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' .u/ p ro.u . _ . . la. 0 - ~ 7 o I.’ O I (’1'. u . . . . a 0.1;. . . .. ... o o 2le . . .. r . . . .v .. .u. .u ..o . . .u. l. A . . a. , tr! .3... . r . . . .o.. .r o —n. I J.‘ u _ . 4 1r,r 4 . ‘ . . -p t . _f o. _ l D . . c . .v I .l a , r , a . . I . r . . I . "o . ... . . . 1 u . o... . v .3 . ., O- v D . . , -. “7- vv --— wv‘w - _ “-‘-«-‘.-¢--,. aunv-c -~ ~ 0... ‘ '°‘-'~."‘-.o- on . ‘fioya4. c BENTHIC MACROINVERTE‘BRATE DIVERSITY IN THREE DIFFERLENT‘IA. PERTURBED MICHIGAN STR - n 3.... ~ _ . 3. . F m: 1; RSSIS M II 3m: UNIVERSITY :3 fl Ln 1 fl ICHI a flu U m REGER ‘ 1973 I I! III IIIZIIIIILIIIII I II I I III III “III I ~ ABSTRACT BENTHIC MACROINVERTEBRATE DIVERSITY IN THREE DIFFERENTIALLY PERTURBED MICHIGAN STREAMS By Scott Jon Reger The macroinvertebrate community structures of three Michigan streams were examined through an annual cycle. Study sites were located above and below known sources of human disturbance on each stream. Tax— onomic composition, standing crops, and diversity of both numbers of individuals and biomass were used in an attempt to describe the effects of cultural development on the streams. Macroinvertebrate diversity indices calculated using numbers of individuals were found to be more sensitive than other indicators of human perturbation, particularly when comparing sections of any given stream. The city of Grayling's sewage treatment plant's conversion to land dis- posal resulted in increased diversity of the lower Au Sable River from an earlier study. This change was not noticeable by direct measurement of chemical water quality parameters. Nutrient enrichment appeared to result in an increased production of macroinvertebrates, followed closely by a decreased diversity of the com- munity. Factors other than enrichment also were shown to be important in controlling the composition and diversity of the communities. Most important of such factors were substrate types and stability and variation in discharge; these may or may not have been a result of human activity in the watersheds. BENTHIC MACROINVERTEBRATE DIVERSITY IN THREE DIFFERENTIALLY PERTURBED MICHIGAN STREAMS By Scott Jon Reger 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 1973 ACKNOWLEDGEMENTS I extend my sincere gratitude to several people who not only helped in the preparation of this work, but also helped further my overall education: Dr. Kenneth W. Cummins for his invaluable help in attacking the many problems that arose in the course of this study with his knowledge and experience in stream ecology, and for making available his time, equipment, and staff to help with the identification of organisms. Dr. Niles R. Kevern for his overall guidance in my course of study, his help with this manuscript, and for making funds, equipment, and en- couragement available whenever I needed them. Dr. Robert C. Ball for his consultation and encouragement, and for providing the facilities of the Institute of water Research when necessary. Dr. Terry A. Haines, project director, for helping with the logistics of the field work, continued consultation, and providing me with other data from the project. I would also like to thank all those who helped gather field data; particularly Dr. wayne L. Smith, Dr. Adam T. Szluha, and Chris Schmitt. Dr. Richard A. Cole has been most helpful in my gaining understanding of stream community ecology, and the many hours he has spent discussing this project with me are most appreciated. Several people have given laboratory assistance, but I want to extend thanks especially to Jon Lauer, Chris Brand, Jane Kotenko, Larry Hater- worth, Kirk Johnson, and Carolyn Goldberg. 11 For help in the identification of organisms I would like to express special thanks to Jack Huycheck, Robert Peterson, Robert King, and Milton ward. This study was supported (in part) by funds from Grant 14-31-0001-3153, provided by the United States Department of the Interior, Office of water Resources Research, as authorized under the water Resources Act of 1964 and (in part) by the National Science Foundation Institutional Grant for Science, both administered by the Institute of water Research, Michigan State University. Funds and equipment were also provided by the Department of Fisheries and Wildlife and Agricultural Experiment Station, Michigan State University. Use of the Michigan State University computing facilities was made possible through support, in part, from the National Science Foundation. 111 INTRODUCTION . . . . . SITE DESCRIPTION . . . Upper Jordan . . Lower Jordan . . Upper Au Sable . Lower Au Sable . Upper Red Cedar . Lower Red Cedar . MATERIALS AND METHODS RESULTS . . . . . . . Jordan River . . Au Sable River . Red Cedar River . DISCUSSION . . . . . . WY 0 O O O O I 0 LITERATURE CITED . . . APPENDICES . . . . . . TABLE OF CONTENTS iv Page 10 10 10 10 11 12 15 15 22 24 27 32 34 37 Table LIST OF TABLES Page Chemical water quality parameters investigated in 1971-1972 at the study sites . . . . . . . . . 7 Maximum and mdnimum water temperatures (°C) and dissolved oxygen concentrations (mg/l) in 1971- 1972 at the study sites . . . . . . . . . . . . . 8 Physical characteristics of the study sites during the summer of 1971 . . . . . . . . . . . . . . . 9 Number of taxa collected at the study sites during each season and overall total . . . . . . . . . . 17 Productivity as standing crops in number of individuals per square meter at the study sites during each season and annual means . . . . . . . 18 Productivity as standing crops in grams per square meter at the study sites during each season and annual means . . . . . . . . . . . . . 19 Shannon diversity indices and equitability in terms of numbers of individuals at the study sites during each season and annual means . . . . 20 Shannon diversity indices and equitability in terms of biomass at the study sites during each season and annual means . . . . . . . . . . . . . . . . 21 Species collected at the upper Jordan site . . . 37-39 Species collected at the lower Jordan site . . . 40-41 Species collected at the upper Au Sable site . . 42—44 Species collected at the lower Au Sable site . . 45-47 Species collected at the upper Red Cedar site . . 48-50 Species collected at the lower Red Cedar site . . 51-52 Shannon diversity indices (using numbers of 1nd1V1d0818 and biomss) o o o o o o o o o o o 0 53-54 .PI';..IA)II’ IIIEI Ital]!!! I’ll: I! List of Tables (con't.) Table A-8 A-9 A-lO Page Equitability values for diversity indices appearing in Table A-7 . . . . . . . . . . . . . . 55-56 Density as standing crOps (using numbers of individuals and biomass) . . . . . . . . . . . . . 57—58 sz of substrate sampled and number of taxa found . . . . . . . . . . . . . . . . . . . . . . 59-60 vi Figure LIST OF FIGURES Page Map of the Jordan River showing the location of the study sites . . . . . . . . . . . . . . . . . 4 Map of the Au Sable River showing the location of the study sites . . . . . . . . . . . . . . . 5 Map of the Red Cedar River showing the location Of the Study Sites 0 O O O O O O O O O O O O O O 6 vii INTRODUCTION Throughout recent history, man has caused stream degradation to occur at an accelerated rate due to changes in agricultural practices and increased industrialization and associated urbanization. One of the more important changes in streams has been enrichment with nutrients from decomposing organic matter in the stream or in sewage treatment plants. Evaluation of the gradual changes in aquatic communities in response to such enrichment is necessary if we are to develop indicators of stream degradation that can be used to properly manage the disposal of such wastes. Sensitive indicators of enrichment are essential if we are to be able to detect and predict degradation of our waterways at an early stage in the process. Hooper (1969) has pointed out the importance of developing indices that will provide a common language for documenting and assessing rates of change in our aquatic communities. Macroinvertebrate populations are often used as an index of stream conditions because they effectively integrate conditions over time and are responsive to critical conditions of short duration that other sampling might miss (Gaufin and Tarzwell, 1956; Gaufin, 1958; Hynes, 1960). The community structure of benthic macroinvertebrates has thus been widely used as one such indicator. Diversity indices, particularly that of Shannon (Shannon and weaver, 1949), have become one of the more pOpular ways of describing such communities (Hooper, 1969; warren, 1971). Such indices are considered among the best and most sensitive indicators of ecological change (Wilhm and Dorris, 1968; Hooper, 1969; warren, 1971). 2 Species diversity was first used by Wilhm and Dorris (1966) to examine the effects of organic effluents in a stream. Harrel and Dorris (1968) used this method to study a stream system within a single drainage basin. Mathis (1968) compared diversity in three unpolluted mountain streams. Mathis and Dorris (1968) investigated the effects of oilfield brine on the diversity of a stream. Gislason (1971) successfully used species diversity of macroinvertebrates collected from artificial sub- strates to show differing levels of human perturbation on four sites in the same three streams as the present study. He also suggested that seasonal instability in the indices occurred before a reduction in the mean value of the index with increased degradation. One of the purposes of this study was to compare the effects of apparent different levels of human disturbance on species diversity of macroinvertebrates. In particular, it was desired to see how the sensitivity of macroinvertebrate diversity indices compared to other indicators of stream enrichment. To help minimize other influences, the sites were located both above and below known nutrient sources; this had not been done in the previous study. 'Also, natural substrates were sampled to see if they produced results different than those found with artificial substrates, particularly in regard to the seasonal in- stability which may have been an artifact of the particular substrate used. A final objective was to see if the use of biomass units, rather than numbers of individuals, in Shannon's formula gave a better indication of the level of human perturbation. SITE DESCRIPTION Six sites were selected on three streams in the lower peninsula of Michigan (Figures 1-3). The sites were chosen to represent a wide range of apparent human disturbance in the watersheds. Certain chemical parameters were studied throughout the course of this project (Tables 1 and 2). Physical parameters and substrate condi— tions during the summer were also recorded (Table 3). In general, the four more northern sites were similar, while differing from the Red Cedar sites. The lower Au Sable site showed marked diurnal dissolved oxygen fluctuations during the summer, which were attributed to macrophyte growths. The lower Red Cedar site had highly variable dissolved oxygen levels, much higher nutrient levels than the other sites, and heavy metal and pesticide residues. Both the Red Cedar sites were subject to highly variable discharge rates. Upper Jordan The upper Jordan runs through uninhabited state forest land, where, in order to preserve the nearly pristine conditions, camping is prohibited along the banks of the stream. Recreational use is limited to occasional wading fishermen. The stream bed is predominately sand with numerous fallen logs and occasional silt deposits. Chara vulgaris is the pre- dominate plant in the silt. Fontinalis g2, grows on the marl concretions on many of the logs. Sculpin and brown trout are the predominate fish. 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It is slightly wider, somewhat more open, has fewer logs, and lacks the silt deposits found in the upper site, which it otherwise resembles. 92231: in 3821.9. The upper Au Sable site receives some nutrients from the town of Frederick and occasional cottages. Some canoeing and fishing constitute the recreational usage. The substrate is largely gravel and sand with frequent silt deposits along the edges. Potomgggton £22, grows extensively in the silt, and to a lesser extent in the sand, seasonally. Predominate fish are brook trout, sculpin, and darters. massage The lower Au Sable receives effluents from the municipal sewage treatment plant of Grayling, a state fish hatchery, and numerous homes and cottages along the banks. Early in the course of this study the Grayling sewage treatment plant completed conversion to a land disposal system. Extensive use is made of the stream by canoeists and fishermen. The substrate is largely gravel with some sand. Potomogeton £22, and Blades 32, grow in both substrates, Potomogeton filiformis becoming very dense over large areas in the summer and fall, causing diurnal dissolved oxygen levels to occasionally fall below 5 ppm. Brown trout and sculpin are the predominate fish, with many other species occurring, some only seasonally. Upper Red Cedar There is considerable agricultural development, including feedlots, several small towns, and a metal plating plant located above the upper 11 Red Cedar site. Substrate is largely sand and some gravel. A large area of the sand has seasonal growths of Potomoggton s22, Diurnal dissolved oxygen fluctuations are pronounced in warm months, but rarely fall below 5 ppm. There is considerable flooding in the spring and occasionally at other times. Characteristic fish are golden redhorse, rock bass, and bluntnosed minnow, but a great variety of other species, including smallmouth bass, occur in smaller numbers. Lower Red Cedar More agricultural land, several subdivision developments, and the towns of Okemos and East Lansing-Michigan State University also add effluents and runoff above the lower site. The substrate is mostly sand with large silt and organic deposits occurring along the edges and at the bottom of the deeper pools during the periods of low flow. Heavy growths of Potoggggton app, occur seasonally in the sand. Dissolved oxygen levels are highly variable and fluctuate diurnally during the warm mouths, occasionally falling below 1 ppm under extreme conditions. Drastic fluctuations in flow occur with runoff, which in combination with the unstable substrate produce a drastic scouring effect. All fish popula- tions are unstable, but many species occur at times, white suckers being predominate. MATERIALS AND METHODS After preliminary studies were made to determine the major substrates, the two or three major types, as mentioned in the site description, were sampled monthly in triplicate. Where macrOphyte growths occurred in a substrate, some of the samples were taken in an area where the plants were growing. Because of the large number of macroinvertebrates collected, the number of samples analyzed completely for species diversity calcula- tions was restricted to those from one month for each season. Gravel was sampled with a modified Surber sampler; this consisted of adding a 0.5 mm mesh net front and sample collection bag to the riffle benthos sampling device described by Coffman, Cummins, and wuycheck (1971). The sample area is dug up to a depth of 10 cm and washed thoroughly. Log samples were taken by carefully enclosing a portion of a log in a bag of the same material used in the Surber, cutting off, and removing. Sand and silt samples were taken with an Ekman dredge mounted on a pole. Samples were taken to a depth of at least 10 cm whenever possible. Samples were sieved through a #30 0.8. standard sieve (0.595 mm openings) to remove small particulate matter, and preserved in formalin. The samples were later washed, placed in an enamel pan, and the inverte- brates removed by hand under an illuminated magnifier.‘ The invertebrates were then separated and identified as far as possible under a dissecting microscope. Each taxa was then counted and wet weights taken, except no weights were taken for clams and snails. Formalin preserved weights 12 13 are generally considered to have less than 5% loss from live weight (Ball, 1973a; waters, 1973; Winberg, 1971). Howmiller (1972), while reporting greater losses, found formalin to show less weight loss than other common preservatives. Diversity indices using both numbers of individuals and biomass were calculated on a Control Data 6500 Computer. The index used is that of Shannon, as described by Wilhm and Dorris (1968), using sample data to estimate the pOpulation diversity. The formula is: s d = - 151 El. log2 El N N 1 per taxa; s = the number of taxa; and d - the estimate of population where N = the total number of individuals; n = the number of individuals diversity. Values range from 0 to any positive number. Wilhm.(l970) has shown that diversity index values rarely exceed nine, and are usually between three and four for clean water streams and below one in polluted conditions. In communities with habitats containing organisms with clumped distribution a method of estimating the actual population diversity has been presented by Pielou (1966), and modified by Mawson and Godfrey (1971). It consists of plotting the diversity indices of all possible combinations of samples. Wilhm and Dorris (1968) and warren (1971) have shown that pooling an adequate number of samples achieves essentially the same results. Three to five samples were shown to be necessary to accurately estimate population diversity. The six or nine samples used in this study thus should give a good estimate of the true population diversity. This particular index has several attributes that should be noted. It makes possible the objective comparison of community structure between 14 different streams in different geographical areas (Mathis, 1968). It takes relative abundance into account and thus is less affected by rare species which might be missed in sampling. It is independent of sample size. It is dimensionless, thus any appropriate units describing each species contribution to the community can be used. This is important in light of the fact that biomass is often considered to be a more accurate way of assessing the ecological impact of a species on t!» community (Wilhm and Dorris, 1968; Wilhm, 1968). The use of biomass units sub— stitutes a continuous for a discrete variable, but redefines diversity in biomass terms, and thus is related more closely to energy distribution among species (Wilhm, 1968). The present study has used an estimate of live weights in the calculation of diversity. It has been suggested that dry weights or ash-free dry weights are better, and that production values are better yet in assessing the contribution of a species to the community (Dickman, 1968). However, production values are at best, estimates, and require more information than available to date in this study (waters, 1969). An index of the evenness of distribution, or species equitability, E, was calculated using MacArthur's (1965) method. If all species are equally abundant, d - log2 s, and s 8 23. The ratio 23/3, or equitability, is thus a measure of the relative evenness of distribution, with maximum and minimum values of l and 0, respectively. water samples were collected monthly, and sediment and fish samples twice annually at each of the sites. Twenty-four hour continuous monitor- ing of dissolved oxygen and temperature were carried out at each site monthly except during winter. Chemical analyses were determined by the Institute of water Research water Quality Laboratory at Michigan State University. Pesticide residue analyses were made by the Pesticide Re- search Center, Michigan State University. RESULTS Jordan River Direct measurement of nutrient levels and other water chemistry parameters did not reflect the effect of the Jordan River National Fish Hatchery (Tables 1 and 2). The high levels of nitrate and apparent increase at the lower station were shown by Szluha (1971) to come from groundwater. He calculated that 898 kg/yr of phosphorus and 4,173 kg/yr of nitrogen were contributed annually by the hatchery effluents, mostly in pulses when raceways were cleaned. These made up 28.32 and 5.0% of the annual budgets of these elements, respectively, in the Jordan River. He showed that periphytic growth was increased by the nutrient additions, but that primary productivity was well within normal ranges throughout the system, and that the oxygen balance of the stream was not affected significantly. Oxygen fluctuations were somewhat greater at the upper site (Table 2), apparently as a result of the more marked temperature fluctuations. The large amount of groundwater entering be- tween the sites has a moderating effect on temperature and oxygen levels at the lower site. The increased discharge did not increase depth or velocity noticeably, but took the form of a wider channel (Table 3). Both sites were characterized by an abundance of trichopteran, ephemeropteran, and chironomid taxa; the lower site showed considerably fewer taxa (Table 4). This may be, in part, a result of the less stable sand and lack of macrophytes. Both sites had large numbers of ephemerellids, baetids, tipulids (Antocha _23), and tantytarsan midges. The upper station 15 16 Table 4. Number of taxa collected at the study site during each season and overall total. Summer Fall Winter Spring Total Upper Jordan 51 56 54 54 87 Lower Jordan 33 30 50 36 63 Upper Au Sable 75 56 56 48 99 Lower Au Sable 63 56 49 43 90 Upper Red Cedar 51 56 73 18* 102 Lower Red Cedar 31 20 13 10 39 *No gravel samples taken due to high water. 17 also had large numbers of sphaerids (Pisidium s23), amphipods (Hyallela azteca), and hydropsychids (Hydropsyche 822.). The lower station had fewer sphaerids, more Gammarus fasciatus than Hyallela azteca, Cheumatopsyche _2, rather than Hydropsyche 322., and had large numbers of diamesian and orthoclad chironomids, brachycentrids, simuliids, and tubificids as well (Appendices 1 and 2). The lower site had far greater productivity than the upper site, both in terms of numbers of individuals and biomass (Tables 5 and 6). This is true of both the log and sand substrates (Appendix 9). Seasonal patterns appear to be similar at both stations but fluctuations are greater at the lower. While a fair amount of the increased production was due to taxa common at both sites, a large contribution is made by the simulids, brachycentrids, orthoclad chironomids, and particularly the tubificids, which were relatively infrequent at the upper site. The lower site showed less diversity than the upper one, especially as numbers of individuals. The lower site also exhibited a greater seasonal variation in diversity (Tables 7 and 8). The lower diversity is apparently due to the increased production of the taxa mentioned above, especially in the sand substrate (Appendix 9). The diversity on logs is similar at both stations despite the increased production at the lower site (AP‘ pendices 7 and 9). The hatchery effluents had an effect on the macroinvertebrate com- munity of the Jordan River. While some of the changes are due to changes in the character of the substrate, a most important factor in the distri- bution of communities (Thorup, 1964; Hynes, 1970), this also may be re- lated to human disturbance in the watershed resulting from earlier logging operations. The instability of the sand makes the logs a most important habitat, as is often the case where other stable substrates are rare 18 Table 5. Productivity as standing crops in number of individuals per square meter at the study sites during each season and annual means (i 1 S.E.). Annual Summer Fall Winter Spring Mean Upper Jordan 4846 2996 9849 6909 6150 i 1469 Lower Jordan 7701 9793 40225 40598 24579 t 9151 Upper Au Sable 8918 2493 2683 2991 4272 i 1552 Lower Au Sable 15722 17000 12266 10751 13935 i 1458 Upper Red Cedar 3098 4553 10770 6029* 6113 t 1664 Lower Red Cedar 3887 99906 79696 105580 81015 t 15104 *No gravel samples taken due to high water. 19 Table 6. Productivity as standing craps in grams per square meter at the study sites during each season and annual means (i l S.E.). Annual Summer Fall Winter Spring Mean Upper Jordan 7.24 8.49 13.81 12.80 10.59 * 1.60 Lower Jordan 14.30 22.23 98.19 47.48 45.55 + 18.92 Upper Au Sable 25.68 13.99 12.76 15.81 17.06 t 2.94 Lower Au Sable 39.54 54.17 46.18 21.41 40.32 i 6.98 Upper Red Cedar 50.21 46.29 78.07 7.28* 45.46 1 14.56 Lower Red Cedar 36.74 72.40 57.09 72.94 59.79 i 8.52 *No gravel samples taken due to high water. 20 Table 7. Shannon diversity indices and equitability in terms of numbers of individuals at the study sites during each season and annual means (i 1 S.E.). Annual Summer Fall Winter Spring Mean Upper 5 3.98 4.01 4.09 4.09 4.04 t 0.03 Jordan E 0 31 0.29 0.32 0.32 0.31 Lower 0 3.63 2.88 3.79 2.01 3.08 5 0.41 Jordan E 0.38 0.25 0.28 0.11 0.26 Upper 0 4.48 4.52 4.49 4.02 4.38 r 0.18 Au Sable E 0.30 0.41 0.40 0.34 0.36 Lower 5 4.16 4.03 4.25 3.28 3.93 3 0.22 Au Sable E 0.28 0.29 0.39 0.23 0.30 Upper 0 3.96 3.72 3.95 3.01* 3.66 r 0.22 Red Cedar E 0.31 0.24 0.21 0.45 0.30 Lower 0 1.67 0.54 0.38 0.61 0.80 r 0.29 Red Cedar E 0.10 0.07 0.10 0.15 0.11 *No gravel samples taken due to high water. 21 Table 8. Shannon diversity indices and equitability in terms of biomass at the study sites during each season and annual means (i 1 S.E.). Annual Summer Fall Winter Spring Mean Upper 0 3.55 3.58 4.00 3.89 3.76 3 0.11 Jordan E 0 23 0 21 0.30 0 27 0 25 Lower 0 3.70 3.24 3.65 2.86 3.36 3 0.20 Jordan E 0.39 0.32 0.25 0.20 0.29 Upper 5 4.10 3.48 3.84 3.45 3.72 3 0.16 Au Sable E o 23 0.20 0.26 0.23 0.23 Lower 0 3.96 3.87 4.15 3.95 3.98 3 0.06 Au Sable E 0.25 0.26 0.36 0.36 0.31 Upper 0 1.05 1.50 2.64 1.78* 1.74 3 0.46 Red Cedar E 0 04 0 05 0.09 0 19 0 09 Lower 5 2.31 0.66 0.53 0.43 0.98 3 0.45 Red Cedar E 0 16 0 08 0.11 0 13 0 12 *No gravel samples taken due to high water. 22 (Hynes, 1970), and increases the importance of the relatively stable areas where silt collects and macrophytes grow. Attempted colonization from these areas also could account for higher diversity in nearby sand areas in the upper site (Hynes, 1970). While the diversity on logs was essentially the same at both stations, production was greatly enhanced by increased enrichment here as well as in the sand. Some change in community composition were noted in both substrates. These changes, especially those due to the increase in filter feeding simulids and hydropsychids, and detrital feeding brachycentrids, chironomids, and oligocheates, in response to the hatchery effluents are the same as have been reported by the Michigan Water Resources Comission (1969) using Hester-Dendy artificial substrates. The completion of settling basins for the hatchery effluents should alleviate the problem. Au Sable River Direct measurement of nutrient levels and other water chemistry parameters did not reflect the input from the town of Grayling and the State Fish Hatchery (Table 1). Nor did they reflect the removal of the Grayling sewage treatment plant's effluent from the stream. Periphytic production was higher at the lower station and was more stable at both sites than in the other two rivers (Ball, 1973b). The lower section was wider and deeper than the upper, and had considerably less silt and detrital deposition areas, possibly as a result of the increased flow (Table 3). The enrichment of the lower site produced a substantial macrophytic growth, and this resulted in a marked diurnal oxygen fluc- tuation during the summer months (Tables 2 and 3). Both sections were characterized by an abundance of trichopteran and chironomid taxa. The upper site also had a large number of ephemerOpteran 23 and gastrapod taxa. Both stations supported a large number of taxa (Table 4). Both sections had large numbers of amphipods (Gammarus fasciatus), hydrOpsychids, elmids, simulids, and chironomids of the tribes Chironomini and Tantytarsini. The upper site also had large numbers of orthoclad chironomids. The lower site had large numbers of diamesian chironomids, glossosomatids (Agapetus illini), isopods (Asellus militaris), and tub- ificids as well (Appendices 3 and 4). The lower section exhibited a greater production both in numbers of individuals and especially in biomass (Tables 5 and 6). This was true of both the sand, where the difference was greatest in numbers of indiv- iduals, and the gravel, where the difference was greatest in biomass (Appendix 9). While much of this increased production was from taxa common to both sites, the lower had significant contributions from 130pods, diamesian chironomids, and tubificids which were unimportant at the upper. The lower section had slightly lower diversity when calculated using numbers of individuals but appears to possibly have had a slightly higher diversity when using biomass (Tables 7 and 8). These differences occurred mostly in the sand substrate, particularly in areas of macrophyte growth (Appendix 7) where large numbers of smaller organisms, such as chironomids and tubificids, lowered the diversity in terms of numbers of individuals but not in terms of biomass. Neither section showed much seasonal variation in diversity (Tables 7 and 8). The sand did show somewhat more seasonal variation at the lower site (Appendix 7). There was little change in diversity between the two Au Sable sites. Part of this was due to the similarity in physical and chemical character- istics. Also, the lower section had been enriched for a long time prior to this study, at least in comparison to the Jordan River, and thus had had time to develop a rich and stable community under such conditions 24 (Hynes, 1970). Nutrient inputs were less localized and farther up- stream from the lower section than in the Jordan River (Figures 1 and 2). The removal of the nutrient input from the sewage treatment facilities at Grayling also reduced the difference between the two sites. It can be shown that some changes in the composition of the community did occur. The increased production was directly related to enrichment. Differences in the productivity of fish pepulations in the Jordan and Au Sable Rivers have also been attributed to enrichment by Quick (1971), and Smith (1972); although other factors also were thought to have effected the brown trout and sculpin populations studied. Such increased produc- tion in both macroinvertebrate and fish populations has been shown experi— mentally in Berry Creek, Oregon, by enriching sections of the stream with sucrose (warren, 1971). Red Cedar River Direct measurement of nutrient levels and other water chemistry parameters clearly demarked the Red Cedar study sites from those on the Au Sable and Jordan, but did not differentiate between the two sections (Table l). The lower study area was slightly deeper than the upper, and exhibited even greater variability in discharge (Table 3). This, in conjunction withthe unstable and substrate conditions, produced a marked scouring effect. During periods of low flow, silt and detrital de- posits accumulated and heavy macrophyte growth occurred at the lower site. Periphytic production was highly variable and showed no difference between the two sections. Oxygen levels were somewhat low at the upper site and quite low at the lower site (Table 2). On August 11, 1971, oxygen de- pletion caused a major fish kill at the lower section and immediate 25 vicinity. Following this the number of taxa of macroinvertebrates found was less, diversity decreased, and the standing crop of the taxa remaining was increased for the duration of the study. Pesticide and heavy metal residues occurred in moderately high levels in sediment and fish, particu— larly at the lower study area (Haines, 1971). The upper site had a large number of taxa, especially of Trichoptera, Chironomidae, and Gastrapoda; the lower showed a distinct paucity of taxa (Table 4; Appendices 5, 6, and 10). The upper station showed an increase in number of taxa, especially more pollution intolerant forms, and numbers of individuals of such groups, during the winter (Appendix 5). Subsequent to the fish kill in August the lower station displayed an even greater lack of macroinvertebrate variety (Appendix 6). The upper site had large numbers of amphipods, hydropsychids (Cheumatopsyche _pp), elmids, pelycepods, and chironomids of the tribe Chironomini. The lower was predominated by naidid and tubificid oligocheates, planarians (Dugesia spp,), and a moderate number of chironomids of the tribe Chironomini (Appendices 5 and 6). The lower section showed higher productivity, particuarly in terms of numbers of individuals (Tables 5 and 6). This was the result of the large numbers of relatively small oligocheates at the lower site. When comparing the sand substrate (in effect the only substrate at the lower site), especially the more stable areas with macrophytes, the difference in production is more marked, both as numbers of individuals and biomass (Appendix 9). Diversity was considerably less at the lower site, particularly when using numbers of individuals. The upper section had fairly high values calculated from numbers of individuals but is low when biomass is used. The lower site had low values in both respects. Equitability was 26 noticeably .ower for biomass indices at the upper, and both numbers of individuals and biomass at the lower Red Cedar than either index at any other site (Tables 7 and 8). The difference between stations was shown more markedly when the diversity of the sand substrata are compared - as was the case with productivity - and for much the same reasons; the lower site having much lower values and exhibiting greater seasonal fluctuation (Appendix 7). The gravel added a great deal of substrate variety and stability to the upper section, thus increasing both the number of taxa and the diversity values. Many parameters showed both the Red Cedar sites to be more enriched than sites on the other rivers, particularly the lower Red Cedar. The oxygen depletion which caused the fish kill in the lower section undoubtedly effected the macroinvertebrate community as well. The unstable sub- strate and fluctuation in discharge were the result of runoff from agricultural and urban develOpments. This runoff, particularly from the combined storm and sanitary sewers, was also responsible to a large degree for differences in water chemistry from the other streams. The fact that coarse gravel supports a more varied and stable community (Hynes, 1970) kept the diversity at the upper site at a moderate level. In fact the sand at the upper study area also had quite low diversity values (Appendix 7). Sand has often been shown to be a relatively poor substrate and to be more susceptible to reduction in numbers and diversity of macroinverte- brates than other substrates (Hynes, 1970; Wilhm and Dorris, 1966). The lack of stable substrate together with an abundant supply of particulate organic material lead to the large production of the few species, such as tubificids, that do well under such conditions: and the subsequent reduction in diversity, particularly in the lower Red Cedar. Similar re— sults have been reported by Jensen (1966) and Talsma (1972). DISCUSSION Macroinvertebrate diversity indices were found to be more sensitive than other indicators of human perturbation when comparing stations on any given stream. Sites were located close enough together so decreased diversity and increased production were largely the result of the known human activity between the sites and not of natural changes encountered moving downstream in the watershed. The increased production and decrease in mean annual diversity, using numbers of individuals, from 4.04 to 3.08, show the effect of the Jordan River National Fish Hatchery on the river. These results agree with those of Gislason (1971), who showed a decrease from 4.33 to 2.98 when using artificial substrates, and with those of the Michigan Water Resources Commission (1969). A small amount of enrichment from the hatchery had a marked effect on this pristine stream where nutrients were the limiting factor to productivity. The irregular pattern of nutrient addition was an unpredictable event to the stream organisms, and thus resulted in an unstable condition. The mean annual diversity values, using numbers of individuals, of 4.38 and 3.93 for the Au Sable sites are not considered different. The 3.93 value for the lower site is considerably higher than the 2.76 reported by Gislason (1971). While some of the difference may have been due to his selec— tion of artificial substrate, the major reason was the reduced level of en— richment subsequent to Grayling's land disposal system becoming operational. It is significant that this reduction in enrichment was detected by 27 28 macroinvertebrate diversity although undetectable when using other para- meters such as water chemistry. The mean annual diversity values, using numbers of individuals, of 3.66 and 0.80, clearly show the degradation between the two sites on the Red Cedar River. Talsma (1972) attributes much of this degradation directly to the input from the large number of drains entering between the sites. The diversity values agree with Talsma's (1972) 4.13 and 1.35 values as well as Gislason's (1971) value of 1.70 for a site further downstream. The Red Cedar was not responding to nutrient enrichment, but was limited by other factors. These factors included the introduction of toxic substances and oxygen depletion, as well as the lack of stable substrate and large fluctuations in discharge. These diversity index values show the changes in the macroinverte- brate communities caused by human activity. They also follow closely the suggestion of Wilhm and Dorris (1968) and Wilhm (1970) that values greater than 3.0 are generally indicative of clean waters, 3.0 to 1.0 of moderately disturbed conditions, and of less than 1.0 of grossly polluted conditions. The ability to compare levels of perturbation between streams by use of diversity indices is less sensitive than the comparison of sites on the same stream, as other variables may also effect diversity values. As mentioned previously, substrate conditions and discharge have signif- icant effects on the macroinvertebrate community and may or may not be related to human usage of the watershed. Current velocity has been shown by Szluha (1972) to be correlated with periphytic production, and Popma (1971) concluded that macrophytic growth was more closely related to discharge and substrate conditions than to nutrient levels in the streams studied. These in turn effect the macroinvertebrate community. Many 29 organisms and communities are closely related to substrate: (Thorup, 1964; Hynes, 1970). The variety of substrate types, including the types of macrophytes, also has a large effect on the community (Cole, 1973; Hynes, 1970). The correlation reported by Gislason (1971) between magnitude of seasonal variation in diversity indices and level of perturbation is believed to have been partially the result of seasonal variation in dis— charge, but largely of the artificial substrates designed to simulate macrophyte beds. Such growths are seasonal in nature in temperature climates and thus would be expected to be used in a seasonal manner by macroinvertebrates evolved under such conditions, probably as a substrate during the summer and as a food source by those utilizing detritus in the fall. This was the case in the Pine River studied by Barber (1970). Samples containing macrophytes in the present study showed less seasonal variation than the unstable sand substrate, but more variation than the more stable gravel or log substrates (Appendix 7). Diversity of numbers of individuals would appear to be more sensi- tive than of biomass when describing differences between two sites on a given stream, thus supporting the current usage of such indices. Diversity in terms of biomass might be considered to be a better way of classifying the areas of the three streams according to the apparent level of human perturbation, at least with respect to where the upper Red Cedar fits in the classification system suggested by Wilhm and Dorris (1968). Enrichment would appear to have caused an increase in production before effecting a lowering of the diversity indices, particularly when using numbers of individuals. This can best be seen in the log samples from the Jordan River and the gravel samples from the Au Sable River (Appendix 9). It has been suggested (Waters, 1961, 1966; Dimond, 1967) 30 that drift of macroinvertebrates might be a function of production in excess of carrying capacity. If this is the case, standing crop estimates alone would not accurately estimate production. Eyman (1969) found that while standing crops of macroinvertebrates were not well correlated with the apparent enrichment in the three streams studied, total drift was. He therefore concluded that drift analysis could give indications of changes taking place at an early stage of enrichment. Diversity indices are a useful tool to show the amount of human perturbation of streams, but they are only a manifestation - they provide no framework for causal explanation. As has been pointed out, many factors which may or may not be man caused can effect changes in diversity indices. Other methods of analysis, such as the tolerance of certain groups of organisms to various substances, the food habits of the community, or methods of obtaining oxygen, may show a closer relation to the cause of the alteration of the community. At times, standing crops or drift might better show enrichment. A11 methods of analysis available should be used, as any method of data reduction does only that, reduce data; hopefully in a concise manner and to a form that is more easily used by workers in other fields. In this study the diversity of macroinvertebrate communities, cal- culated using numbers of individuals, was found to be the most sensitive indicator of human perturbation of streams, particularly enrichment. The effect of the hatchery on the Jordan River and the removal of sewage effluents from the Au Sable River demonstrate the sensitivity of Shannon's diversity index to enrichment. Results were most clear when other dif- ferences were minimal; this limits the use of such an index in comparing different types of streams. 31 As Hynes (1960) has warned, it is a mistake to rely on any one formal method of data analysis, as doing so tends to lead to rigidity of thought and approach. The investigator should use all means feasible under the prevailing conditions. Warren (1971) concurs when he states that such indices "should not replace the use of other knowledge about the biology and environmental requirements of the species contributing to diversity." Both agree that the presentation of tables of species and numbers permit the use of many methods of analysis and interpretation. As Warren (1971) has so succinctly put it - "There is no single path to understanding." SUMMARY Benthic macroinvertebrate communities in three Michigan streams were studied through an annual cycle. Natural substrates were sampled both above and below known areas of human activity on each stream. Taxonomic composition, standing crop, and diversity indices were used in an attempt to find a sensitive indicator that would show the varying levels of pertur- bation of the streams. The Jordan River exhibited the effects of the National Fish Hatchery's enrichment of the stream. Standing crop was increased, number of taxa found decreased, and diversity values were lower below the hatchery. The slight enrichment of this pristine stream had a measurable effect on the macroinvertebrate community. The Au Sable River had a higher standing crop below the city of Grayling and the state fish hatchery, but showed little difference in number of taxa collected and diversity from the section above these inputs. The important disclosure here was the marked increase in diversity from an earlier study conducted by Gislason (1971). This was a result of the Grayling sewage treatment plant's completion of conversion to a land dis- posal system. The diversity index responded to this change that water chemistry data failed to show. The Red Cedar River exhibited a dramatic change in taxonomic compo- sition and a marked reduction in diversity at the lower site. Here sub— strate stability, variable discharge, periodic oxygen depletion, and occasional introduction of toxic substances were the cause of the reduced macroinvertebrate communit stability. An oxygen depletion caused fish and 32 33 invertebrate kill in the lower section early in the study led to a de- creased number of taxa and decreased diversity, but an increased production of the remaining taxa for the remainder of the study. Diversity indices were found to be the most responsive to changes in stream conditions of the indicators examined. As macroinvertebrate communities integrate all the conditions they are exposed to, they are useful for detecting subtle changes in the lotic environment. They have been shown useful in detecting the effects of enrichment in an above and below situation on the Jordan River, a before and after situation on the Au Sable River, and to respond to other conditions in the Red Cedar River. However, as these communities act as integrators, care must be used when attempting to attribute changes in diversity to a specific cause. This makes comparison among streams difficult, especially if the streams are physically or chemically disimilar. Whenever possible other parameters should be studied and other indicators used in conjunction with diversity indices. LITERATURE CITED Ball, R. C. 19733. Personal communication. Ball, R. C. 1973b. Unpublished manuscript. Barber, W. E. 1970. Ecological factors influencing macroinvertebrates in the Pine River. Ph.E. thesis. Michigan State University, 77 p. Coffman, W. P., K. W. Cummins, and J. C. Wuycheck. 1971. Energy flow in a woodland stream ecosystem 1. Tissue support trophic structure of the autumnal community. Arch. Hydrobiol. 68(2): 232-276. Cole, R. A. 1973. Variations in stream community response to nutrient enrichment. 4, water Poll. Contr. Fed. In press. Dickman, M. 1968. Some indices of diversity. Ecology, 49(6): 1191-1193. Dimond, J. B. 1967. Evidence that drift of stream benthos is density related. Ecology, 48: 855-857. Eyman, L. D. 1969. A comparison of invertebrate drift in three Michigan streams. M.S. thesis, Michigan State University, 44 p. Gauffin, A. R. 1958. The effects of pollution on a mid-western stream. The Ohio Journal gf_Science, 58(4): 197-208. Gaufin, A. R. and C. M. Tarzwell. 1956. Aquatic macroinvertebrate communities as indicators of organic pollution in Lyttle Creek. Sewage and Ind. Wastes, 28(7): 906-924. Gislason. J. C. 1971. Species Diversity of Benthic Macroinvertebrates in Three Michigan Streams. M.S. thesis, Michigan State University, 53 p. Haines, T. A. 1971. An ecological evaluation of stream eutrophication. Interim completion report, Office of Water Resources Research, project C-2205. Harrel, R. C. and T. C. Dorris. 1968. Stream order, physico-chemical conditions, and community structure of benthic macroinvertebrates in an intermittent stream system. Am, Mid. Nat. 80: 220-251. Hooper, F. F. 1969. Eutrophication indices and their relation to other indices of ecosystem change. pp. 225-235 In: Eutrophication: Causes, Consequences, Correctives. National Academy of Sciences. Washington, D. C. 34 35 Howmiller, R. P. 1972. Effects of preservatives on weights of some common macroinvertebrates. Trans. Amer. Fish Soc. 101(4): 743-746. Hynes, H. B. N. 1960. The Biology of Polluted Waters. Liverpool Univ- ersity Press, Liverpool, England. Hynes, H. B. N. 1970. The Ecology of Running Water. University of Toronto Press, Toronto, Ontario, Canada. Jensen, A. L. 1966. Stream water quality as related to urbanization of its watershed. M.S. thesis, Michigan State University, 128 p. MacArthur, R. H. 1965. Patterns of species diversity. Biological Reviews, 40: 510-533. Mathis, B. J. 1968. Species diversity of benthic macroinvertebrates in three mountain streams. Transactions g£_the Illinois State Academy gf_Science, 61: 171-176. Mathis, B. J. and T. C. Dorris. 1968. Community structure of benthic macroinvertebrates in an intermittent stream receiving oil field brines. Am. Mid. Nat. 80: 428-439. Mawson, J. C. and P. J. Godfrey. 1971. DIVERSE: A fortran IV program to calculate diversity indices of stream bottom organisms. Water Resources Research Center. Univ. of Mass., Amherst. Michigan Water Resources Commission. 1969. Biological monitoring of the Jordan River, vicinity of the Jordan River National Fish Hatchery, Elmira, Michigan. Dept. of Natural Resources, Lansing, Michigan. Pielou, E. C. 1966. The measurement of diversity in different types of biological collections. g, Theor. Biol. 13: 137-144. Popma, T. J. 1971. A comparative study of standing crops and of phos- phorus and nitrogen contents of four macrophyte stream communities. M.S. thesis, Michigan State University, 45 p. Quick, Robert F. 1971. The age and growth of brown trout (Salmo trutta) - and sculpin (Cottus £22,) as it relates to eutrophication in the Jordan and Au Sable Rivers. M.S. thesis, Michigan State University, 86 p. Shannon, C. E. and W. Weaver. 1949. The mathematical theory of communica- tion. University of Illinois Press, Urbana. Smith, Wayne L. 1972. The dynamics of brown trout (Salmo trutta) and , sculpin (Cottus 322.) populations as indicators of eutrophication. Ph.D. thesis, Michigan State University, 43 p. Szluha, Adam T. 1972. Potomological effects of fish hatchery discharge on the Jordan River, northern lower Michigan. Ph.D. thesis, Michigan State University, 54 p. 36 Talsma, A. R. 1972. The characterization and influence of domestic drains on the Red Cedar River. M.S. thesis, Michigan State Univer- sity, 127 p. Thorup, Jens. 1964. Substrate type and its value as a basis for delineation of bottom fauna communities in running water. In: K. W. Cummins, g£_§l. (ed) Organismic-Substrate relationships in streams. Pymatuning Symposia in Ecology Special publication #4. Warren, C. E. 1971. Biology and Water Pollution Control. W. B. Saunders and Co., Philadelphia, Pennsylvania. Waters, T. F. 1961. Standing crop and drift of stream bottom organisms. Ecology, 42: 532-537. Waters, T. F. 1966. Production rate, population density, and drift of a stream invertebrate. Ecology, 47: 595-604. Waters, T. F. 1969. The turnover ratio in production ecology of fresh- water invertebrates. The American Naturalist, Vol. 103, No. 930, pp. 173-185. Waters, T. F. 1973. Personal communication. Wilhm, J. L. 1968. Use of biomass units in Shannon's formula. Ecology, 49(1): 153-155. Wilhm, J. L. 1968. Range of diversity index in benthic macroinverte- brate populations. J, Water Poll. Contr. Fed. 42: R221-R224. Wilhm, J. L. and T. C. Dorris. 1966. Species diversity of macroinverte- brates in a stream receiving domestic and oil refinery effluents. Am. Mid. Nat. 76: 427-449. Wilhm, J. L. and T. C. Dorris. 1968. Biological parameters for water quality control. Bioscience. 18(6): 477-481. Winberg, G. G. (ed.). 1971. Methods for the Estimation of Production of Aquatic Animals. Academic Press, New York. 175 p. APPENDICES 37 Table A1. Species collected at the upper Jordan site. Species 8/12/71 Number Collected 10/09/71 1/30/72 5/20/72 HYDRACAR INA ISOPODA Asellus milltaris 57 AMPHIPODA Gammarus fasciatus 24 Hyallela azteca 117 ODANATA *Enallagg _sp. Cordulegaster _sp. *Gomphus _p. PLECOPTERA Pteronarcys _p. Nemoura _p. l Isogerla _spp. 2 *Acroneuria _p. Paragn_etina _p. 1 TRICHOPTERA Lype _p. 1 unknown psychomyiid Psychomia _2. l Polycentrogus _2. Cheumatopsyche _p_. ifldropsyche _p. 8 fl. slossonae Rhyacophila _p. Glossosoma .B- 1 Agaylea multipunctata Neotrichia _p. Phryageniadae l Brachycentrus americanus Micrasema _p_. Lepidostoma _p. 5 PyncnOpsyche _p_. Mollana _p. 4 Mystacides _sp. Oecetis _p. EPHEMEROPTERA Ephemera simulans 2 *Hexagenia limbata Caenis _p. Tricorythodes _p_. 33 12 “#H 236 10 12 15 36 80 25 260 127 13 22 10 27 3O 38 23 H0 0001—11-0 NHN ll 38 Table A1 (con't.) EPHEMEROPTERA (con' t. ) Stenonema _p, 2 l4 Epeorus _p. l l 27 46 Ephemerella _p. 4 160 824 258 Baetisca _p, 1 1 Baetis_Jg. 134 8 79 72 Pseudocloeon _p_. 18 Paraleptophlebia s2. 35 1 E, debillis 2 g. mollis 21 4 HEMIPTERA Corixidae 2 DIPTERA Tipula _p. 1 l Antocha s2. 41 186 196 61 #Hexatoma _p, 3 2 l Pedicia _p. l 8 Liriope sp. 4 Simulium _p, 6 3 Prosimulium _p, l 12 Odontomyia _p, l 1 Tabaninae 8 4 Chrysops _p, 4 3 2 Atherix variegata 2 2 20 Epididae 9 2 118 15 Unknown Diptera 'A' 33 24 135 Prodiamesia _p, 2 2 10 6 Orthocladiinae 85 55 184 318 Cardiocladius 42. 28 13 22 95 Cricotomis _2. 10 Tanypodinae 37 34 206 102 Polypedilum _p, 7 1 50 320 Microtendipes _p, 42 131 84 Cryptochironomus _p, l Tantytarsini 283 ll 95 221 Ceratapogonidae ‘ 6 6 26 29 COLEOPTERA Unknown Coleoptera 1 Optioservus _p, 17 35 40 47 Haliplus 32, l l MEGALOPTERA Sialus “p, l 6 l Nigronia _p_. 2 2 Table A1 (con't.) 39 GASTROPODA Aplexa hypnorum Amnicola _p, Promentus exacuous PELECYPODA Sphaerium _p. Pisidium _p. HIRUDINEA unknown Hirudenea Hellobdella stggnalis fl. fusca TRICLADIA Dugesia tigrina unknown Turbellaria OLIGOCHEATA Lumbriculidae Tubificidae Naididae 93 276 45 mN 11 32 *Species found in the study site that was not present in the samples analyzed quantitatively. #Includes Eriocera. 40 Table A2. Species collected at the lower Jordan site. Number Collected Species 8/12/71 10/09/71 1/30/72 5/20/72 AMPHIPODA Gammarus fasciatus 84 100 257 9 Hyallela azteca 4 PLECOPTERA Pteronarcys s2, 2 4 Nemoura s2, 1 l N. venosa 4 6 Brachyptera _p, 1 IsoBeria £22, 5 60 189 32 *Acroneuria s2, Paragggtina _p, l TRICHOPTERA unknown Trichoptera 3 Psychomia _p, 1 l 8 psychomyiid genus 'A' l 12 56 2 Polycentropus _p, 4 2 4 Cheumatopsyche _p, 7 9 762 25 Hydropsyche s2, 20 4O 2 _H. recurvata 1 Rhyacophila _p, l Agapetus illini 1 2 Glossosoma _p, l Brachycentrus americanus 7 69 503 201 Micrasema JR. 4 82 57 Lepidostoma _p, 2 EPHEMEROPTERA Tricorythodes s2. 17 Stenonema _p, 6 Epeorus _p. 1 6 Ephemerella 522. 31 256 1876 514 Baetis _p, 298 80 738 139 Pseudocloeon 43. 182 Paraleptophlebia _p, 5 116 HEMIPTERA Merragata s2. 1 l DIPTERA Antocha _p. 83 60 140 19 ’Hexatoma _p. 3 Dicranota _p_. 4 Simulium _p, 178 8 8 107 Prosimulium _p. 46 11 Odontomyia _p, 8 5 41 Table A2 (con't.) DIPTERA (con't.) Tabanus _p, 2 Atherix variegata l LimnoPhora aquifrons l l Epididae l 2 54 3 Diamesia s2, 194 25 Prodiamesia _p, 136 26 175 29 Orthocladiinae 144 29 11 272 Cardiocladius _p, 43 3 384 26 Metrocnemius _p, 106 268 Cricotopus _p, 1483 Tanypodinae 31 34 486 41 Polypedilum _p, 5 8 16 Microtendipes _p. 2 101 9 Tantytarsini 295 4 153 94 Ceratapogonidae 1 8 COLEOPTERA unknown Dytiscid 2 Acilius_Jg. l Optioservus 32, 42 10 211 35 GASTROPODA Physa £2, 4 Aplexa hypnorum 4 PELECYPODA Sphaeriumisp, 1 3 Pisidium _p, l 3 4 9 TRICLADIA Dugesia tigrina 8 OLIGOCHEATA Lumbricuidae 4 Lumbriculidae 6 6 57 3 Tubificidae 483 681 1820 3793 *Species found in the study site that was not present in the samples analyzed quantitatively. #Includes Eriocera. 42 Table A3. Species collected at the upper Au Sable site. Number Collected Species 8/11/71 10/02/71 1/29/72 5/19/72 HYDRACARINA l AMPHIPODA Gammarus fasciatus 146 14 29 8 ODANATA Agrion _p. 8 l Ishnura _p, 11 Boyeria JR. 5 1 Qggphus _p, 1 1 Ophiggomphus s2, 1 1 PLECOPTERA Nemoura _p, 84 Taeniopteryx s2. 8 5 Isoperla JR. 8 1 TRICHOPTERA Chimarra_Jp. 2 psychomyiid genus 'A' l l Polycentropus J2. 1 3 E, glacialus 1 *2, flavus Cheumatopsyche gp, 229 94 133 41 unknown hydropsychid l Hydropsyche ER, 222 15 60 10 fl, recurvata 120 58 27 18 fl. slossonae 1 UL bifida group 61 12 6 7 fl. betteni 3 Arctopsyche _p, l Rhyacgphila _p, 5 3 4 6 Agapetus illini 1 Glossosoma _p, l 2 Protoptila 32, 1 ‘Aggaylea multipunctata 3 phyraganeid genus 'A' 1 Brachycentrus americanus 6 2 B. numerosus 3 B. lateralis 30 l 10 Lepidostoma _p, 3 ' 7 Goera _p, l Limnephilus 32, 6 Pyncnopsyche _p, l 2 Neophylax 32, 1 1 *Ganonemauyg. Leptocella.Jp. 3 43 Table A3 (con't.) EPHEMEROPTERA Ephemera simulans 17 140 6 2 Hexagenia limbata 21 12 12 Caenis _p, 48 104 3 33 Tricorythodes _p. 77 13 Stenonema JR. 5 15 2 Ephemerella _p, 3 6 57 76 Baetisca _p, 2 13 3 Baetis _p, 9 l 13 Pseudocloeon _p, 257 4 l Siphlanuuus 52. 1 l Isonychia _p_. 98 l Leptophlebia _p. 5 l ParaleptOphlebia praepidita 1 39 HFMIPTERA Trichocorixa _p, 5 2 2 DIPTERA Tipu1a_Jp. l Antocha _p, 7 23 7 9 #Hexatoma‘_p, 9 11 5 l Simulium JR. 315 8 29 8 Prosimulium _p. 117 Tabaninae 4 2 Chrysops _p, 13 25 17 12 Atherix variegata 4 4 2 7 Epididae 4 6 3 Diamesia _p, 3 1 5 Prodiamesia JR. 5 64 2 Orthocladiinae 109 66 109 106 Cardiocladius _p, 4 24 Tanypodinae 224 31 33 61 Conchepeloyia _p_. 2 Chironomini l4 2 33 Polypedilum _p_. 255 147 161 106 Microtendipes ya. 261 19 58 8 Cryptochironomus _p. 10 Tantytarsini 671 41 15 261 Ceratopogonidae 14 8 7 11 COLEOPTERA thioservus 32, 53 37 62 9 Dubaraphia _p, 7 2 2 8 Stenelemis _g. 2 4 Donacia £2. 1 MEGALOPTERA Sialus _p. 6 7 1 2 4 Nigronia 92. 5 Table A3 (con't.) 44 DECAPODA Orconectes virilis GASTROPODA Physa _p. Ammicola s2, Somatggyrus _p, Promentous _p, Gygaulus s2. Lioglax _p. Egrrisia 52. PELECYPODA Sphaerium s2. Pisidium‘gp. Lampsilis g2, HIRUDENIA Hellobdella fusca TRICLADIA Dugesia tigrina unknown Turbellaria OLIGOCHEATA Lumbricuidae Lumbriculidae Tubificidae Naididae 20 259 29 36 22 17 33 32 15 11 ll *Species found in the study site that was not present in the samples analyzed quantitatively. #Includes Eriocera. 45 Table A4. Species collected at the lower Au Sable site. Number Collected Species 8/10/71 10/02/71 1/29/72 5/19/72 HYDRACARINA l 1 ISOPODA Asellus militaris 91 193 125 7 AMPHIPODA Gammarus fasciatus 251 627 ' 7O 5 LEPIDOPTERA Paragyractus s2, 1 PLECOPTERA Nemoura s2. 3 IsoEerla s2, 4 Paragggtina _p, 1 TRICHOPTERA Chimarra _p. 1 2 E, feria 1 Q, alterrima 1 Q, obscura 1 1 unknown psychomyiid l Psychomyia _p, 8 14 10 psychomyiid genus 'A' 29 29 1 Polycentropus _p. 4 3 P, centralis l 2 l Phylocentropus s2, 2 Neuriclipsis _p, 1 Cheumatopsyche _p, 831 502 275 19 Hydropsyche _p, 272 129 245 32 H, recurvata 504 180 289 68 fl, slossonae 87 45 51 11 fl. hifida group 403 381 301 62 Arctopsyche _p. 1 Rhyacophila sp, 1 l3 Aggpetus illini ~ 106 482 270 908 Glossosoma _p, 6 40 12 Protoptila ya. 78 1 28 Hydroptila _p, " 3 2 Agraylea multipunctata 44 Brachycentrus _p, 2 B, americanus 16 26 21 26 B, numerosus l 2 B. lateralis 20 3 8 Micrasgg§__p, l Lepidostoma _p, 7 1 Neophylax _p, l 46 Table A4 (con't.) TRICHOPTERA (con't.) Ganonema s2, 7 Heliocopsyche borealis 1 Leptocella _p. 3 EPHEMEROPTERA Ephemera simulans 1 1 l Tricorythodes _p, 4 Stenonema _p, 17 15 34 l Ephemerella _p, 5 20 219 26 Baetisca _p, 2 l Baetis _p, 439 82 7 4 Pseudocloeon £2. 79 30 2 Siphlonurus g2, l Paraleptophlebia mollis l HEMIPTERA Corixidae 1 DIPTERA Tipula _p, l 5 l 1 Antocha s2, 4 61 87 17 Simulium _p, 988 924 867 133 Prosimulium _p, 1 l 1 Chrysogsin. 7 l 2 3 Epididae 2 5 5 6 Diamesia _p, 332 165 214 210 Prodiamesia _p, 50 7 138 6 Orthocladiinae 143 32 103 94 Cardiocladius_Jp. 12 40 340 11 Conchepelopia_Jg. 15 1 l Chironomini 8 24 Polypedilum _p, 228 114 69 400 Microtendipes _p, 9 47 243 24 Tantytarsini 216 125 192 119 COLEOPTERA thioservus _p, 112 317 160 69 Dubaraphia s2. 1 2 Haliplus _p. 1 MEGALOPTERA *Sialus _p. Nigronia _p, l DECAPODA Orconectes virilis 1 1 Table A4 (con't.) 47 GASTROPODA Physa _p, Amnicola._2, Somatogyrus _p, Planorbula _p, Gflaulus _p. Helisoma antrosa Lioglax _p, Ferrisia _p. PELYCEPODA Sphaerium‘sp, Pisidium s2, HIRUDINEA Erpobdella punctata *Nephelopsis obscura *Hellobdela nepheloidea TRICLADIA Dugesia tigrina 2, microbursalis OLIGOCHEATA Lumbriculidae Tubificidae Naididae 99 10 602 h‘h‘£~\l l'" 51 96 166 82 1209 16 19 542 *Species found in the study site that was not present in the samples analyzed quantitatively. 48 Table A5. Species collected at the upper Red Cedar site. Species 8/18/71 Number Collected 10/19/71 1/18/72 5/10/72 HYDRACARINA AMPHIPODA Gammarus fasciatus Hyallela azteca LEPIDOPTERA Paragyractus _p, ODANATA Agrion,_p, Hetaerina _p, unknown Coenagrionid Ishnura_Jg. Enallagga _p, Gomphus.Jg. PLECOPTERA Taeniopteryx _p. Perlinella drymo Acroneuria _p, Phasgonophora _p, Paragnetina _p. Classenia.Jg. TRICHOPTERA unknown Trichoptera Psychomyia _p, psychomyiid genus 'A' Polycentropus _p, P, remotus P, cineirius Neuriclipsis _p, Cheumatopsyche_Jp. Hydropsyche _p, _H. recurvata fl, bifida group H. aerata Rhyacophila _p, Glossosoma _p, Orthotrichia _p, Agraylea_Jg. Brachycentrus _p, _B. americanus Sericostoma _p, Helicopsyche borealis Athripsodes ancylus _A_. dilutus Oecetis eddlestoni 353 one: O‘h‘h‘ 36 50 514 O‘UOl-‘H P'P‘P‘ w 14 11 12 NM 0‘ MO‘ NHl-‘N 21 49 Table A5 (con't.) EPHEMEROPTERA Hexagenia limbata 3 18 l Caenis _p, 94 8 1 2 Tricorythodes.§p, 8 l Stenonema sp. 20 26 26 Baetis _p, 21 4 41 l Siphlonurus £2. 1 Isonychia s2, 2 Leptophlebia _p, 5 13 Paraleptophlebia s2, 8 l HEMIPTERA Corixidae 2 DIPTERA Tipula _p, 1 Antocha £2. 1 48 #Hexatoma s2, 23 l 15 Psychoda _p, l Simulium s2, 9 Chrysops _p. 7 Atherix variegata 1 Epididae 5 Prodiamesia JR. 9 Orthocladiinae 50 445 Cricotopus _p, 838 Tanypodinae 35 34 46 3 Chironomini 40 192 242 160 Chironomus _p, 7 232 466 21 Polypedilum _p, 23 3 Microtendipes _p, 1 69 44 Cryotochironomus _p. 1 26 Endochironomus s2. 1 Tantytarsini 50 56 335 36 Ceratopogonidae 7 COLEOPTERA Optioservus _p, 1 3 Dubaraphia JR- 25 47 32 8 §£ene1emis _p, 127 62 270 Macronychus _p, l 3 Psphenus herriki 2 3 2 MEGALOPTERA Sialus _p. 12 16 2 Corydalus cornutus l Nigronia _p, 2 DECAPODA Orconectes propinquis 10 5 4 Table A5 (con't.) 50 GASTROPODA Physa _p, Aplexa hypnorum Amnicola s2, Somatogyrus _p, Promentus exacuous Cyraulus _p, Helisoma antrosa LiOplax _p, Stagnicola s2, §, emarginata Viviparous _p, Campeloma _p. Ferrisia _p. Pleurocera acuta PELYCEPODA Sphaeriumi_p. Pisidium s2. *Antodontoides _p, HIRUDENIA unknown Hirudenia Hellobdela _p, TRICLADIA *unknown Turbellaria Dugesia tigrina Q, microbursalis OLIGOCHEATA Tubificidae Naididae bid-b 77 17 14 NW 30 31 N 143 17 bNbLflH 64 56 28 ll 32 58 13 *Species found in the study site that was not present in the samples analyzed quantitatively. #Includes Eriocera. 51 Table A6. Species collected at the lower Red Cedar site. —.- Number Collected Species 8/19/71 10/29/71 1/19/72 5/10/72 ISOPODA Asellus militaris l ODANATA Enallagma _p, 5 Ishnura _p. 2 TRICHOPTERA Cheumat0psyche _p, 2 1 Hydr0psyche slossonae l DIPTERA unknown Diptera 2 Psychoda _p, 1 Simulium _p. 6 5 4 Muscidae 2 2 Orthocladiinae 191 25 5 9 Tanypodinae 58 43 15 Chironomini 158 22 Chironomus _p, 202 16 2 17 Polypedilum _p. 124 1 Cryptochironomus ER, 1 Tantytarsini 61 l COLEOPTERA Dubaraphia _p, 5 9 3 Stenelemis _p. l DECOPODA Orconectes propinquis l GASTROPODA Physa _p. 2 2 Aplexa hypnorum l Amnicola _p, 20 Planorbula _p. l Promentus exacuous 1 Cyraulus _p, 4 Helisoma antrosa l l PELYCOPODA Sphaerium‘sp, l 4 Pisidium JR- 92 103 15 191 Lampsilis _p, l Table A6 (con't.) 52 HIRUDENIA unknown Hirudenia Hellobdella stagnalis .fl. fusca Glossiphonia complanta TRICLADIA Dugesia tigrina Q, microbursalis OLIGOCHEATA Lumbriculidae Tubificidae Branchiura sowerbyi Naididae 3942 11 466 199 12708 667 3078 268 6547 492 53 00.0000.0 00.0 00.0 50.0 00.0 00. 0000. 0 00.0 00.0 00.0 00.0 moaeamm 00¢ 00.0000.0 00.0 00.0 00.0 00.0 00. 0000. 0 00.0 00.0 00.0 00.0 00\3 0am 00.00HH.0 05.0 00.0 50.0 00.0 00. 0000. 0 00.0 05.0 00.0 00.0 0105mm 00.0005.H 00.0 05.0 55.0 00.0 00. 0000. 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