriched section of the river in 1961 by Vannote and Ball (1972). Rock bass growth in 1961 was not significantly different at the two sites sampled during the present study (Linton and Ball, 1965). Both 1961 populations exhibited growth patterns resembling computed growth at the lower site for 1971 (Figure 3). The accelerated growth of lower Red Cedar rock bass in the present study is attributable to rapid growth Figure 3. 26 Back-calculated total length (TL) (mm) at annulus formation 1 l S.E. and instantaneous growth rate (G) of Red Cedar River rock bass at two locations, 1961, 1971, 1972 (1961 data from Linton, 1964). Back-Calculated Total Length (mm) 27 200- 150 d 100- 50-- 1! . / A o ’1 x / I/ ,9: / I / A I J #1 r O [In A I I / A I, I. G, l9?! :3 ' TL,I97I I A TL,I972, Upper x a TL,I972, Lower o TL,I96I, Lower X TL, I96l, Upper ----— Lower \\ Upper 0% .‘x‘ .\ .\\0—O \ \ 1a.. ' o)‘ -2.0 ~1.5 PLO ~05 I l I I I l I I I I II III IV V VI VII VIII Annulus Instantaneous Growth Rate, G Figure 4. 28 Back-calculated total length (TL) (mm) at annulus formation t l S.E. and instantaneous growth rate (G) of upper Red Cedar River smallmouth bass, 1971-72. 29 0 .22... 5390 969.353.: IVVVI Annulus 0‘. 1 4| 0. a a . a o A/ I 2 O l . 1 may / A '9' / I 1 . Lulu .lloiA TGT \ / OOA . vlloll. .‘l‘ — 4 a - O O O W O 0 O 4 3 2 1 AEEV 59.3 .58. 8332.3 -xomm 30 Figure 5. Back-calculated total length (TL) (mm) at annulus formation i 1 S.E. and instantaneous growth rate (G) of Red Cedar River white suckers at 2 locations, 1971-72. 31 0 .22”. £320 maoocmacmams 0. 5 0 5 2 1 4|. 0 _ a 7 _ a m I O . L. L. W w \x G T T U L \ a A _ u o u \\ no \ \ \ \ \ \‘uI. V .. 7m! — a — a 0 0 0 0 0 0 0 0 4 3 2 1 AEEV 5 3:3 .50... “5.23030 .53 IV Annulus Figure 6. 32 Back-calculated total length (TL) (mm) at annulus formation 1 l S.E. and instantaneous growth rate (G) of Red Cedar River golden redhorse at 2 locations, 1971-72. Back-Calculated Total Length (mm) 33 A 4001 I . .2,0 0/ AI/1 1 r / 1F AI 1 300-4 -1.5 A I, , I / I 2001 g - 9.1.71 " 3.0 A - 11,1971 A 11,1972, Upper a TL,1972, Lower 100- "pp" -o.5 --- Lower .\ e\. \ o r r l r r r TLI‘ o I II III IV V VI VII VIII Annulus G Instantaneous Growth Rate, 34 .va aamw uswao3 Hmucoeouuaa o .va measaan um unwaoB woumasuamo m .AEBV :Hmw zuwcoa aspen HousmaouusH c .anoa Ga mcomum>ummno mo Honadz m . .suwcoa HeuOu wounaaoamolxomn come we pound pumpamum N .msascan um Aaav :uwcoa amuOu nonmasoamonumn new: a I I a.mq a.am a.ma m.m~ H.mH m.oa q.N I I H.moa o.aHH m.ma o.mm w.om a.NH <.N I I oo.HN oo.HN mm.ma Nd.a~ mm.mm mm.¢m mm.mm I I N a o «N we «a maa I I oom.m moo.m qu.m Hm¢.N Hom.a 0ao.H com. I I om.o¢a om.maa om.qma ao.oea mm.maa mm.ma mm.ma qaa qq.mm «.moa um3oa 0.0 o.qm m.mm m.¢m o.o~ w.~a a.w N.q o¢.N m.oqa m.oqa m.ooa «.mw m.aq m.aN a.¢H 0.0 mq.m oc.o oo.aa mq.ma mm.ma qq.~m mo.a~ om.o~ mm.ma am.om a CH mm mm «ma owa «mm oom mmwm I oaa.m 0mm.N mam.H omo.a mmm. mmm. Hmm. Noaa. oo.oma oo.oma cm.maa mm.mma ~o.oma ma.woa mo.am aa.oo Hm.o¢ mad am.am o.aaH Head: xa aaa> aa> a> > E aaa aa a noneoz a3 35 3am ousuamu manuamo um msasac< um unwaoz nuwcoq Hench and: and: .aama .mcoaumooa N scam mmmn xoou uo>am umvoo pom Mew mcoaumusano cusouw was om< .m manna 35 .va camw pnwmos ampCoanocH m .awv mzasscm pm unwaoz commasoamo m .AEEV :Hmm gumsoa HopOp aspcoanosH : .cmoe Ca m20apm>aomno mo LonEsz m .zpwsoa ampop poemHSOamonomn some mo aoaao pamocmpm m .msasccm pm AEEV npmcoa ampOp ooumasoamolxomo smog a m.mmm m.mma m.:ma a.mm m.mm mm.m m.:ma m.wa: o.m©m m.mm m.:m mm.w oo.mm oo.o: mm.ma mm.mm ma.mm :am.za a m m a m mqa I ooo.m: moa.mm aom.:a mmo.m m:mo.m oo.amm mm.mam mm.aem mm.:ma mm.mma aam.:m om :m.sm mm.mm a> > >a aaa a a amv asev oaoodeo LonEsz emanamo pm pm camcoq msassc< unmaoB coo: ampoe :moz .aama .mmmo EpsoEaamEm ao>am amooo pom Loads pom mcoapmpSQEoo npzoam new om¢ .o oaome 36 Table 7. Age and growth computations for Red Cedar River white suckers from 2 locations, 1971. Mean TL Mean Wt Annulus at at Number Site Capture Capture I II III IV V Upper 249.50 359.38 24 93.311 179.25 262.58 341.50 483.00 2.2132 7.907 10.683 10.689 - 163 12 12 1o 1 78.004 85.94 83.33 78.92 141.50 8.05 55.8 175.1 387.4 1176.0 8.06 47.8 127.3 212.3 788.6 Lower 155.8 88.93 245 107.46 208.87 295.93 351.00 - 1.356 6.393 12.965 - - 155 38 14 1 - 82.46 101.41 87.06 55.07 - 13.1 108.5 293.2 495.0 - 13.1 95.4 184.7 201.8 — Mean back-calculated total length (mm) at annulus. Standard error of mean back-calculated total length. Number of observations in mean. Incremental total length gain (mm). Calculated weight at annulus (g). Incremental weight gain (g). O‘U‘I-FUNH .va cflmw uswwmz HmuamEmuucH .va moasacw um unwwma wouwanoamo .AEEV :flmw nuwama Houou HmuamEmuoaH .cmwa aw macaumpumwno mo umnasz .Suwama Hmuou woumasoamolxomn amma mo Houum puvaMum .msasaam um AEEV SuwGoH Hmuou wmumaaoamolxomn coo: 37 Fihlflfid'V\€> I I 1 1 w.amm w.HmH 0.8H q.m 1 I 1 I o.mom m.HnH o.o~ «.m I 1 1 I cm.ooa om.HNH ma.~q mm.mm - I u u u N 3 R u n a u u om.mw omm.~ mmm.~ I I u 1 oo.mqm on.qq~ ow.N~H mo.ow Hm o.oq w.Hma H.on m.me m.mHH N.om o.wH ow.q m.amo m.mqo m.mHm «.mqm m.oom o.mm «.mm mw.q oo.m HH.mN mm.mc Ha.m¢ Hm.Ho mm.mm oo.mm com.~m H N m Hm mm we Ha mcqa 1 ooo.¢H qma.m m~m.q anm.c qu.q ch.N ~N¢H.H 00.5mm oo.wmm mm.mmm om.mam mn.oom mq.mo~ o~.omH Hom.um Nma HHH> HH> H> > >H HHH HH H m HGQBSZ UN um msasac< u: cmmz AH cmmz .Hmma .mcofiumuofi N scum mmuonumu cmuaom um>flm umwmo wmm new mcowumu=QEoU susouw cam mw< .w manme 38 during the first two years of life since the specific growth rate computed for upper Red Cedar rock bass equals or exceeds that of the lower population after two years or 80 mm total length are attained (Figure 3). §pecies Composition The species composition and frequency of occurrence for the fish communities sampled are presented in Appendix A. All the species encountered by Linton and Ball (1965) and Vannote and Ball (1972) were found during the present study. These authors did not, however, differentiate minnow and darter species. Horton (1969), in a detailed investigation of species composition, found 8 species at the lower site which were not encountered during the present investigation. These species, which represented a very small portion of the 1962 community, include Umbra limi, Campostoma anomalum, Notrqpis rubellus, fl. spilopterus, fl. stramineus, Rhinichthys atratulus, Lepomis gulosus, and Labidisthyes sicculus. Species present at the upper site in 1962 that were not encountered in 1971 or 1972 include Notemigonus chrysoleucas, Notropis spilopterus, and Eucalia inconstans. These species also accounted for a small proportion of the total 1962 community. No new species were encountered. As in 1962 the common Shiner, Notropis cornutus, and the bluntnose minnow, Pimephales notatus, were the most frequently encountered minnow species (Appendix A). The other cyprinids contributing substantially are the carp, Cyprinius carpio, creek chub, Semotilus atromaculatus, and the chubs of the 39 genus Nocomis (micropogon and biguttata) (Appendix A). Among the percids the blackside darter, Percina maculata, was in 1962 and is presently the most abundant at both sites followed in abundance by the Johnny darter, Etheostoma nigrum (Horton, 1969) (Appendix A). The rainbow darter, Etheostoma caeruleum, was encountered at the upper site in 1971 and 1972 but was not found at the lower site (Appendix A). This species was not captured in the mainstream of the Red Cedar at any point below Okemos in 1962 but was present in Sycamore Creek (Figure l) (Horton, 1969), which joins the Red Cedar some 3 km downstream from the lower site. In general the species composition is similar at the two Red Cedar River sites investigated. The relative abundance of the species present correlates well with the results of Horton (1969), Linton and Ball (1965), and Vannote and Ball (1972), indicating that the species composition has not been greatly altered since 1961—62. The rainbow darter, while present in a tributary of the Red Cedar which has its conflu- ence downstream from the lower site and at the upper site in 1962, 1971, and 1972 is consistently absent at the lower site. Estimates of Population Density and Standing Crop The results of the population density and standing crop estimation computations are presented in Tables 9 and 10. The 1971 pooled values represent weighted means for the two stations sampled at each site. Since Linton and Ball (1965) 40 66 6.66 66 a666 6 6.66 66 666 66366 6a66 66 6.66 6 a666 6 6.a6 6 666 666666 6666666666666 66 6.66 66 66a6 66 6.66 6 a66 66366 6a66 a 6.6a 6 6666 6 6.66 66 666 66666 6 66.6 6v 66 66 6.a6 66 a66 66366 6a66 6 6.66 66 66 6 6.a6 6 666 666666 66666666666 66 6.6 6v 6 6 6.66 6 666 66366 6a66 6 6.66 6v 66 6 6.66 6 666 66666 66 6.66 66 6666 66 6.66 66 6666 66366 6a66 66 6.66 66 a66a 66 6.66 66 6666 666666 6666666666 66 6.66 a6 666 a6 6.6a 6a 6aaa 66366 6a66 66 6.666 66 6a666 6 6.66 66 6a66 66666 66 6.66 6 666 66 6.666 66 6a66 66366 6a66 66 6.666 66 a666 6a 6.a66 66 6666 666666 666666666666 66 6.666 66 666 66 6.a6 66 a666 66366 6a66 6a 6.66a 66 6a66 66 6.a66 66 aa66 66666 N ouo<\mnq N 0604\663832 N ouo<\mnA N .muo<\uona:z coaumum umow AHHEMm N H umwmo pom 60mm: N Havoc com umaoa .6616666 .666666066 6 66 666:3saoo Smfim uo>6m umpmo com 656 m0 .6HHEmw kn .GOHuwmoafioo N van .oocmvasnm Hmofiumasc .aouo wafipQMum .m manna All .m£\wx u mwONH.H x muom\mpcdom 6 .ms\uonaad oqa.o N muom\uonE:zH 666 6.a66 666 a666 666 6.666 666 6666 66366 6a66 666 6.6a6 666 a6666 666 6.666 666 6666 666666 66666 666 6.666 666 6666 666 6.666 666 6a666 66366 6a66 666 6.6666 666 66666 666 6.66a 666 66a6 66666 6 6.a 66 66 6 6.6 6v 6 66366 6a66 6 6.a6 66 66 6 6.6 66 66 666666 666666 666 6 6.66 66 66 6 6.6 6 6 66366 6a66 6 6.6 6v 6 6 6.66 66 66 66666 a 6.66 66 6666 66 6.6 6 66 66366 6a66 6 6.66 66 6666 66 6.6 6 666 666666 66666666 6 6.6 66 66a 6 6.6 66 66 66366 6a66 6 a.66 66 6666 6 6.6 6 666 66666 6.6.6666 6 66666 42 0.00 00 000 0.00 00 000 60300 0000 0.00 0 00 0.6 6 00 000000 000600 0.00 0 06 0.0 0 00 60300 0000 000006000 0.0 0 0 0.0 0 0 00000 0.00 0060 0000 0.00 000 0000 60300 0000 0.06 0000 00000 0.00 0000 00000 000000 60600600 0.6 060 0000 0.60 0000 00000 H0300 0000 00006602 0.00 0006 60000 0.00 000 6000 60000 0.0 00 000 0.0 0 00 60300 0000 0.0 00 000 0.0 00 000 000000 00000008 0.00 00 600 0.0 00 60 60300 0000 08066000: 6.6 6 66 6.66 66 666 66666 0.0 06 000 0.0 0 0 60300 0000 6.00 000 0000 0.0 0 00 000000 0:000uw00 0.0 0 0 0.0 0 00 60300 0000 8000000000: 0.00 000 6000 0.0 0 00 6000: 0.00 006 0000 0.00 600 0006 60300 0000 0.000 0000 00000 0.0 000 000 000000 6060606060 6.00 600 0060 0.6 00 000 60300 0000 05060000: 6.666 6666 6a666 6.66 666 6666 66666 0.00 00 000 6.00 000 0000 00300 0000 «0.00 00 066 0.000 0000 00060 000000 0000605600 0.00 00 000 0.06 0000 00060 00300 0000 0580600600 . . 6000 00 0 0000 0000 00 000 00600 000060 0 60000 000 6000: 00000 000 60300 0006060 6000 0000000 .0010000 .0000u0000 0 60 .000u0 0000000 no 0000000 00 .00Lm00 60>00 ~000U 000 00 0060 00000060 000 000000500 000060806 00u080umm .00 00008 Fir-CH0 COCO NNNO 0000 0000 0000 O‘Q’l‘h O‘QMO OHCI-fi NH \0 “3 m 0000 .6 {00¢ COCO 0000 0000 66000 0006 06000 000 0006 000 6000 OOOO 0000 GOOD \DOMO COCO NOI-IH O O... 0606 r-i OH\D\D 0000 NH 0006 0000 GOOD I-IOHO MONO 0000 QOO‘O 6 0666 0000 0000 6006 0000 6 GHQ 60 66 60 006 6060 00000 00600 06006 066 060 000 HOOD 6 60306 006000 60306 6000: 60306 000000 60306 6000: 60306 000000 60306 60009 60306 006000 60306 6000: 60306 000000 60306 6000: 60306 000000 60306 6000: 6000 0006 6006 6006 0006 6006 0000 6000 0606 6000 0000 6000 0806 0668: 0000006050 0000 000006 x006 0000006000 60060 .000 0050002 000 0060600080660 006060800 0060600 000000600 0.6.0000 00 00600 141$ mam omq mwm mmq moNN m¢a mama nmo «N mm co qwmm mmq Hmm wmm HH Ha Ha Ha mm cad mm when mama comm omom ©OOmH mmmo mnmm QNNQ me NMN we mm: OOOM whmm 0000 00ch ommo 0000 GOOD COCO NwON met-IQ OOOO OOOLO OHHO 0mm mmm mmm 0H¢ mmo cow wee «mm Nmmm wno 0mm mwa OOON 0000 quN omqm mmow mmmm qoqq mHNm mnqc Hood um3oq vmaoom um3oq “mam: uw3oq vmaoom um3oq “mam: umzoq meoom umaoa umam: H0304 vmaoom um3oq umaa: umaoq vmaoom um3oq Hung: “@304 vmaoom HG3OA Hmmaa NN¢H Huma Numa Hnmfl Numa Hmma Nan HN¢H Nm¢a Hmma Nan Hmma .nam mafiaqu mdumaaumaouwwc mfionom mmvfioaamm msumumouoaz HamaEOHov m5u0umouuaz mfiuummmsu mmufiaaoana< .mam mausamuoH A.u.aouv OH magma “5 Hmn\wx mmONH.H xv muum\mnA m Awn\* u oq.o xv muum\uwnE:zN Hex\H u qH~o.o xv mHHe\Hmpa=zH m.HH~ Hmow wmmom H.qom mooq HHHHm “msOH HHmH m.HHo HmNHH oqmoOH n.0mq moqw oaowm meoom H.¢Hm Hmmm Hommm o.~mH aHmOH mmomn uszH HHHH Hmuoe o.HmoH mmmqm qoqmmH w.mHH Hme ommoq “mam: o.o o o o.o o o “maoH NHmH o.o o o H.o H o cmHoom mcmumaooaH o.o o o 0.0 o o pmon HHoH mHHmusm o.o o o H.o N HH “mam: o.o o o 0.0 o o .333 NEH o.o o o m.o m oH umHoom mamowm>mHH o.o o o 0.0 o o umaoH HHHH muumm o.o o o m.o m Hm “mam: H.o qu Ham 0.0 o o HmaoH NHmH H.o OH Ho o.o o o umHoom asmHapmu o.o o o o.o o o “msOH HHaH «acumomnum H.o mH HNH 0.0 o o “mag: ¢.H Hoq mHoN 0.0 o o pmsOH NHmH H.o He mHH H.o H ow cmHoom ausHc H.o e we H.o H HH “maoH HHmH «soumomnum ~.o on mom H.o c me “man: ~.HH mmo~ NHomH «.0 ac wmm umsoH ~H¢H o.H~ ommm ommqm H.o wHH mHOH umHoom mumHsuma H.q mmn oooe H.o mm HoN “msoH HHHH maHoumm q.wm o~oo NHmmo N.H Ham HHHH “mag: H.u.:ouV OH oHan U6 found most assumptions of parametric analysis of variance to be violated when mark and recapture estimates are employed, non-parametric among location comparisons of pooled 1971 estimates were used. The Wilcoxon two sample unpaired obser- vation method for a posteriori comparisons of ranked data (Sokal and Rohlf, 1969) indicates that differences in standing crop are significant at the 0.001 level. Differences in numerical population density are significant at the 0.1 level. The Mann-Whitney U test for two samples of ranked unpaired observations (Sokal and Rohlf, 1969) indicates that differences in standing crop and numerical density among years are not significant at either location. Linton and Ball (1965), utilizing similar tests, found significant differences in numbers but not standing crop among 5 Red Cedar River locations in 1961. The Cyprinidae and Catostomidae are generally the most abundant families although the darters (Percidae) may be equally abundant at the upper site (Table 9, Figure 7). The Centrarchidae at the upper site and Ictaluridae at the lower site are third; the sunfishes are twice as abundant at the upper site as at the lower while the bullheads are more than 10 times as abundant at the lower site (Table 9). These 5 families account for at least 99% of the individuals present in all samples. Total standing crop estimates are highly variable due largely to variation in the estimated abundance of the sucker species (Table 10). While Linton and Ball (1965) and Larimore (1961) discuss the many difficulties encountered 147 Figure 7. Relative numerical and standing crop abundance of the most frequently encountered Red Cedar River fish families at 2 locations, 1971-72. Biomass Numbers 148 maiwx 3 2 8 S 3 :1 I'L |972 l 30 20 10 0 ’1 g; 919L110 IIV s\\\\\\\\\§:§2; up...” . o...o.o.v.v.o.9.v.vovo'.'.'o'o'0 a. 009 I, o o o o o o o o ’.,’of0’02°.‘.°o'< 3.9.03.0...0‘5 7/////. 2:: ’6 .' r///////,;.;.;.;.~;.;. aepunlezol A a 0 ¢ ...0‘- eapguomnueo '0'. 5'0'0'9'0'0'4 N 5202.19.26.30? 99 p g u g 1 d A 3 k\\\\\\\\\\\V:f:3:= V//////A:§3: egpngISOIBO 7///////////fi:o':3:§1‘2§:§t§:§ft 18990 pea Jaddn JBDOO pea JOMO‘] I97| 919U10 IIV .\\‘ '. 'a 'A '0 W‘ gappjgd "as: espluOJBJWOO eepunlezm .\\\\\\\\\\\\\ 99p .u . “Mo x\\\\\\\\\\\\\\\\\\\\\\ ‘2265939 ' ' '//////////////////////////z‘°‘"""’: eepgwozsmao lelol 10 3140‘ “9 when attempting to compare stream standing crop estimates, comparisons may be Justified if restricted to similar loca— tions and estimation procedures. Since Linton and Ball (1965) did not include minnows and darters in their 1961 estimates the data presented in Table 11 have been corrected by subtraction for comparative purposes. The standing crop estimates computed during the present study are generally higher than those of earlier Red Cedar River investigators. Only two total standing crOp estimates computed by Linton and Ball (1965) fall within the range of adjusted 1971-72 values. Vannote and Ball (1972) estimated the total standing crop of Red Cedar fishes in 1961 at 195 lbs acre.l (218.6 kg ha-l) including minnows and darters, which is also considerably lower than the unadjusted estimates computed for 1971-72 (Table 11). The greatest deviations from earlier estimates are in the suckers and rock bass. As in 1961 (Linton and Ball, 1965) great variability exists in the estimated abundance of the various sucker species. Standing crops are, however, generally higher for 1971-72. The standing crop of rock bass at the upper site is consistently greater than any values obtained in 1961 (Linton and Ball, 1965; Vannote and Ball, 1972). In 1964 the standing crop of rock bass was estimated as 65 lbs acre (72.9 kg ha-l) at the upper site (Linton, 1967), which approaches the 65 to 82.3 lbs acre-l (92.2 kg ha—l) computed for 1971—72 (Table 10). The standing crop of rock bass estimated for the lower site in 1971 is about the same as the 1961 estimates (Linton and Ball, 1965) while 50 .Hm:\wx .Qopo wchcmpm Hmqu wmpmsmpm Ho & + mmON.H xv whom\.mnH :H oopo wCHpcmpm * .AOH mHomBO msmppmp cam mzoccHE Oo prHm3 OmmeHpmm mchompppzm an Ompm5nOHm memO pom LOOOE mzu mo COszoHchoo .HH mHomB 51 1972 values are considerably lower (Table 10). In 1962 this species was virtually absent at the lower site (Horton, 1969; Linton, 1967) but was recovering by 196“ (Linton, 1967). The relative abundance of the four sucker species present in the Red Cedar River demonstrates several con- sistent trends (Table 12). The northern hog sucker, Hypentelium nigricans, was virtually absent from all samples taken at the lower site. Only one young-of-the-year was captured in 1971 while none were captured in 1972. This species was completely absent at the lower site in 1962 (Horton, 1969) but in 1959-61 it represented as much as 100% of the sucker biomass at the lower site (Linton and Ball, 1965). The spotted sucker, Minytrema melanops, has apparently always been of minor importance, contributing about 1% to the total standing crop of suckers (Vannote and Ball, 1972; Linton and Ball, 1965; Horton, 1969). Few were captured at either location during the present investigation (Table 12, Figure 8). In 1972 the golden redhorse, Moxostoma erythrurum, was the dominant sucker species at the upper site despite great variability in the estimates (Table 12, Figure 8). This is generally consistent with the findings of Linton and Ball (1965) and Horton (1969), but Vannote and Ball (1972) found the white sucker, Catostomus commersoni, to be the dominant sucker at an enriched location several km upstream from the upper site in 1961. White suckers are generally the most abundant sucker at the lower site even though there were more redhorse present in 1972 (Table 12, Figure 8). 52 .OO\OO n OOOO.H O OO9O\OOHO .EX\% u :Hmw.o x mHHE\%H OOH O.Ou HOH.O OOH O.OOH OOO.H Hmsoe O O.O mOm O O.O O OOOOHHOHO EOHHOOOOOHO m 0.0 OOH Hv 0.0 mm mmmOOHme OEOHOHOHE OHOH mO H.HH OOO.O Hm 0.00 mmm.: Espsszpmpm OEOOxoxoz : m.mH OmH m: O.Om OOH.m HcomHmEEoo OSEOHOOHOO OOH H.O:H OOO.OH OOH O.Hmm OOO.OH Hmsoe O O.Om HHO.H Hv H.O HH OOOOHLOHO EOHHOOcmmHm m O.O OOH H H.H OOH mmosOHOe OEOHHHOHH HHOH HO O.OOO OOO.HH O 0.0 OOO SOLOHOHHHO OEOOmOHox O 0.0H OOH OO O.OHO OHH.HH Hcomsmesoo mOsOpmosmO R opom\mpH OHHE\O O whom\mpH HmHHE\m mmHoQO Hmmw Homo: sozoq .meHOOH .mCOHHOOOH m pm mmnHEoumOHOO Hm>Hm HmUmO pom esp mo COHpHmanoo mmHommw .mH meme Figure 8. Species composition of the Red Cedar River Catostomidae at 2 locations 1961, 1962, 1971, 1972 (1961 data from Linton and Ball, 1965; 1962 data from Horton, 1969). 511 2.3 2.3 «2: 6-33 I I O . In I I I TON f T .- 10' I 1 T I: l W I IO. r on. I I T T I T Io. O 1. l I IO. 1 OO— ia.a.....>uo . utoclzl 3““ 2.001.... '50:...01 y// 0505;}: .4045 3.3.0.309»... _..u.........u............w V\\k I. 19 p93 pea JSMO'I spguouong lilo; Jepeo pau Jeddn 55 The estimated standing crop of smallmouth bass at the upper site compares favorably with the earlier estimates of Linton and Ball (1965) of 5.6 lbs acre.l (6.3 kg ha—l) using the Peterson method and 11.0 lbs acre"1 (12.3 kg ha-l) using rotenone. Vannote and Ball (1972) estimated 18.6 lbs acre-1 (20.8 kg ha-l). Smallmouth bass were virtually absent at the lower site in 1971, 1972, and 1961 (Linton and Ball, 1965). Horton (1969) found up to 2.8% of the specimens in an electrofishing sample captured in the vicinity of the lower site to be composed of smallmouth bass. Largemouth bass (Micropterus salmoides) young-of-the-year were captured in small quantities at both locations during the present investigation but no adults were captured (Table 10; Appendix A). Northern pike, Esox lucius, were found at both sites in 1971 and 1972 but were generally more abundant at the upper site (Table 10). This species was absent at the lower site in 1962 (Horton, 1969) and in the enriched zone investigated by Vannote and Ball (1972). Linton and Ball (1965) found several specimens at the upper site in 1961. The total contribution of the major game species present in 1971-72, smallmouth bass and northern pike, combined represent 2-12% of the total standing crop at the upper site and only 0.3-2% at the lower site. Vannote and Ball (1972) estimated the contribution of smallmouth bass in 1961 in an enriched zone as 9.5%. Diversity Diversity and equity indices computed using 1971 and 1972 population density and standing crop estimates and the relative abundance of fishes captured in 1962 by Horton (1969) are presented in Table 13. Standing crop diversity exhibits some variation at the upper site as does numerical diversity at the lower site. No consistent trends are apparent; d values do not exhibit the downstream increase or positive response to enrichment observed by Cole (1973), Harrel (1967), Sheldon (1968), and Whiteside and McNatt (1972). 57 .AOOOHV coupon Eosm mumm* I mmm. I m:.m *NOOH mmm. FOO. HO.m om.m NOOH Lozoq mHO. mzm. HO.m OO.m HOOH I mmm. I :O.m *mOOH OOH. HOO. Om.m OO.m NHOH HOOOO mam. Omm. OO.m OH.m HOOH mono wcHocmpm Hpstm mHmDESZ HpHsvm QOHO wchcmpm O mpmoasz m Hmmw mpHm .HOOOH .coOsom SOHO OOOO mOOHO OOOH .HHOH .OOOH .OOOHOOOOH O OO OHHCOEEoo gmHO Lm>Hm Hmvmo Omm 63p mo HOHHHOOHHSOO new Amv OprHm>HO mmHomom .MH mHomB DISCUSSION Natural woodland streams are heterotrophic systems which utilize autumnal leaf litter as a trophic base (Minshall, 1967; Cummins, 1972). A complex decomposer- detritivore interaction is essential for the incorporation of these allochthonous materials into the community food web (Cummins, 1972). Deforestation, intensified agriculture, and urbanization tend to reduce autumnal leaf fall, stimulate autochthonous primary productivity, and produce large quanti- ties of allochthonous dissolved and fine particulate organic material (Cummins, 1972) (Figure 9). This sequence of events leads to a subsequent breakdown of the detritus-oriented food web. Although the heterotrophic state produced by this temporal series receives organic inputs which are similar to those of the original heterotrophic woodland stream, the large particle detritus input necessary as substrate material for efficient assimilation of organic inputs is absent (Cummins, 1972). This temporal series is represented by longitudinal succession in the Red Cedar River and is charac- terized in Table 15. Tributary headwaters represent original heterotrophic woodland streams. Upstream agricultural develop- ment and small ammounts of urbanization (Figure 9) have caused the Red Cedar at the upper site to be autotrophic although considerable large particle terrestrial input is 58 59 Figure 9. Hypothetical temporal-longitudinal stream succession. A "natural" woodland stream with relatively stable water levels and low sediment loads is degraded to an autotrophic stage characterized by high nutrient levels, sediment loads, and primary productivity and greater dis— charge variability because of deforesta- tion and intensive agriculture. Further degradation results in a heterotrophic state based on organic wastes where urban runoff and resultant storm drain overflow along with water removal, cause severe water level fluctuations and high sediment loads. 1 - complex, heterotrophic, shade adapted, cool water system charac- teristic of Red Cedar River tributary headwaters; 2 - simplified, autotrophic, light adapted, warm water system repre- sented by the Red Cedar River in the vicinity of the upper site; 3 - simplified, heterotrophic, warm water system repre- sented by the Red Cedar River at the lower site. Redrawn from Cummins (1972). 60 Funded Idldrohdd low nght 1 % .’lni"-. . ___‘ Clo-r Wold! 8H... .Ioond '00" laHl-l Hoidrdfirophic $9.00 Agricoliovdl lads-var .fl.‘ “.‘. Doiorolidd Idldrlhdd .N‘“-' . . - aim-‘5‘“ 3w . ‘ J ,I‘ I? 1‘ -' Agile-"Ital .I Ida." ”I hubld 'C'Ol In... Ioducod Ground '00.: Intotnddldu Autoirophlc Stat. Urbonlldd Vanni-dd “H A III-I e— an Glamor, ...-. _ Hi.” I7 Lula,” _ a: fill High tlih! WWW-V lndulfrldl and 3 "Poo-dunno I 9001:11: In". Vuy lurbld Sud-III CFO-Hy loddcod .ndud Inter ‘ Urbdnlzod Hot-renophlc $9.9. 61 present (Table 1”). An urbanized heterotrophic-autotrophic state (Cummins, 1972) characterized by high primary produc— tivity, high levels of fine particulate and dissolved organic input, and low levels of large particle terrestrial organic input is represented by the Red Cedar River at the lower site (Table 14). Reger (1973) characterized the macrobenthic inverte- brate fauna at the upper site as consisting of many amphipods, hydropsychids, elmids, pelycopods, and chironomids of the tribe chironomini. The lower site is characteristically inhabited by naidid and tubificid oligochaetes, planarians, and chironomini. Large detritivorous invertebrates, such as the amphipod Gammarus fasciatus and the crayfish Orconectes propinquis (Minshall, 1967), are abundant at the upper site (Reger, 1973). The large detritivorous invertebrates are represented at the lower site by few specimens of Q. propinquis and the isopod Asellus militaris (Reger, 1973). The Red Cedar River, then, demonstrates through longitudinal succes- sion the temporal succession characteristic of many streams; large detritivores, abundant at the upper site, have been replaced by smaller benthic macroinvertebrates in the absence of large particle detritus input at the lower site (Figure 9). Vannote and Ball (1972) have emphasized the importance of large invertebrates in the support of Red Cedar River smallmouth bass and rock bass populations. They concluded that although extensive fish forage exists, this resource is largely unexploited. Smallmouth bass and, to a lesser 62 mmI E Hmo m , HI - ml H r .OmLSmmoE Hos eoHposnosq mpmzaomomz .msHm> OmmeHpmm * OOH.OHN OOO.OO Hausa: OHOOOHO Om>HommHO OO0.00 OO0.0H Hausa: OHOOOHO OOOHOOHOHOO Och mmm NON HzacH HmHHpmmHHmB mHOHuHmm mmpmq O0.0 HH.H OHOOm O\m O.m H.m COHHOHHQmmm HHHQ:EEQO m.m 0.0 mHmmnpczmoponm mmOHO HHH.OOO O0.0 HOH.OOO H0.0 OOHHOzOOOO OHOEHOO Omz Hmsoe HOO.mH H0.0 *Hmm.mn *H0.0 coHposeosm mngaohomz Hmz HOO.mOm O0.0 HOO.OOH O0.0 coHpozuosm coOHnOHsmm Omz pmpoo com pmzoq Hmpmo pom HAHQO .HmOmHO cpm>wx new HHmm Eopm .HIOOO mIE Om CH .UOHOOHOCH open: pamoxm ammsHm> HH< .mOIHOOH .mQOHHOOOH m pm Hm>Hm Hmpmo mom opp pom umwwso mmpmcm CO .:H mHnt 63 .mmummz OHHOmEoO Eopm mSHOLQmOQQ ch :mmopuHc mo mHm>mH szm .mpSQCH mmmzmm Eopm mpcmEHOmm OHcmmHo mEom .muomnopo 20H» Ioshpmcoo OmwmcmEmHE an Ommsmo QOHmOHm Eopm mOOOH psoEHOmm anm .pcmEOczogeH oEOO Ocm .pszH OmmmmmocH .Hmpms Ocsopw Omoswmn memme Eopm Hmanm .mHQosm Loam; Ocsopw Omoswmh memmHO .Hmccmno Emmpum mo COHH ImpmpHm HOOHOHQQ 5038 new mmocsm smog: on mzu mHompmcz chmHm .OOOCSH HmpsuHSOHpmm Eopm mngSQmon Ocm cmwoauH: mo mHm>mH anm .Ho>mmpcm HOLSHHSOHHMO HQ Ommsmo COHm loam Eopm mOdOH pcmEHpmm HmcmHm .COHpmemcma pzmHH OmwmmpocH Ocm scum: Ocsopw noosc Imp Eopm Hmcme .OmSmHmpmz HOLSH IHSOHHmm mo moHpmH upmpompmco MCHOHon Hmpmz Loom .Hmpmz Ocsopm Omoswmh 0p mzv mempch .mzhozamozm Oam cowOHpH: mo mHm>mH 30H .HHmz HHow mOHOg Omnmmopmz Umpmmpoa mumpwz pmmHo Ocm OOOH pcmEHOmm 30H .mHo>mH Hmumz Ocsopw an5 com Emmhpm Ompmnm mo mmsmo Imp 30H mHo>HpmHmm .mOOHHoa mac :H Hm>mH Hmpma mchchmE mHm>mH nouns Ocsopm OOH: .Hmsmz OOHOO Omnmpmpmz Ompmmpom mmHQmpm sz>HpmHmm mHCmHHpsz mpcmEHOom manpmHmQEmB mEHmmm onm OmNHchHO Ompmmpomoo Omuooz Oocmpmpmz Ampmpm Ampmpm Ampmum OHpmHHmpomezo OHnQoppopmpmm OHEQOHHOHSO OHQQoppopmpom OmNHcmnHDV mpmHOmEHmszv HmeHch HmOmO Omm pmzoq HmOmO Omm Hogan msmumznmmm .COHmOOOOSm HmOHHOSHOQOS mo mommpm m :H Hm>Hm Hmpoo Omm mzp mo OOHpmHHmpompwno .mH mHomB u. 6 .mmpwpnmppm>cH wwHwH Sou mpm> .meHmewE 0chwpo msococp IQoOHHw mpwHSOHprQ mcHO ch Oo>HommHO co Ummwn HHHQOEEOO OOHHHHOEHO HHOOOHO .Empmzm OHQQopuopopmz mHv .pcoEOCSOQEH mEom ch mHo>mH pcmepsc ch pcwHH OOH: OH OOO OOH; HHO> .mmpmws OHpmmEov Eopw OOOOOH OOO OOHH sopa 30H mHHwOHpHpo zHHwCOHmwooo .ome mzoszmsw OHnQopp Iopmuon zcwz .mnaopp0psw mp COHpospopQ ch COprHHQmmH Eopm OQOHO stuosHm chszp oOHB .mmpmwz OHummEOO Eopm meHHmqu Ochmpo Om>HowwHO ch mpstOHH lawn wch mo mHm>mH OOHm .OOOOH mOpan ImO Omoswmp memeO .mmpwpnmupo>:H mmpwH Hosea .mLQQHpOpsw Iconpnoopsw co Ommwn mpchs IEOO OOHOHHOEHO .Empmmm OHQQopuousw mHA .mHm>mH HemHHpsc new panH OmmwmpocH 0p mzp gmHn mpm> .30H mHHwOHpHHo Hm>mz .mcooppOHSw mp COHposcoua OOOHHOOO OOH: ocw COprHHQmmH HwCHSQ Iooc SOHO OCOHuwSH Iong HwCLSHU ®UH3 mchHSOm meszSOHwa ch OH» ImmEov Eonm pcmEQOHH Icm OOHHEHH .OOOOH mapHHme noosnmm .mOpwsompHo>cH mwpwH mew: .psa IcH mszHme mch Immoopa mHucmHOHm Imm mo mHowao mpHQSEEoo mHowpm nmeQEoo .mmpm>HQ .Empmzm OHQQOHH Iopmpm: mHv .mHo>6H pzmHHCZO pew pamHHpsz 30H 0» @3O 30H Hm> .30H mHHwOHquo Lm>mz .mCOprzuosz HwCHsHO HHwEm .HHOHHHH OwOH HwHHpmmLHmp mHoHuLwQ mwpva OOHHHOOO HHOmoz mwpwzpmppo>CHopowE OHepcmm OHuwm m\m HpH>Hposvopm OHwEHHm cowhxo Um>H0mmHQ pjocH OHewmso H.O.coov OH OHOOH 65 .mmwp canoe IHHwEm oz .OSOCCHE ch nmpwmzHHzn .mpmxosm mo macho wcHchpm szs mHmpmeOoE .m>Hpo:OopQ IQ: mHHmecmO .mHHHx anw HwCOHmwooo on can stO mo mpwpm pcwpm Icoo w 2H mH £0H£3 mmHoQO pcwpmHOHQH mHSp ImeQEmp ann ch ..O.Q on .OHHO OcHxOmH HOHCSEEoo prwpwco .mpmnESC mpwmeoE :H Hammond mmHoQO pcwHOHOHCH .O.Q 30H ch pHHm .COprH Isaoa mmwp cusoeHHwEm wcHOSOOHQmH OHHwaspwc .mchprmzmImHmm w ch mmeOHn BOCCHE pew .meosm .mmwo Hoop OOH: HO OONH Inmpowpwno OHHQSEEOO mewpm mHmuwmeoz .mmHooam pcwpmHOHCH .O.Q 30H ch .oHSHwHwQEmp OOHO .OHHO Hews mo wchmHmcoo OHHQSEEOO mHowpm anm H.O.coov OH OHOOH 66 extent, rock bass were found to feed extensively on crayfish and other large invertebrates. Since 1961, primary productivity decreased from 1156 g cal m—2 day"1 to 509.4 g cal m-2 day"1 at the upper site and from 768 g cal m-2 day-1 to 655.1 g cal m-2 day-1 at the lower site (King, 196“) (Table 1“). The standing crop of fishes, in particular that of rock bass, increased during this time period at the upper site and at the lower site prior to the August 11, 1971 fish kill (Table 10). Increased autochthonous primary productivity, then, is not reflected in increased standing crop at the primary and secondary consumer level. Greater fish (Table 10) and benthic macro- invertebrate (Reger, 1973) standing crops are present at the upper site which, although strongly autotrophic, has less net autotrophic production than the lower site (Table 1“). Estimates of fish standing crop for 1971-72 are also greater than those computed for the same sites in 1961 when primary productivity was greater (King, 196”; Linton and Ball, 1965). Clearly then, stimulating autochthonous primary production either through nutrient enrichment, increased light penetra- tion, or both (Figure 9) does not generate increased fish standing crop in this warm water stream ecosystem. Rather, the increased rock bass biomass and generally high total fish standing crop at the upper site can probably be attributed to relatively high levels of large particle terrestrial input (Table 1A) and the associated presence of many large benthic macroinvertebrates (Reger, 1973). The lower fish biomass at the lower site, along with reduced quantities of 67 large particle terrestrial input, heavy loads of dissolved and fine particulate organic material (Table 14) and the presence of few large invertebrates (Reger, 1973) indicates that reduced fish standing crops may be due to a breakdown of the heterotrophic detritivore community. Longitudinal succession in streams usually results in a downstream increase in the number of fish species and computed species diversity (Cole, 1973; Whiteside and McNatt, 1972; Sheldon, 1968). Species richness, however, is dependent upon sampling efficiency; rare species, which might be overlooked through experimental error, are weighted equally with the most abundant species when species richness is used as a measure of succession. Diversity indices, such as a, are employed to weigh the contribution of each species by its relative abundance. Neither quantitative method, however, adequately describes the type of succession demonstrated by the Red Cedar River. Here, the number of species present (Appendix A) and the computed diversity index (Table 13) are nearly equal at the two sites although the non-parametric tests of rank clearly illustrate a significant difference in the relative contribution of each species to the total fish abundance (Table 10). These differences in rank are probably due to the physical-chemical characteristics of each site. Although gradient and discharge are greater at the lower site than at the upper (Table l) the effects of flooding and siltation are more severe at the lower due to the presence of several small impoundments in Okemos and on the Michigan State University campus (Figure 2). 68 Starrett (1951) found the most successful minnows in a prairie stream prone to frequent spring flooding and silta- tion to be those which had either a prolonged spawning period or one which commenced late in the growing season. Increased sediment loads during spring floods, which scour the benthic community (King and Ball, 196”), have the effect of reducing the amount of food available to emerging larval fishes and high water levels may wash some eggs and larvae downstream (Starrett, 1951). This situation is demonstrated by the Red Cedar River Cyprinidae. Those minnows which are most suc- cessful at the lower site, where water levels fluctuate greatly (Table l) and silt loads are high, are those which can avoid exposing eggs and young to the detrimental effects of these conditions. The common shiner and bluntnose minnow, the most abundant cyprinids at both locations, spawn after the usual spring flood period; the common shiner spawns from late June through July and the bluntnose minnow throughout the summer (Starrett, 1951). Additionally, both species behaviorally protect their eggs from silt deposition, the common shiner by fanning with the pectoral fins and the blunt- nose minnow by depositing its eggs on the underside of sub- mersed stones and debris (Starrett, 1951). Those cyprinids which are moderately successful at both sites, including the chubs Semotilus atromaculatus and Nocomis spp., construct nests in stream areas of considerable current (Trautmann, 1957) but frequently leave the stream proper, along with carp and goldfish, to spawn in backwaters and tributaries where the effects of flooding and siltation are not so 69 severe (Starrett, 1951). Those minnows which are only slightly successful at the upper site and rare at the lower, including the rosy face shiner (Notropis rubellus) and the mimic shiner (Notropis volucellus) broadcast their eggs over gravel (N. rubellus) or weed beds (N. volucellus) (Carlander, 1969). A similar relationship between egg protection and rela- tive abundance exists among the darter species. The rainbow darter, which has been consistently absent at the lower site while present at the upper, broadcasts its eggs over gravel (Winn, 1958). The johnny darter, abundant at the upper site and present in small quantities at the lower, protects its eggs by adhering them to the underside of stones (Winn, 1958). The blackside darter, abundant at both locations, can utilize the nests of other fish species (Hoover and Cooper, 1936). Similarly, the most abundant ictalurid at the lower site is the yellow bullhead (Ictalurus natalis) followed by the brown bullhead (I. nebulosus) and black bullhead (I. melas). Lower Red Cedar yellow bullheads captured in late July and August were in pre—spawning condition indicating that this species avoids adversely high spring water levels by spawn— ing late in the growing season. Black and brown bullheads, however, are spring spawners (Carlander, 1969). All bullheads build and guard nests (Trautmann, 1957). Hog suckers, characteristically inhabitants of the upper Red Cedar River and rare in the lower reaches, are also silt intolerant (Trautmann, 1957). 70 The reproductive success of Red Cedar River smallmouth bass, along with other centrarchids, has been closely cor- related with spring water levels (Vannote and Ball, 1972). Spawning is temperature stimulated but may be deterred by high discharge (Vannote and Ball, 1972). The limited success of this important game species in the lower Red Cedar River can probably be attributed to a lack of large invertebrates as well as high spring water levels, high sediment loads, occasional low dissolved oxygen levels, and the lack of suitable spawning substrate; the preferred gravel and rubble areas comprise only 8% of the bottom area (Reger, 1973). Maintenance of a non—reproductive population is unlikely due to the distance to the upstream, self-sustaining popu- lations which, in 1961, appeared not to exist downstream of Okemos (Figure l) (Horton, 1965). Additionally, stream fishes seldom actively move downstream (Hynes, 1969). The highly variable year class strength exhibited by rock bass at both locations (Appendix A) and the virtual disappearance of this species during the highway construction of 1961-6“ (Horton, 1965; Linton, 1967) indicate that rock bass repro- duction is also affected adversely by spring flooding and high sediment loads. Reproductive success among Red Cedar River fishes, then appears regulated by the amount of spatial, temporal, and behavioral protection each species offers its progeny. Although diurnal dissolved oxygen levels observed at the two Red Cedar River locations are generally similar (Table 2) occasional low levels, caused by high BOD inputs from combined 71 sewer and stormdrain overflow (Talsma, 1972) and respiratory oxygen demand at times of low light, are characteristic of the urbanized lower site. The fish community at this site, then, is highly unstable due to periodic oxygen sags and subsequently high mortality such as that which occurred on August 11, 1971. The makeup of this fish community at any time is dependent on the severity of the last depletion, the length of time between major depletions, and the rate of recolonization. The composition of the 1972 lower site community indicates that post fish-kill recovery occurs rapidly. Although the age structure of the rock bass, white sucker, and redhorse populations were altered severely (Appendix A), all were present in 1972 and only minor changes in species composition were evident (Appendix A; Table 10). Repopulation by younger individuals is indicated by the absence of older rock bass, redhorse, and white suckers at the lower site in 1972. This is consistent with the findings of Engstrom-Heg and Loeb (1971), who found repopulation of a rotenone treated stream to be by smaller individuals than had been present earlier, and Larimore, et a1. (1959) who noted the presence of many young individuals in a small warm water stream one year after a severe drought. Unlike the lower site the fish community at the upper site has been relatively stable in the last ten years and is characterized by the presence of extensive rock bass populations with con— siderable longevity, 3 darter species, A sucker species, several silt-intolerant minnow species, a limited number of northern pike, and a reproducing population of smallmouth 72 bass. The relative stability of this community suggests that it represents a steady state toward which fish communi- ties in other enriched areas, like the lower Red Cedar, will tend in the absence of specific perturbations. In general the communities sampled were found to be similar. Differences in the relative abundance of the species present at each site appear to be related to the physical- chemical attributes of the eutrophication process. The con— clusion of Larkin and Northcote (1969) that "The many changes involved in the process of eutrophication may have their greatest effect on fish populations by influencing survival of the young" is supported when the effects of flooding and siltation are assessed. Concurrent breakdown of the complex detritivore oriented food web, essential for maximum support of primary and secondary consumers in the stream ecosystem, is the probable reason that the fish community does not respond positively to the increased primary productivity associated with enrichment and urbanization. The community at the upper site may represent a steady state towards which the community at other Red Cedar River locations will tend in the absence of specific perturbations, such as the fish kill of August 11, 1971. SUMMARY AND CONCLUSIONS Growth, as revealed by back-calculated lengths and weights at annulus formation, is generally faster for lower Red Cedar fishes than for upper. Sucker growth is highly variable at both locations while computed length at annulus II is significantly different among locations for rock bass. Species composition differences between the sites are slight. The most notable difference is the complete absence of the rainbow darter at the lower site concurrent with consistently high levels of abundance at the upper site. Non-parametric tests of rank indicate that the relative con- tribution of each species to the total abundance and standing crop differs significantly among locations. Species diver- sity, as measured by the Shannon-Weiner index, 5, is similar for all locations and years. Fish standing crop estimates computed for both Red Cedar River study locations are greater than any made in previous years by other authors. Although standing crop at the upper site is generally higher than at the lower, considerable variability, due largely to the unpredictable abundance of catostomids, exists in the estimates. Rock bass biomass has increased substantially at both locations since earlier investigations, approaching 80 lbs acre"l (89.7 kg ha-l) at the upper site and 25 lbs acre.l (28.0 73 74 kg ha-l) at the lower exclusive of age 0 individuals. Game fish, including smallmouth bass and northern pike, represent 2—12% of the total fish biomass at the upper site and only O.3-2% at the lower. Fish standing crop, representing secondary productivity in the warm water stream, is not elevated by the increased primary productivity resulting from enrichment. It appears that autochthonous materials contribute a relatively small proportion of the energy utilized in supporting the fish community of this stream. The loss of large particle ter— restrial organic input and subsequent breakdown of the detritus- oriented food web of a heterotrophic stream resulting from urbanization, along with other physical and chemical perturba- tions, appear to be causitive factors limiting the produc- tion and distribution of warm water fishes. The distribution of species at the upper site, which has exnibited relatively high stability in the last 10 years compared to the lower, is suggested as a possible steady-state community for these enriched reaches of the Red Cedar River. Occasional low dissolved oxygen levels and other perturbations, such as the fish-kill of August 11, 1971, appear to prevent the distribution of species at the lower site from ever reaching this equilibrium. Results of 1971-72, as well as those of earlier, similar investigations, indicate that the process of urbanization involves complex interaction of physical, chemical, and biological effects which are reflected in the fish community structure of the Red Cedar River. Streambed alterations, 75 watershed deforestation, and reduced ground water levels are secondary characteristics of urban growth which alter the natural trophic structure of the stream ecosystem and reduce its ability to absorb the effects of subsequent organic and nutrient enrichment. The biomass increase realized at the upper site, especially the increased standing crop of rock bass, and the consistently high relative abundance of such desirable species as small— mouth bass and northern pike indicate that the detrimental effects associated with urbanization are not part of an irreversible process. Rehabilitation of the warm water stream, as demonstrated by the upper Red Cedar River, is possible through conscientious management practices and effective sewage treatment. Results indicate that while the lower Red Cedar has generally improved since 1961 the improvement is far short of what is possible based on upstream responses. Formulation of effective stream management policies would be greatly simplified if the effects of the many interacting factors which alter stream fish communities were being under— stood. LITERATURE CITED Bailey, N. J. J. 1951. On estimating the size of mobile populations from recapture data. Biometrika, 38:293--~ 306. Ball, R. C., N. R. Kevern, and K. J. Linton. 1969. The Red Cedar River Report 11. Bioecology. Pub. Mus. Mich. St. Univ., Biological Series A(A): 107-157. Ball, R. C., and N. R. Kevern. 1973. An ecological evalua- tion of stream eutrophication. Mich. St. Univ. Inst. Wat. Res. Tech. Rept. 36. Ball, R. C., K. J. Linton, and N. R. Kevern. 1968. 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APPENDIX 81 O.HON I O O H O Oma m.NNm Om.mm O OO :H m OOO 0.000 mm.ma: m OOH mm m OOH 0.00 O0.00 m mam ON H OOH 0.0 NN.HOO m ONO HO O OOH I I I OmHH mmH AHwHOOV HcomHmEEoo OSEOHOOHwO 0.00H I m om O mmocmeE wEOHeNeHz I I H H H mchHHch EsHprcmmmm I I I I H EusEosw wEoumomeO I I I I Nm mspwpoc mmezmmEHm H.m mm.HOO H OOO HO HOOHOOOO OOOHOHOOOO OOOOO m.N I H H H mSOszw msHmwwth O.N I H N N .mm OHEOOoz 0.0 I m ONm om mapscpoo mHmdppoz 0.00 I H m m wschHHOEw xowm 0.000 I m NH O OOHosH xomm vapcmez .m.m mama mpwEHpmm OmpmpczoOCM mmHomam :wmz prezz .HNOH .OIO pmsw5< .Opr 0mm LOBOH nCOprpm Loam: map pw OmpomHHoo mmHoQO .HI< mHowB 82 O.H 0.0: O.H O :TO mmo MNKO (I) O0.0 m:.O OOHOOOHOHOOz OOHHOO OOH: OOOOOOOO u comhmpmm OOHOHOoECO u :oprHmEOQm powHHQ u INC‘JNNMI—lm (\l (\l INCUr—l OOH mNm OOH ON OHH HOH r r-‘IKMWW H m m mm OO O HO HOOHOOOO H m N O HH om mO OOH AHwHOOV N mO OO AOmHooav H mH mm Om HHOOOOO C‘ [.1 Q E [11 (1) pwEHumm mo mameH mcwpmcoocH wHHwowm memomm>wHO wopwm ESHOHQ wEOOOOOLHM wuwHSOwE wCHonm msHHmchd OHEOQOH mamonnmw mHEommH OwOHonprcmO Hmcpo mw< mw< mw< mw< mwa wwd Om< r_‘| (\l m: LOKON mHHmeMdH mmpHHQOHOE< mewE mzdewHOH mHkuwc mzthwHOH vaHaszHOH 83 I I I I H OwHOE OOHSHwHOH I I I I Om OOOOHSQO: OzawHwHOH I I I I Om OHprwc mzpusHOH 0.0 I O OOH NO HOOHOOOO OOOHOOHOOOH O.OOH I H H H O OOO O.H: I H O O m OOO 0.00 HO.OH O O: NO H OOO I I I m: om AkuoBv ESHOHLHOHO w80pmoxoz 0.00N I O m H m OOO O.OO O0.0m. m OOH OO H OOO 0.0 OO.HOO m OmOH OO O OOO I I I OOmH OOH AproBV HCOOHOEEOO m550pw0pwo H.Om I H N N wmocwHOE wEmeHcHz O.H I H H H mchHHch EsHHmpcmmzm 0.0mO I H O O OHQHwO OSHCHLQOO 0.0 I O OOOO OHH OOOOOOO OOHOOHOEHO H.m I H Om Om .mm mHEoooz 0.0H I H O O mspstowEoppw msHHuoEmm 0.0 HH.NOHH m OHNH HOH OOOOOOOO OHmmOOOz vauanmz .m.m OQ>B OpwEHpmm Ompmpcsoocm OOHowam :wmz amnesz .HNOH nOlm pm3w3< .mpHm LwOmO Omm Hmon aCOprpm HOBOH map pw OmpomHHoo mmHoQO .mI< mHan 84 OOHOOOHOHOoz HOHHOO OOH; 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O OOH H OOH OHLOOOOHL OOHHHQOHQEH OOOHoEHOO OSLOOOOOOHS OOH OOH OOH OOH OOH HOHHEOHOU OHOOWWOLOHE OI—i—‘J‘LOKO ‘ OHOOHOOOE OHEOQOH OSHHOQOHO OHEOQOH mzmonnHO OHEommm OSLHQOOLHOOE OHEOQOH OHOOHHQOC OHLHHOOOH OHHOOOQ OSLHHOOOH OOOHLdHOpOH H.O.coov OIH OHOOO \ 95 O.HOO LOOOELOOO I pOz0OESSOm HOQOQQOO coHuOOHOHuoz OOHHOO cqu compOpOm cothpOm OOHmeoECO COHOOLOESCO OOOLHQ O HO m mOm O OOmH II II H (\l m: Ln COOS UOHOOQ * ”OOOOHQEO OOOEHOOO mo OOOOH ESOHHLOOO OEOHOOOSOO ESLOH: OEOpmongm OpOHSUOE OcHohOm mdpmHfiomfiopOHc meofiom H.O.zoOO OIH OHOOO HICHIGQN STQTE UNIV LIBRQRIES 1293103583989